THE CRANIAL BASE

J. H. SCOTT
Anatomy Department, Queen’s University, Belfast, Ireland

SIX FIQURES

INTRODUCTION

The midline cranial base, extending from the anterior edge

of the foramen magnum (basion) to the fronto-nasal suture
(nasion), is one of the most important regions of the skull
from the point of view of anthropological studies, while in its
growth it affects both the cranial and facial regions of the
skull. It is proposed in this paper to collect together data
from various sources and discuss its development, its growth,
and its morphological and phylogenetic significance.

T h e parts of the cranial base

I n the human adult the cranial base consists of the following
elements from behind forwards (fig.1):
(a) The basal part of the occipital bone.
(b) The spheno-occipital synchondrosis. This persists until
about 17-20 years of age.
(c) The body of the sphenoid bone.
(d) The spheno-ethmoidal suture. This is continuous lat-
erally with the fronto-sphenoidal sutures on the floor of the
anterior cranial fossa.
(e) The cribriform plate region of the ethmoid bone uniting
the midline mesethmoid (perpendicular plate and crista galli)
with the bilateral facial parts of the ethmoid.
(f) The fronto-ethmoidal suture and the foramen caecum.
(g) The frontal bone (glabella region).
319
320 J. H. SCOTT

The structure of the cranial base in the anthropoid apes is

the same as in man, but in the monkeys, lower primates and
other animals a second synchondrosis, situated between the
presphenoid and postsphenoid elements of the body of the
sphenoid, persists until after the eruption of the second per-
manent molars. Its significance will be discussed later.

Fig. 1 Human mid-line cranial base. 0, baaioccipital; 8,sphenoid; E, ethmoid;

F, frontal; N, nasal; 1, posterior cranial segment; 2, middle cranial segment;
3, anterior cranial segment.

I n order to study the growth of the cranial base it can best

be divided into three segments. These are (fig. 1):
1. From basion to pituitary point (the posterior segment).
2. Pituitary point to foramen caecum (the middle segment).
3. Foramen caecum to nasion (the anterior segment).
I n the majority of studies on the cranial base (2) and (3)
together make up the anterior segment.
The development of the crafinial base
The cranial base from the foramen magnum to the region
of the foramen caecum is pre-formed in cartilage which is
continuous with the cartilage of the nasal capsule, the latter
including the cartilage of the nasal septum (fig. 2). The frontal
bone develops in membrane above and slightly in front of the
anterior end of this great mass of cartilaginous tissue ex-
tending from the foramen magnum to the front end of the
nasal septum. I n this mass of cartilage centers of ossification
appear from behind forwards in the following order (Frazer,
'40) :
 CRANIAL BASE 321

1. A single center for the basioccipital about the middle of

the third month of fetal life.
2. Two to four centers for the postphenoid about the end of
the 4th month of fetal life.
3. Two centers for the presphenoid during the 4 4 t h months
of fetal life.
4. A single center f o r the mesethmoid during the first year
after birth.

Fig. 2 Human mid-line cranial base in late fetal life. The ossification centere
for the basi-occipital, postsphenoid and presphenoid elements are shown in the
cartilage. V, vomer; M, maxilla; F, frontal; N, nasal.

These various parts unite with one another as follows:

1. The two parts of the sphenoid unite in man during the
last month of fetal life. They are united at birth.
2. The mesethmoid unites with the facial parts of the eth-
moid by ossification of the cribriform plate between the first
and third year.
3. The occipital unites with the sphenoid between 17 and
20 years of age.
At birth the basal part of the occipital is ossified and the
two parts of the sphenoid have just united. The mesethmoid
has not yet ossified so that the nasal septum is still entirely
cartilaginous except for the vomer at its lower edge. The
spheno-occipital synchondrosis is in the form of a bilateral
epiphyseal cartilage contributing to the growth of both the
occipital and sphenoid bones.
322 J. H. SCOTT

The growth of the cramnial base

In tables 1 and 2 the growth of the whole cranial base as
measured from basion to nasion is shown in fetal and post-
natal life. For comparison, data on the growth of body length
(crown-heel) and of the cranial circumference are also in-
cluded. In all cases the figures are the percentage of the
adult size reached at each period of growth. At the 14th week
TABLE 1
Growth of body length, cranial baae, and head oircrvmference
Fetal life
(Shown as a percentage of adult dimensions)
AQE EEAD
I N WEEXS BODY LENQTH 'ItANIAL
LENQTH amawxmmmaz
14 7 21 19
16 9 26 23
18 12 30 28
20 14 34 33
22 16 37 38
24 18 41 42
26 19 44 -
28 21 45 47
30 23 47 51
32 24 50 55
34 26 53 57
36 27 54 60
38 28 56 62
'Data for body length from Scammon and Calkins in Krogman ( '41).
*Data for cranial base from Ford ( '56).
'Data for head circumference from Pfuhl in Krogman ('41).

