Brain Research, 509 (1990) 31-40 31

Elsevier

BRES 15179

The ipsilateral corticocortical connections of area 7 with the frontal

lobe in the monkey

J.W. Neal ~, R.C.A. P e a r s o n 2 a n d T.P.S. P o w e l l 1

IDepartment of Human Anatomy, Oxford (U. K.) and 2Departmentof Anatomy, St Mary's Hospital Medical School, London (U. K.)

(Accepted 1l July 1989)

Key words: Corticocortical connection; Area 7; Frontal lobe

The corticocortical connections between area 7 and the frontal lobe have been studied in the monkey. Injections of HRP were made into
area 7 of the parietal lobe or into area 46 in the walls of the principal sulcus. The two subdivisions of area 7, 7a ox? PG and 7b or PF, are
connected with different parts of the frontal lobe, and each subdivision is connected with two distinct areas. Area 7b, PF, is connected in a
well organized and somatotopic manner with the lower premotor area and with the lower part of area 46, below the fundus of the principal
sulcus. Area 7a, PG, is connected with area 8a and with the upper part of area 46, above the fundus of the principal suclus; it is suggested
that the lower part of area 8a and the posterior part of area 46 are related to the central visual field, while the medial part of area 8a and
the anterior part of area 46 are related to the periphery of the visual field. The corticocortical connections between area 7 and the frontal lobe
are reciprocal and those passing from area 7 to the frontal lobe are 'feed-forward' and those to area 7 are 'feed-back'.

INTRODUCTION MATERIALS AND METHODS

Most of the material is that which has been described in earlier

T h e c o r t e x o f a r e a 7 o f t h e p a r i e t a l l o b e in t h e m o n k e y
b r a i n is c o n n e c t e d w i t h c e r t a i n p a r t s o f t h e f r o n t a l l o b e ,
a n d t h e t w o m a j o r s u b d i v i s i o n s of a r e a 7 d i f f e r in t h e s e
connections. Area 7a o r P G , w h i c h is p r e d o m i n a n t l y
r e l a t e d t o t h e v i s u a l s y s t e m , is c o n n e c t e d t o t h e f r o n t a l
e y e f i e l d a n d t h e s u p e r i o r p a r t o f a r e a 46 (refs. 2, 3, 15,
16, 19, 29, 31), w h i l e a r e a 7 b o r PF, w h i c h is s o m a t i c
s e n s o r y in f u n c t i o n , is c o n n e c t e d t o t h e l o w e r p a r t s o f t h e
premotor c o r t e x a n d a r e a 46 (refs. 3, 13, 15, 22, 29).
T h e r e is also e v i d e n c e t h a t w i t h i n e a c h o f t h e s e sets o f
c o n n e c t i o n s t h e r e is a n o r g a n i z a t i o n 13'15'22 t h a t m a y b e
b a s e d u p o n t h e r e p r e s e n t a t i o n s o f t h e v i s u a l field 26 a n d
t h e b o d y 3° in t h e t w o s u b d i v i s i o n s of a r e a 7. C e r t a i n
f u r t h e r d e t a i l s o f t h e o r g a n i z a t i o n of t h e s e c o n n e c t i o n s
between area 7 and the frontal lobe have now been
o b t a i n e d f r o m a s t u d y o f t h e d i s t r i b u t i o n of t h e l a b e l l e d
cells in t h e f r o n t a l l o b e a f t e r i n j e c t i o n s o f h o r s e r a d i s h
p e r o x i d a s e ( H R P ) in a r e a 7, a n d t h e y c o n f i r m a n d e x t e n d
previous observations on these projections. In order to
determine whether these corticocortical connections are
r e c i p r o c a l l y a r r a n g e d in a s i m i l a r l y well o r g a n i z e d m a n -
ner, injections of HRP h a v e also b e e n p l a c e d in t h e
frontal lobe.
brief reports 25'26 of the corticocortical connections of area 7. Under
general anaesthesia and with aseptic precautions, injections of 15%
HRP or wheat germ agglutinin ( W G A ) - H R P were made into
different parts of area 7 in 14 monkeys, and into the cortex above
or below the principal sulcus in the frontal lobes of each hemisphere
in one monkey. It was felt justifiable to make the two injections into
the frontal lobes on opposite sides of the same brain because the
purpose was to study the corticocortical connections between the
frontal and parietal lobes, and not the interconnections within the
frontal lobe, and also to compare directly in the same brain the cell
labelling after injections on either side of the principal sulcus. The
injections were of varying size and were made iontophoretically by
passing current of between 2 and 5/~A for 10-20 min in pulses of
7 s with 7 s intervals; the pipette was left in the cortex for 20 min
after the injection. The animals were allowed to survive for 24-72
h, the majority for 48 h, and were then deeply anaesthetized and
perfused through the heart with White's saline and a mixture of 2%
paraformaldehyde and 1% glutaraldehyde for the HRP injections or
1.25% paraformaldehyde and 1% glutaraldehyde for the W G A -
HRP injections. The brains were removed, photographed and cut
into blocks before being immersed in 30% phosphate buffered
sucrose at 4 °C until they sank. The blocks of the frontal lobes were
cut from the rest of the hemispheres immediately posterior to the
genu of the arcuate sulcus. Frozen sections of the frontal lobes were
cut at 40 ~m in the coronal plane, and of the other blocks in either
the coronal or sagittal plane. A 1:20 series was processed with the
dianisidine and nitroprusside (DAS) method 8 and an alternate 1:20
series with the tetramethylbenzidine (TMB) method 24. The distri-
bution of the labelled cells was transferred to a planar reconstruc-
tion of the cortex on the lateral and medial surfaces of the frontal