of fetal life both the cranial base and head circumference

have reached approximately the same proportion of their
adult size, the cranial base being slightly ahead of cranial
circumference. By the 22nd week the head circumference
begins to draw ahead in the rate of its growth, and is some-
what ahead of the cranial base at birth. During the first and
second year after birth, however, there is a great spurt in
brain growth, as illustrated by the increase in head circum-
ference from 65 to 90% of the adult size. The cranial base
does not reach 90% of its adult size until about the 13th year.
 CRANIAL BASE 323

Growth of the body as a whole is more constant than growth

of the cranial base and head circumference. While head cir-
cumference shows a great spurt in growth immediately after
birth (first to third year in association with growth of the
brain (Keith, '31), the cranial base grows most rapidly from
the 14th to the 32nd week of fetal life, with a period of less
rapid growth during the last two months, a second spurt dur-
ing the first years after birth, and a gradual slowing down
after about the 7th year.
TABLE 2
Growth of body length, crank1 base and head circumference
Birth to adult life
(Shown as a percentage of adult dimensions)
~~~

AGE BODY LINGTH OUANIAL BAS1 Horn

(B-N) OIUOUMFEEZNO1

Birth 28 55 (21)' 65
2 yrs. 50 70 (22) 90
4 yru. 59 75 (15) 93
6 yrs. 66 80 (15) 95
8 yrs. 72 86 (32) 97
10 yrs. 78 89 (12) 98
13 yrs. 85 90 (12) 99
15 yrs. 91 91 (10) 99.6
18 yrs. 98 96 ( 8 ) 99.8
I Data for body length and head circumference from Kornfeld in Krogman ('41).
'Figures in parentheses = number of skulls measured.

Table 3 shows the growth of the various segments of the

cranial base. The figures given are the actual measurements.
It will be seen that, while the growth of the posterior (basion-
pituitary) and anterior (frontal) segments show a growth
pattern similar to that of the cranial base as a whole, the mid-
dle (ethmoidal) segment reaches adult dimensions by about
the 7th year. (See also De Coster, '51; Scott, '54;Bjork, '55).
This early stabilization of the ethmoidal region is overlooked
in the use of measurements of the anterior cranial base taken
from the pituitary fossa to nasion (Brodie, '41 ; Grossman and
Zuckerman, '55) as this measurement includes the frontal
bone which continues to increase in thickness until adult life.
324 J. H. SCOTT

The available growth sites in the human cranial base after

birth are:
1. At the foramen magnum. Unlike the anthropoid apes
(Keith, '10) the foramen magnum in man does not show any
evidence of backward migration.
TABLE 3
Growth of cranial base segments

AGQ B-NI B-Pl w* *N 1

Birth to mean 54 22 29 4
two weeks (4)' range (48-58) (21-23) (25-32) (3-5)
1-3 yrs. mean 73 32 39 7
(16) range (68-79) (28-35) (3444) (@)
4-7 p. mean 80 35 44 9
(11) range (72-91) (31-38) (40-48) (7-11)
8-13 yrs. mean 90 42 46 10
(12) range (83-96) (38-45) (43-50) (9-12)
14-20 yrs. mean 97 46 47 14
(15) range (86-106) (43-49) (45-50) (11-20)
Adult M +F mean 98 47 47 16
' B = basion, P = pituitary point, N = nasion, C = foramen caecum.
a Figures in parentheses in first column = number of skulls measured.

2. At the spheno-occipital synchondrosis. This is an impor-

tant growth center until the beginning of adult life. It has
been shown (Scott, '54) that it is situated in such a position
relative to the coronal and lambdoid sutures as to influence the
growth of the cranial vault in the antero-posterior dimension
(fig.3).
3. The spheno-ethmoidal and fronto-ethmoidal sutures.
These with their lateral extensions, the fronto-sphenoidal
sutures at the floor of the anterior cranial fossa, join the
coronal suture system at pterion and are situated in the same
coronal plane as the retro-maxillary suture systems (in the
pterygo-palatine fossa). As, however, the distance from pitu-
itary fossa to foramen caecum remains constant after about
the 7th year, there is probably little gr0wt.h at these sutures
after the end of the first decade (fig. 4).
 CRANIAL BASE 325

4. The frontal bone increases in thickness a t the glabella

region by surface deposition until adult life (Keith and Cam-
pion, '21; Bjork, ' 5 5 ) , and this part of the bone is invaded by
the frontal air sinus, especially during adolescence.

Fig. 3 Diagram to show the position of the apheno-occipital synchondrosia

relative to the coronal and lambdoid euture systems and the cranial segments.
P,posterior cranial segment; M, middle cranial segment; A, anterior cranial seg-
ment; F, upper facial skeleton; 8, synchondrosis.

Fig. 4 Diagram to show the position of the spheno-ethmoidal suture relative to