Correspondence: T.P.S. Powell, Department of Human Anatomy, South Parks Road, Oxford OX1 3QX, U.K.

0006-8993/90/$03.50 © 1990 Elsevier Science Publishers B.V. (Biomedical Division)

32

lobe made from coronal sections. The limits of the architectonic experiment in which both areas 7a and 7b are involved.
areas described by Walker 34 have been superimposed upon the
Three separate injections were placed so that altogether
reconstruction (Fig. 1), but we have not studied the cytoarchitecture
of this region. they spread to involve most of area 7b, PF, except for the
anterior end, and a considerable part of area 7a, PG,
from the middle of the posterior wall of intraparietal
RESULTS sulcus anteriorly to the posterior margin of this area (Fig.
2). A correlation of the site of the injection in area 7b
The greater part of the extent of the cortex of the with the map of the representation of the body 3° indicates
frontal lobe that is related to area 7 is given in an that most of the representation has been involved except
that of the face. In the frontal lobe, labelled cells are
present in the anterior and posterior walls of the arcuate
sulcus and in the upper and lower walls of approximately
the posterior half of the principal sulcus (Figs. 2 and 3).
When transferred to the planar reconstruction, it can be
seen that there are cells in the posterior wall of a large
extent of the inferior limb of the arcuate sulcus, in what
is area 6 of Brodmann 6 or the lower premotor area, and
in the upper limb of the arcuate sulcus cell labelling is
seen only in the anterior wall where it is in most of its
mediolateral extent, in area 8a of Walker ~4. The labelled
cells in the walls of the principal sulcus are in area 46
(Walker34). No orthograde granularity was seen in the
cortex of the frontal lobe either in this or other
experiments.
The parts of the cortex in the frontal lobe that are
!
related to area 7b, and the somatic sensory system, have
,1 been determined in experiments with injections limited to
this area. A large injection which involves the greater
part of area 7b resulted in cell labelling along most of the
posterior wall of the inferior limb of the arcuate sulcus
and on the surface immediately below and behind the
MED
lower end of this sulcus (Fig. 4). There are also cells in
] //
Q the posterior half of the inferior wall of the principal
/ / ~ 24 CG sulcus and on the lateral surface of the hemisphere
inferior to the sulcus; this cell labelling is in area 46. A
small injection in the anterior part of area 7b, which
probably involved the adjoining representations of the
10 9 ~ face and arm 3°, resulted in cell labelling in the lower part
of the posterior wall of the arcuate sulcus and on the
• AS