the retro-maxillary sutures in the pterygo-palatine fosea region of the upper
facial ekeleton. 0,basioccipital; S, sphenoid; E, ethmoid; R,lateral pterygoid
plate; F, frontal; N, naaal; M,maxilla; P,palatine L, lacrimal. Arrow indicates
position of spheno-ethmoidal suture.
326 J. H. SCOTT

The cranial base and the cranium

Table 4 shows the degree of correlation between the size of
the cranial base and cranial capacity (brain size). The data
are from the great series of 19th century Scottish skulls
measured by Young ( '17). Three hunrded and seventy-seven
male skulls have been divided into four groups according to
their cranial base (basion to nasion) length.
(a) Cranial base length 90-95 mm (37 skulls).
(b) Cranial base length 96-100 mm (167 skulls).
(c) Cranial base length 101-105 mm (141.skulls).
+
(d) Cranial base length 106 mm (32 skulls).
The average length of the cranial base for the series is 100.37
mm.
It will be seen that the larger skulls tend to have the larger
cranial base but the range of variation for cranial size is
quite great for each group. That is, the cranial base is not
directly correlated with growth of the brain. The relative
independence of the size of the cranial base and cranial capac-
ity is further illustrated in table 5 from data supplied by
Hrdlicka ('39) on micro- and macro-crania. It Will be seen
that in the female microcrania the range for cranial base
length, 82-95 mm, does not differ much from the normal range
of 8 6 1 0 0 mm, while the average value of 85.1 mm is not greatly
below the normal of 93 mm as compared with the differen&&
the cranial capacity. The comparison between the large crania
and the normal shows even less difference in regard to the
cranial base measurement.
Tables 6 and 7 show the degree of correlation between
cranial base length, cranial capacity and the cephalic index.
It will be seen that for skulls of a given cranial base length
there is a slight tendency for the larger crania to be brachy-
cephalic. Howells ('41) found a correlation coefficient of
-0.54 between cranial base length and cranial index in the
Gallen crania. Taking 0.5 f as signscant (Smart, '38) this
is just significant. Again, there is a very wide range of
variation in these correlations as was shown by Huxley in
 TABLE 4
Correlation of cranial base length and cranial capacity. Scottiah aeries. Y a l e skulls
(Data from Young, '17)

OEANIAL B u m CEANIAL CAPACITY, CaL'

LENaTH
-1200 1210-1300 1310-1400 1410-1500 1510-1600 1610-1700 1700 f
d

mm
90-95 (37)' 3 27 24 24 19 3 0
W
96-100 (167) 4 8 34 32 17 5 1 P
ul
M
101-109 (141) 0 3 21 37 24 11 3
106 + (32) 0 0 5 19 41 17 19

'Distribution shown as a percentage of total in each group.

'Figures in parentheses = number of skulls.
338 J. H. SCOTT

TABLE 6
Cranial base length in microcrania and macrocrania
(Data from Hrdlicka, '39)

Rang0 of cranial capacity 940-1050 1750-2100

Range of cranial base length 82-95 94-122
Average of cranial base 85.1 104.3
Normal skulls
(Data from MacCurdy, '23 and Young, '17)
60 P E B U V U N 100 BCOTTIBE
(mar-) (nm)
Range of cranial capacity 1020-1410 1250-1930
Range of cranial base length 84-100 92-112
Average of cranial base 93 100.96

TABLE 6

Correlation of cranial base, crania2 capacity and cephalic index

Male eknrlk
(Based on data from Young, '17)
Cranial base = 100 m m (46 skulls)
CEANIAII OXPHm,O CrnmAL CEPEALIO
CAPACITY ' RTDEX CAPACITY INDEX

1200 76.8 1460 (2) 74.7'77.5

1290 73.2 1470 (2) 74.1,74.5
1300 73.4 1480 (2) 76.4,75.8
1310 (2) 72.9,74.7 1490 74.3
1320 (2) 74.5,69.4 1500 (2) 72.3,74.5
1330 71.6 1510 77.0
1340 (2) 73.8,72.9 1520 (2) 72.9,73.1
1350 (2) 73.9,74.5 1530 70.6
1370 (2) 70.7,73.1 1540 76.5
1380 (2) 75.5'75.0 1550 (2) 76.3,71.5
1390 (2) 72.9,73.3 1570 77.6
1410 69.8 1630 (2) 74.0,77.6
1420 (3) 69.8,71.3, 1710 76.0
77.2
1430 77.6
1440 72.2
1450 (2) 73.1'74.1
Range of cranial capacity 1200-1710.
Range of cephalie index 69.4-77.6.
 CRANIAL BABE 329

1867 when he compared in detail two skulls, one the widest

(C.I. =97.9) and the other the narrowest ((3.1.=62.9) he
had met. The cranial base length was the same in both. I n
commenting on this he wrote : “Brachycephaly and dolicho-
cephaly are not necessarily connected with the shortening and
.
lengthening of the base of the skull . . their most extreme
forms may arise exclusively from modifications of the side
walls and roof of the cranium.”
TABLE 7
Correlutwn of cranial base, craniul capacity and oephalia index
Cranial base = 100 mm (48 skulls)
CEPHALIC INDBX
W I A L OAPAOITY
72 and - 74 and - 75 and + 77 and +
1440- (26) 6 16 5 2
1450 + (22) 2 7 9 4

Among different living human races the average cranial base

length for male skulls extends from 95 mm in Bushmen to
108 mrn in the Eskimo. In females the measurements are
always slightly less ; the average being 93 mm in Bush skulls
and 103 mm in the female Eskimo (Martin, ’28). As there
is some correlation between brain size and body size, it might
be expected that a certain degree of correlation would exist
between the length of the long bones and the cranial base.
Howells ( ’41),however, found the very low value of - 0.015
for the correlation coefficient between length of the femur
and the basion-nasion length, and stated that stature has no
apparent relation to either the size or the shape of the head for
Early Christian Irish.

The cranial base and the facial skeleton

The facial skeleton (excluding the mandible) is closely re-
lated to the anterior cranial segment of the skull (fig. 4) which
consists of the frontal, mesethmoid and sphenoid bones, and
this upper part of the face is closely related in its develop-
330 J. H. SCOTT

ment and growth with the anterior half of the cranial base
(from pituitary fossa to nasion) that is, the middle and an-
terior segments. I n early fetal life the cartilage of the cranial
base is continuous with that of the nasal capsule (fig. 2). The
ossilication centers which appear in the cranial base have
already been described. I n the cartilage of the nasal capsule
the ossification centers of the facial ethmoid and the inferior
turbinate bones appear during fetal life and are fully ossiflecl
by the time of birth. Centers for the maxillae, zygomatic,
palatine and lacrimal bones appear in condensations of meso-
derm in close relation to the nasal capsule early in fetal life,
while the vomer appears as an ossification center in the peri-
chondrium bordering the lower edge of the cartilage of the
nasal septum. W'ith further development these facial bones
come into relationship with one another at various sutures
which together make up two great facial suture systems. One
of these, the circummaxillary system, separates the maxilla
from the frontal, lacrimal, facial ethmoid, palatine, zygomatic
and vomer, while the craniofacial system separates the lac-
rimal, facial ethmoid, palatine, zygomatic and vomer from the
bones of the anterior cranial segment: the frontal, sphenoid
(greater wing of median pterygoid plate and body) and me-
sethmoid (perpendicular plate of ethmoid). These suture
systems are arranged so as to permit growth of the upper
facial skeleton to take place in a downward and forward
direction (fig. 4) and it has been suggested (Scott, '53, '54)
that this growth is regulated to a considerable extent during
fetal life and early childhood by the growth of the cartilage
of the nasal septum.
I n man the sutures between the back of the maxilla and the
vertical plate of the palatine and between the palatine and
the median pterygoid plate and the suture line across the floor
of the anterior cranial fossa (spheno-ethmoidal and fronto-
ethmoidal) make up a suture system which would permit the
bones in front of the system (frontal ethmoidal, palatine and
maxillae) to grow forward from the sphenoid. I n so doing
space would be made for the maxilla to grow backwards and
 CRANIAL BASE 331

thus make room for the erupting molar teeth. This is the clas-
sical theory of maxillary growth (Keith, ’02 ; Brodie, ’42).
We have already seen, however, that growth of the cranial
base from pituitary fossa to foramen caecum, that is, across
this suture system, ceases about the 7th year and, while it is
possible that the maxilla map continue to move forward rela-
tive to the other facial bones after this period, there is no
evidence that this takes place to any considerable extent. It
would appear that growth of the upper face can be divided into
two phases: (a) an early phase during fetal life and early
childhood when growth of the anterior half of the cranial
base and nasal capsule (especially the nasal septum) thrusts
the facial bones downwards and forwards and allows growth
to take place at the facial and anterior cranial sutures, and
(b) a later phase, from about 7 years until adult life, when
growth at the facial sutures has ceased and growth of the
upper part of the face is produced entirely by surface deposi-
tion associated with internal absorption of bone to allow for
increase in size of the nasal cavities, air sinuses and the oral
cavity.
Although growth of the upper facial skeleton is closely
related with the growth of the anterior half (middle and
frontal segments) of the cranial base, it is possible for a
normal facial skeleton to develop in relation to a reduced