Fig. 1. The extent of the cortex on the medial and lateral surfaces
of the frontal lobe of the macaque monkey (within broken lines),
above, that has been included in the planar reconstruction made
..."---2. from coronal sections, below. The limits of the architectonic areas
described by Walker 34 have been superimposed upon the planar
reconstruction by broken lines. Note that the principal sulcus is
orientated more obliquely, anteromedial to posterolaterali on the
• 12 ~ reconstruction than on the surface view because the medial margin
of the hemisphere is directed anteroposteriorly on the reconstruc-
tions rather than sloping inferiorly to the frontal pole; the genu of
POST the arcuate sulcus appears more inferiorly on the reconstruction
than on the surface view for the same reason. AS, arcuate suicus;
CG, cingulate sulcus; CL, calcarine sulcus; CS, central sulcus; IOS,
inferior occipital sulcus; IPS, intraparietal sulcus; LS, lateral sulcus;
LUNS, lunate sulcus; PS, principal sulcus; POS, parieto-occipital
sulcus; STS, superior temporal sulcus.
 33

adjoining surface, as well as in the superficial part of the

inferior wall of the principal sulcus, just behind the
middle of its anteroposterior extent, and again on the
adjoining surface (Fig. 5). Two smaller injections, in
separate experiments, which are respectively confined to
the representations of the face and the arm in area 7b
t °°
.
.AS

AS

produced two small patches of cell labelling in the

posterior wall of the arcuate suicus, and the labelling
related to the face extended on to the surface (Fig. 6); - 2
3
two small groups of labelled cells were also present in the
inferior wall of the principal sulcus and on the surface of
the cortex below the sulcus, and that due to the injection
in the representation of the face was again situated lower
on the surface than that related to the arm. The probable

Fig. 3. The distribution of the cell labelling (dots) on outlines of

AS CS IPS coronal sections at the levels shown on the outline of the hemisphere
in the preceding figure.

*o

Fig. 2. The site of the large injection of HRP in area 7 (stipple) on

an outline of the lateral surface of the hemisphere, above, and the
distribution of the cell labelling (dots) in the frontal lobe on the
planar reconstruction, below. In this and the subsequent figures
showing the distribution of cell labelling each dot, circle or triangle
represents between 20 and 30 labelled cells.
Fig. 4. The site of a large injection (stipple) in area 7b, PF, above,
and the distribution of the cell labelling (dots) in the frontal lobe on
a planar reconstruction, below.
34
r e p r e s e n t a t i o n of the trunk and leg in these frontal areas
is shown by the distribution of cell labelling after a
relatively small injection which straddles the b o r d e r
b e t w e e n areas 7a and 7b. The distribution of the cells due
to the involvement of 7a will be considered later, but
within the frontal areas which have already been shown
to be related to area 7b there are labelled cells in two
s e p a r a t e sites; in the p o s t e r i o r wall of the arcuate sulcus,
close to the bank and on the adjoining surface just above
its genu, and on the inferior bank of the posterior third
of the principal sulcus (Fig. 5). As the interpretation of
the s o m a t o t o p i c r e p r e s e n t a t i o n involved by the injections
in area 7b is based upon a correlation with the maps of
R o b i n s o n and Burton 3°, and not from direct recording, it
is possible that the interpretation of the representation of
individual parts of the b o d y in the frontal lobe is not quite
accurate, but it is in general a g r e e m e n t with o t h e r studies
CS
J J -
P$
AS
: ".o,
Fig. 5. Top: sites of two injections (stipple), in separate experi-
ments, in area 7b; the smaller involves the representations of the
face and arm and the other overlaps the boundary of areas 7a and
7b. Bottom: the distribution of the cell labelling in the frontal lobe
following these two injections; the labelling after the injection in the
representations of the face and arm is indicated by dots and that
after the injections at the boundary of areas 7a and 7b by circles.
on area 6 where the r e p r e s e n t a t i o n has been d e t e r m i n e d
electrophysiologically.
The extent of the cortex of the frontal lobe that is
related to area 7a, PG, and the visual sensory system, has
been defined in experiments with injections of variable
sizes that were confined to this area. A n injection of
m o d e r a t e size involves both the surface of the inferior
parietal lobule and the posterior bank of the intraparietal
sulcus just medial to the middle of its mediolateral extent
(Fig. 7). As after all injections in area 7a the labelled cells
in the frontal lobe are anterior to the fundus of the
arcuate sulcus; they are in the a n t e r i o r wall of the arcuate
sulcus, in the walls of the principal sulcus or on the
surface posterior to it (Fig. 7). In this e x p e r i m e n t there
is cell labelling in the medial half of the anterior wall of
the upper limb of the arcuate sulcus and within the
principal sulcus; in the latter the cells form an oblique
).-..,
/aS
AS
°,°°°
Fig. 6. The distribution of the cell labelling in the frontal lobe,
below, after two separate injections (stipple) in area 7b, above. One
injection is in the representation of the face and the resulting cell
labelling is shown in dots; the other injection is in the representation
of the arm and the triangles show the distribution of the labelled
cells after this injection.