cranial base. This is shown by a study of the microcrania and
macrocrania described by Hrdlicka ( ’39). He writes : “ I n no
one of the specimens, small or large, is there any significant
abnormality of the palate, the dental arches or the teeth.”
This relative independence of the growth of the cranial and
facial parts of the skull is illustrated in table 8, in which cer-
tain measurements of 10 of the microcrania from Hrdlicka are
compared with the range of variation of 50 normal Peruvian
skulls (from MacCurdy, ’23). F o r cranial length, 8 of the
microcranial skulls are below the normal range; for cranial
width, none; for cranial base length, two; for orbital height,
palate width and bisygomatic width, one; and for upper facial
height and nasal height, none.
 TABLE a

Cranial and facial measurements in microcrania

(Data from Hrdlicka, '39 and MacCurdy, '23)
IZANQEor
NOILYAG
1 a 3 4 6 6 7 8 9 10 PERUVIAN
BKULLB

Cranial capacity 920 940 955 970 970 970 980 990 995 1000 1020-1410
9
Cranial length 144 156 152 153 148 147 159 157 157 159 158-180
w
Cranial width 130 130 130 130 143 134 124 132 130 126 119-139
%
Cranial base length 88 88 92 82 88 88 82 89 91 94 847100

Upper face height 61 64 64 57 61 64 56 63 60 65 5S71

Orbital height 31.5 33 30.5 33 33 34 32.5 30.5 32.5 32 31-41

Nasal height 44 46 45.5 41.5 43 49.5 42.0 42.5 43.5 45.5 40-52

Bizygomatic width 121 116 120 113 123 122 117 121 127 124 115-130

Palate width 56 63 63 52 57 57 - 66 60 59 53-66

 CRANIAL BASE 333

The relationship between growth of the cranial base and the

facial skeleton is further illustrated by conditions such as
achondroplasia in which there is a failure in the growth of the
cartilage of the cranial basc. I n two newborn children de-
scribed by Huntcr ('33), the cranial base lcngth was 23 m
and 25 mm. As, however, the cartilage of the nasal capsule is
also involved, the reduction of the upper facial skeleton
found in this condition (Brash, '56) is probably more directly
related to this site of cartilage growth failure than to the
failure of growth at the spheno-occipital synchondrosis. The
ability of the upper facial skeleton to compensate for failure
of growth of the cranial base and to a lcsser extent for failure
of growth of the cartilage of the nasal capsule, is probably
dependent on surf ace deposition, which is independent of
suture growth. Even in skulls which show a marked reduc-
tion in the size of the upper part of the face, in the region
of the bridge of the nose, there is often a normal development
of the alveolar process and of the lower part of the face. This
is well illustrated in the case of a microcephalic skull from
the Anatomy Museum of the Queen's University (fig. 5). The
cranial capacity is 455 mm, cranial length 140 mm, cranial
width 103 mm, cranial base length 94 mm, upper facial height
68 mm, orbital height 32 mm, nasal height 51 mm, bizygomatic
width 106 mm, and the external palatal width 63 mm. Of these
measurements, cranial capacity, cranial length, cranial width,
and bizygomatic width are well below the normal range, while
the other measurements are within the range of normal, al-
though the cranial base is short f o r a male skull. The alveolar
arch, however, is very large arid perfectly formed.

The flexure of the cranial base

I n man the cranial basc shows a higher dcgrec of flexure of
the cranial base than in any other animal. Duckworth ('04)
states that the gradual decrease in the sizc of the spheno-
ethmoidal angle from the lo\wr mammals to man gives a good
indication of the gradual increase in the dcvelopmcnt of the
334 J. H. SCOTT

frontal lobes of the brain. Like Huxley (1867) and Topinard

(1890) he measures this angle between the two lines, one from
basion to spheno-ethmoidal suture (prosphenion) and the
other from the spheno-ethmoidal suture to nasion. Ford ('56)
shows that during fetal life the angle increases from 131.5'

Fig. 5 Microceplialie skull with a well devclopcd dental arch.

at 10 weeks to 150.5" at birth, a total increase of 19". This,

of course, involves a flattening or straightening out of the
cranial base during fetal life. What happens after birth is
less certain. Topinard, quoting Welcker, gives average values
of 140" for 6 newborn infants, 137" for 10 children from 10-15
years of age, 138" for 30 German women and 134" for 30
German men. The number of skulls in each group is, however,
 CRANIAL BASE 335

quite inadequate to allow for the wide range of individual

variation. Zuckcrman ( ' 5 5 ) , using prosphenion as the place
of inflexion gives a value of 151.9" as the mean for 8 skulls