band which is orientated from posterolateral to antero-
medial in the posterior half of its extent. The labelled
cells in the anterior wall of the arcuate sulcus are in area
8a of Walker 34, and those in the principal sulcus are in
area 46. There would appear to be some degree of
organization in these corticocortical connections of area
7a because an injection restricted to the part of this area
on the surface of the hemisphere (Fig. 8), and only
slightly more lateral than the previous one, results in the
cell labelling in the principal sulcus being deep and
around the fundus, just behind the middle of its antero-
posterior extent (Fig. 9). In the arcuate sulcus the
distribution is very similar to that after the previous
injection. After an injection that is largely confined to the
posterior wall of the intraparietal sulcus (Fig. 8), at the
same level as that on the surface in the preceding
experiment, the cell labelling is in area 8a on the anterior
wall of the arcuate sulcus just below the genu; there is
also cell labelling on the surface immediately in front of
the sulcus and in the posterior part of the lower wall of
the principal sulcus (Fig. 9). A n injection that is even
?-.
P$
AS
Fig. 7. The site of an injection (stipple) in area 7a, PG, above, and
the distribution of the cell labelling (dots) in the frontal lobe, below,
35
further lateral and straddles the boundary between areas
7a and 7b, and which has already been described with the
connections of area 7b (Fig. 5), extends deeply into the
posterior wall of the intraparietal sulcus. The cell
labelling that can be ascribed to involvement of area 7a
is on the anterior wall of the arcuate sulcus, immediately
above and below its hilum, and on the surface between
the arcuate and principal sulci. To what extent the cell
labelling in the inferior wall of the principal sulcus is due
to involvement of area 7a or 7b cannot be determined,
but is probably the result of both.
The variation in the distribution of the cell labelling in
these experiments indicates that there is an organization
ID~ C$
Fig. 8. Photomicrographs of the sites of two injections of HRP in
the cortex of area 7a, PG. Above, the site of an injection deep in
the posterior wall of the intraparietal sulcus (×3) and, below, a
small injection on the lateral surface of the hemisphere (×2). The
photomicrographs have been taken at the levels of the maximum
extent~ of the injections.
36
of these corticocortical connections in the anteroposterior
dimension at least. After injections that extend deeply
into the posterior wall of the intraparietal sulcus, the cell
labelling in the anterior wall of the arcuate sulcus is
around the genu and it continues across the surface of the
hemisphere into the walls of the most posterior part of
the principal sulcus. With involvement of the surface of
the inferior parietal lobule, the labelled cells become
more medially placed in the arcuate sulcus and more
anteriorly in the principal sulcus. When the injection is
confined to the surface of the inferior parietal lobule
labelled cells are present only in the medial part of the
arcuate sulcus and about the middle of the anteroposte-
rior extent of the principal sulcus, and the injection that
extends most posteriorly in area 7 results in the most
medially placed labelling in the arcuate sulcus and the
most anterior in the principal sulcus. Whether this
organization in the anteroposterior dimension is related
J c____
PS
AS
Fig. 9. The distribution of the cell labelling in the frontal lobe,
below, after two separate injections, at the same mediolateral level,
in area 7a. The circles indicate the labelled cells after an injection
in the cortex of area 7a on the surface of the hemisphere (stipple),
above, and the dots the cell labelling after an injection in the
posterior wall of the intraparietal sulcus.
to a representation in the visual field must remain
tentative, but a small injection in the representation of
the central part of the visual field in the visual area V4
(ref. 33) results in cell labelling in a localized patch in the
anterior bank of the arcuate sulcus, the adjoining surface
and extreme posterior end of the principal sulcus (Fig.
10). These labelled cells are in the inferior part of area
8a and in the most posterior part of area 46. It is
significant that the distribution of this cell labelling is
almost identical with that after an injection confined to
the posterior wall of the intraparietal sulcus, and also that
after this injection in area V4 there are labelled cells in
the same part of the posterior wall of the intraparietal
sulcus. With the material available it has not been
possible to discern an organization of the connections in
the mediolateral dimension, but in view of the differen-
CG
PS
AS
Fig. 10. The site of a small injection (solid black) in the
representation of the central part of the visual field in area V4,
above, and the distribution of the resulting cell labelling (dots) in
the frontal lobe, below.