under one year ; 148" for 12 skulls between one and two years ;
144.1" for 9 skulls between 3-5 years; 144.5" for 28 skulls
between 6-8 years; 142.6' for 4 skulls between 9-14 years;
145.1" for 20 skulls between 15-21 years and 148.8"for 99 adult
skulls. The slight differences in the angle at various ages
are probably the expression of the range of variation. Bjork
( ' 5 5 ) , using the middle of the pituitary fossa as the meeting
point of the anterior and posterior cranial axial lines, gives
the mean valuc of the angle as 130.8" at 12 years of age, and
131.6" at 20 years of age. He states that in some indviduals
the angle increases (maximum 5"), while in others it decreases
(5.5"). He used serial x-rays of the skull in his studies. Young
('17) in 98 adult male skulls from his Scottish series, using
phosphenion as the site of inflexion, found the range of varia-
tion to extend from 137" to 170".
It should be pointed out that nasion is not altogether a sat-
isfactory point for measuring the angle of flexure. It varies
in its relationship to the foramen caecum (the true anterior end
of the cranial base), and it also changes its position with age
(Scott, '56). The position of nasion relative to the anterior
end of the true cranial base also shows wide variation in dif-
ferent animals.
A more important question, however, is the exact site at
which the bending process occurs. It is usually considered that
this takes place at the meeting of the two bounding lines, that
is at the spheno-ethmoidal suturc (prosphenion). This is un-
likely as it would involve a thrusting backwards of the maxilla
and palatine bones against the median ptcrpgoid plates in
the facial skeleton. Bjork ('55) considers that the rotation
takes place at the spheno-occipital synchondrosis. There is,
however, another site which persists in man up to birth and in
monkeys and lower mammals for a much longer period, that is,
at the spchondrosis between the presphenoid and post-
sphenoid elements of the body of the sphenoid bone.
336 J. H. SCOTT

Cameron ('24) believed that the flexion of the cranial base

occurs in the region of the pituitary fossa and it is inter-
esting to note that the synchondrosis between the two parts of
the sphenoid as seen in the Rhesus monkey (fig. 6) is related to
the front of the fossa. It is probable that this is the main
site of cranial base flexure during fetal life. Changes after
birth in man, if in fact they do occur, and if they are not due

Fig. 6 Cranial base in Rhesus monkey about three years of age. The spheno-
occipital and sphenoidal synchondrosis arc both shown. The latter is closely
related to the front of the pituitary fossa above and to the septa1 cartilage below.

to changes in the position of nasion, are probably due, as

Bjork suggests, to changes at the sphenoccipital synchon-
drosis which can act as a secondary site of flexure. It would
appear that the failure of the angle to increase in man after
birth is due not to growth of the frontal lobes of the brain
but to the early union of the two parts of the body of the
sphenoid bone which unite the anterior and posterior parts of
the cranial base across the main site of flexure. The straight-
ening out of the cranial base in animals other than man and
the anthropoid apes so that it reaches 180" or more in adult
 CRANIAL BASE 337

animals (Duckworth, '04) appears to be related to the direc-

tion of growth of the cartilage of the nasal septum. I n a
fetal dog the direction of growth of the cartilage is vertical,
while in an adult animal it is horizontal (Bolk, '26; de Beer,
'51). I n man it remains more or less vertical. As, however,
the nasal cartilage is itself an extension of the cartilage of the
cranial base, a change in its growth direction will involve the
anterior half of the cranial base. I n the Rhesus monkey, in
which the synchondrosis between the two parts of the sphenoid
persists until after the eruption of the second permanent
molars, Ashtoii ('57) describes a steaay opening out of the
angle of the cranial base up to this period. One hundred and
forty-nine degrees is the average for 14 female animals be-
tween birth and alignment of the first permanent molars;
155" is the average for 9 animals between eruption of the
permanent incisors and eruption of the second molars, and
164" is the average before eruption of the canines and third
molars. After this time his figures show a slight reduction
but this may be due to individual variation among the small
numbers of animals used. The average given for 12 adult
animals is 160".
Comparative anatomy of the cralzial base
Table 9 gives some information from various sources for