tial connections of the medial and lateral parts of area 7a
with the prestriate visual areas e6 this possibility should be
examined further.
The experiments that have been described have shown
the differential projections from the cortex of the frontal
lobe to the two major subdivisions of area 7 and also
certain details of the organization within them. The two
injections in the frontal lobe indicate the reciprocity of
these connections. One injection involves the posterior
half of the inferior wall of the principal sulcus together
with the immediately adjoining surface. In the parietal
lobe (Fig. 11) there is cell labelling throughout most of
area 7b (PF), and this has also extended into two
adjoining areas in the lateral sulcus, the second somatic
(SII) and retroinsular (Ri) areas. There is also a slight
extension of the labelling into the lateral part of area 7a
(PG) where it occupies a narrow band from the fundus of
the intraparietal sulcus anteriorly to the anterior bank of
the superior temporal sulcus posteriorly. This cell label-
ling in area 7a is probably due to the injection extending
into the fundus of the principal suicus. An injection of
comparable size in the upper wall of the principal sulcus
is mainly in area 46 but also encroaches upon area 8a.
Following this injection most of the labelled cells in the
CG
Fig. 11. The distribution of the cell labelling in the parietotemporal
lobe after injections in the frontal lobe. The labelling is shown on
a planar reconstruction m a d e from sagittal sections, and the
triangles indicate the distribution following an injection in the
inferior wall of the principal sulcus and the dots that result from an
injection involving the superior wall of the principal sulcus. Ri, the
retroinsular area; SII, the second somatic sensory area.
37
parietal lobe are in area 7a where they fill most of the
exposed surface of the inferior parietal lobule, the
anterior wall of the upper part of the superior temporal
sulcus and a small focus in the posterior wall of the
intraparietal sulcus. Some labelled cells are also present
on the medial surface of the parietal lobe, and these are
in area PE of von Bonin and Bailey 5. The distribution of
the cell labelling in this experiment is in agreement with
the interpretation of an organization of the corticocortical
connections in the anteroposterior dimension from the
findings after injections in area 7a. Most of the labelled
cells are on the surface of the inferior parietal lobule
because the greater part of the injection is in area 46,
while the small focus of cells in the posterior wall of the
intraparietal sulcus is the result of the slight involvement
of area 8a. The presence of labelled cells in area PE is
also in accord with previous observations on the projec-
tions of this parietal area to the upper bank of the
p r i n c i p a l s u l c u s 15"28'29.
Following the injections in area 7, the labelled cells in
the frontal lobe were distributed throughout the depth of
the cortex in areas 6, 8a and 46, but were predominantly
in laminae III and V, with the greater number in lamina
-~22,"
D" . - " . " " -?.-
\ .\ • ",o, • ~ ~,.'%-.\',,
" ~ / ," ~ --.,'_ • ,',~,'N--,~...~"
,.x~.., ,. ~ e V , V ~ ,'
-"-': .' i',
\ " • t s • i
~ ~ / s #¢ # /s # s#/I
Fig. 12. T h e parts of the cortex in and around the arcuate and
principal sulci that would appear to be related to the somatic and
visual sensory systems from the results of the present study and
those described in the earlier literature. T h e areas of cortex related
to the somatic sensory system are shown by broken lines, and those
related to the visual system by dots and circles; the parts of the
cortex related to the central visual field are indicated by small dots
or circles, and those related to the peripheral visual field by large
dots or circles. T h e upper p r e m o t o r area is related to the somatic
sensory system by its connection with area 5 and the lower premotor
area and area 46 through their connections with area 7b, PF. Area
8a and the upper part of area 46 (dots) are related to the visual
system through their connections with areas 7a and PE, and areas
45 and the lower part of area 46 (circles) by their connections with
area T E in the temporal lobe.
38
V. The cell labelling in both areas 7a and 7b after
injection in the frontal lobe again involved mainly
laminae III and V, but here the majority were in lamina
III. In all areas the labelled cells were clearly distributed
in bands of 0.5-2.0 mm width.
DISCUSSION
The results of the experiments that have been de-
scribed confirm and extend the observations made in
earlier studies on the connections between area 7 and the
frontal lobe in the monkey. With their predominant
relationship to either the visual or the somatic sensory
system, it is not surprising that areas 7a and 7b are
connected to different parts of the frontal cortex and to
areas that are known to be important in the control of
movements of the eye and body. Perhaps one of the most
significant features of the relationship between the
subdivisions of area 7 and the frontal lobe is that each of
them, together with other areas in the parieto-occipital
lobe that are interconnected with them, is connected to
t w o areas of the frontal cortex. It is not known whether
this is due to different cells projecting to different frontal
areas or to branching of the axons from individual
neurons in area 7; nor is the functional significance of this
relationship clear.
There is a somatotopic organization in the lower
premotor area 11'13'2°'22 (which is largely in the posterior
wall of the inferior limb of the arcuate sulcus and the
adjoining part of the exposed surface near its lower end)
and with one qualification, the present results are in
agreement with the previous observations on this orga-
nization. The parts of this area and area 7b, PF, that
contain the representations of the same part of the body
are interconnected and while in both areas the represen-
tation of the arm has been shown to be relatively large,
there appears to be a discrepancy about the representa-
tion of the leg. The distribution of labelled cells after an
injection in the representation of the leg in the primary
motor area 13 and single unit recording upon movement of
the hindlimb 2° indicate that in the premotor area the
representation of the leg is in the region of the superior
precentral sulcus. Some authors consider there is no
representation of the leg in the inferior part of area 6 (ref.
11). In the present experiments both the large and the
small injections that overlapped the boundary of areas 7a
and 7b, and which should have involved the representa-
tion of the trunk and leg, did not result in cell labelling
above the level of the genu of the arcuate sulcus. It is not
possible to explain this difference at present, but it is
interesting that although Godschalk et al. ~3 found that
stimulation 'in the superior limb of the arcuate sulcus was
generally ineffective in exciting antidromic response' in
the parietal cortex, their Fig. 5 does show one example
of a response in the representation of the leg in area 7b
after stimulation just above the genu of the arcuate
sulcus. The present results are in agreement with the
electrophysiological observations on antidromic re-
sponses in the premotor area after stimulation of area 7
(ref. 13) and of labelled cells in area 7b and SII after
injections in the lower premotor area ~322.
Area 7b is also reciprocally connected with that part of
area 46 that is in the lower wall of the posterior half of
the principal sulcus and the adjoining surface below it.
From the distribution of the labelled cells after the small
injections in area 7b and a correlation of the sites of these
injections with the map of the representation of the body
in this area 3° it seems reasonable to conclude that there
is also a topographic representation in area 46. The face
is presented below and anteriorly and the leg above and
behind, with the representation of the arm again large. In
all areas of the parietal and frontal lobes the represen-
tation of the body is therefore essentially the same, with
the face below or in front and the leg above or behind,
and that of the arm is the largest.
In considering the connections of area 7a, PG, with the
frontal lobe it should be emphasized that here this area
has been treated as a single entity with the limits given by
von Bonin and Bailey 5, rather than in terms of the several
subdivisions that have recently been recognized in this
region of the parietal lobe 15'28. The finding that only area
8a, and not area 8b as well, is related to area 7a is in
accord with recent studies on the extent of the frontal eye
field7'~6; area 45 is not involved. The organization that
has been found in the connections between area 7a and
areas 8a and 46 is in agreement with earlier anatomical
and functional studies. The deep part of the posterior
wall of the intraparietal sulcus, related to the central part
of the visual field 26, is connected to the lower part of area
8a, in which the cells have small receptive fields in the
central field of vision 32 and where microstimulation
results in minimal amplitudes of saccadic eye move-
merits 7. More posterior parts of area 7a are related to the
peripheral field of vision and the cells have large
receptive fields, and these regions are connected with
more medial parts of area 8a and more anterior portions
of area 46. This relationship is also in accord with
functional observations; in the medial part of area 8a the
cells have large and more eccentric receptive fields 32 and
microstimulation gives the greatest amplitudes of sacca-
dic eye movements 7, and in area 46 the receptive fields
of the cells become progressively larger more anterior-
ly32. This interpretation of the relationship between area
7a and areas 8a and 46 in terms of the representation of
the visual field is confirmed by the findings in the
experiment with a small injection in the representation of