cranial base length and the spheno-cthmoidal angle in the dog,
Rhesus monkey, baboon, chimpanzee, orang and gorilla. Much
of the apparent difference between the values for the angle
shown among the primates appears to be due to the position of
nasion relative to the foramen caecum. I n the adult baboon and
Rhesus monkey, nasion, as in man, is at about the same level
as the floor of the anterior cranial fossa, while in the gorilla
and orang it is well above this level. I n table 9 the values of
the spheno-ethmoidal angle provided by Cameron ( '30) and
Ashton ('57) are both given. Cameron used nasion as his
anterior point; Ashton in an attempt to avoid the faults of
this position used a point where the mid-sagittal plane is
crossed by a line joining the upper limits of the frontal pro&
 w
w
00

Comparative measuremelcta of cranial boss

(Data mainly from Ashton, '57 and Cameron, '30)

Do0 BHlSUS BABOON OBANB 6H)EILLA OHIYPANmH YAN

?
Cranial base length 95 120 108 100.38
ul
Basion-pituitarypoint (Ashton) 27 44 47 58 44 48 d
0
H
Pituitary point-nasion (Ashton) 37 46 55 73 56 59 H
Pituitary point-foramencaeeum 32 35 41 48
Spheno-ethmoidalangle (Ashton) 161" 148' 167" 169" 159" 133"
Spheno-ethmoidalangle (Cameron) 208" 173" - 161" 178' 159" 131'
 CRANIAL BASE 339

esses of the maxillae. He states that this lies approximately

along the floor of the anterior cranial fossa, but even this point
lies well above the level of the foramen caecum in these ani-
mals, (anterior end of the cribriform plate). This is because
in many animals, including the anthropoid apes, but not in
man, the cribriform plate and foramen caecum lie at the
bottom of a deep olfactory pocket (Cameron, '30). Even in
these cases, however, the foramen caecum probably repre-
sents more accurately than any other point the true anterior
end of the cranial base.

TABLE 10

The spheno-ethmoidal angle in 6 chimpanaee skulls, wring foramen caecum

a8 the anterior point

NO. DENTAL CONDITION ANGLl

1 Deciduous dentition complete 155"

2 Deciduous dentition complete 140"
3 Deciduous dentition complete 160"
4 1st permanent molar erupting 145"
5 1st permanent molar in occlusion but deciduous
molars still in place 145"
6 Adult 160"

I n the higher anthropoid apes, as in man, the synchondrosis

between the two parts of the sphenoid closes early, probably
at or about the time of birth. However, Ashton ('57) shows
an increase in the angle up to adult life in the anthropoid
apes as in Rhesus monkeys. I n table 10 I have given tho
measurements of the angle in 6 chimpanzee skulls using fora-
men caecum as the anterior point. Although the number of
skulls is quite inadequate, the table shows the need for measur-
ing a much larger number of skulls in order to eliminate the
individual variation factor and indicates that more work still
requires to be done in regard to the comparative anatomy of
the cranial base.
 TABLE 11

Cranial base, cranial capaaity and cephdic indez in human fossils*

UPPER XODERN
PITAEOANTB&OPVS SINANTHROPUS H.SOWENSIS NEANDERTHAL P~O LITHIa
MAN

Cranial base length:

Range 108-113 - 113-1 14 98-125
Average 110.5 105.5 113.5 111 104.8 (M) 102.7
99.0 (F) F-l
Cranial capacity: F
Range 775-900 915-1225 rn
1035-1255 1220-1610 c,
Average 822 1043 1100 1400 1300
Cranial index:
Range 73.2-79.3 71.4-72.6 66.2-76.7 68.2- 76.3
Average 76.3 72.2 72.0 73.3 72.8
- - .
-- .

Cranial base angle Early

101-1 17
Late
123-1 35
From Weidenrcich ( '43), Howell ( '51) and Morant ( '30).
 CRANIAL BASE 341

Cranial base in fossil man

Table 11gives the cranial base length, cranial capacity and
cranial index in some human fossils (from Morant, '30;
Weidenreich, '43, '45), and the cranial base angle in Neander-
thal skulls (Howell, 5 1 ) . Cranial base length was greater in
early man and has tended to decrease with increase in cranial
capacity. Although the number of skulls, on which the evi-
dence of the change in the cranial base flexure in Neanderthal
man is based, is limited the difference is significant and inter-
esting.
DISCUSSION

Perhaps the most interesting problem regarding the morph-

ology of the cranial base is concerned with the high degree
of flexure which is characteristic of the human skull. The de-
crease in the angle of flexure is associated with an increase in
the height, and a reduction in the length, of the cranial vault
(Weidenreich, '41, '47 ; Bjork, '55), but from what we know
of the development and growth of the cranial base, there is
no evidence that these changes are the direct result of brain
growth. The brain appears to adapt its form to that of the