the central part of the visual field in area V4; the
distribution of the labelled cells in the frontal lobe was
virtually the same as after injections involving the d e e p e r
part of the p o s t e r i o r wall of the intraparietal sulcus, and
there was also cell labelling in this part of the same
sulcus. It has not been possible to d e t e r m i n e whether the
m e d i a l and lateral parts of area 7a are also differentially
c o n n e c t e d to the frontal lobe, and therefore possibly in
terms of the r e p r e s e n t a t i o n of the superior and inferior
halves of the visual field, but it is significant that
microstimulation showed that saccade direction ' h a d no
global direction across the frontal eye field 'v.
A r e a P E is also reciprocally connected with the upper
parts of areas 8a and 46, and although there is insufficient
evidence to decide w h e t h e r or not it overlaps with the
part of a r e a 46 that is related to area 7a it certainly does
in a r e a 8a (ref. 29). T h e r e is p r o b a b l y a similar
organization of its connections to that of area 7a, because
the injection in the u p p e r wall of the principal sulcus
resulted in cell labelling in the parts of the two areas that
are k n o w n to be interconnected.
The p r o j e c t i o n s f o r w a r d to the frontal cortex from
areas 7a and 7b show m a n y aspects of convergence; each
of the subdivisions of area 7 receives corticocortical
connections from several adjoining areas 25"26, but each
p r o j e c t s to only two areas in the frontal lobe. Some of the
areas of the cortex in the parietal and occipital lobes that
are c o n n e c t e d with a r e a 7a or 7b also send fibres to their
areas of p r o j e c t i o n in the frontal lobe. A s examples of
such p r o j e c t i o n s are SII and Ri which send fibres to area
7b and which also project to the lower p r e m o t o r area 13'22
and area 46, while areas PE and V4, which are connected
to a r e a 7a, send fibres to the frontal eye field and area
46.
The l a m i n a r distribution of the labelled cells was
different in the frontal and parietal cortex. In the parietal
cortex there were some cells throughout the depth of the
cortex but with the m a j o r i t y in layer III and fewer in layer
V, and this a r r a n g e m e n t would indicate that the cortico-
cortical connections to the frontal lobe are of the
f e e d - f o r w a r d type. In the cortex of the frontal lobe there
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basis for the functional relationship between the neurons
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