skull base upon which it rests, and both brain growth and
cranial base growth contribute to the form of the cranial vault.
If the cranial base is grossly underdeveloped, as in achon-
droplasia, or if one of its growth centers is destroyed growth
of the brain tends to provide the necessary room by a rounding
(brachycephalization) of the vault. Weidenreich ( '45) has
pointed out the correlation between the expansion of the brain
(relative to body size) and reduction of the jaws among cer-
tain breeds of dog. He states that the changes are due to the
expansion of the brain and that the brain of a dwarf breed
of dog, such as the King Charles spaniel, is much too large
for the reduced skull and consequently needs for its accomoda-
tion all the space in the skiill which can be made available.
The result is an enormous expansion of the brain case in all
its dimensions and enormous reduction of the jams and teeth.
He concludes that the principle of the correlation shown
342 J. 11. SCOTT

among different races of dog is exactly the same as that which

is manifested in the phylogenetic transformation of the hu-
man skull. As, however, the cause of the variations shown
on the skulls of dwarf breeds of dog are probably related
to achondroplasia and the reduction of the facial skeleton is
a consequence of this rather than an increase in brain size,
phylogenetic comparisons are perhaps somewhat inapt unless
one postulates that modern man is an achondroplastic mutant
as Keith (’11)suggested that Xeanderthal man showed some
of the characteristic traits of acromegaly.
Elsewhere, however, Weidenreich ( ’43, ’47) suggested that
the bending of the cranial base is a final step in the adaptation
of the skull to new static and dynamic conditions necessitated
by the acquisition of the upright posture. That there is some
correlation between cranial base flexure and prognathism of
the facial skeleton is indicated by the phylogenetic changes
in the skull form shown by human evolution from Pithecan-
thropus to modern man, and the very interesting “reversion”
on the part of late Neanderthal fossils in which it appears that
the face became more prognathous and the cranial base angle
more opened out. This correlation although suggestive is not,
however, an inevitable process of evolution. The baboon, which
develops a very massive and prognathous facial skeleton, does
so in relation to a cranial base angle of 148” (Ashton, ’55)
which is considerably less than that of the Rhesus monkey
and is in fact closer to the human average of 133” than any
other higher primate.
Table 12, constructed from the data supplied by Young ( ’17),
shows the correlation between the spheno-ethmoidal angle
and the spheno-maxillary angle. The latter is a measurement
of facial prognathism. It will be seen that there is a general
but not exact correlation between cranial flexure and facial
prognathism.
From a consideration of the available evidence it would
seem that the cranial base angle is, however, in a general way
related to the growth of the upper facial skeleton. It has been
pointed out that the anterior part of the chondrocranium is
 CRANIAL BASE 343

continuous with the cartilage of the nasal capsule and that

growth of the nasal septum involves the cranial base from
the sphenoid to the frontal bone. I n some animals (de Beer,
'37) the mesethmoid is absent and the presphenoid ossifica-
tion centers invade the septal cartilage replacing the per-
pendicular plate of human anatomy. This spread of ossifica-
tion from the sphenoid region of the cranial base into the
nasal septum emphasizes the structural continuity of these
parts. In animals other than man and the higher anthropoids,
the synchondrosis between the postsphenoid and presphenoid
persists for a considerable time after birth and even in man it
is an active site of growth during fetal life when the cranial
TABLE 12
Correlation of cranial base flexure and facial prognathiem
(Data from Young, '17)

BPHCNO- 8 P H E N O ~ Y M I L W K YA N Q U
ETHYOIDAL
ANGLB 07-17" 18-830 84-89' 900 +
136-141" 4 1 0 0
142-147" 3 9 8 0
148-153" 1 15 13 1
154-159' 1 5 19 6
160" + 0 2 2 10

base flexure is established. This synchondrosis is continuous

with the septal cartilage in most animals until after the union
of the perpendicular plate with the romer, and in many ani-
mals the septal cartilage continues to extend back to the
sphenoid in a bony canal or tube even after union of the septal
bony elements (Scott, '53). Bolk ('26) has suggested that the
essential nature of the cranial base flexure is due to a failure
of the angle to open out during fetal life. In early fetal life
all mammals show a flexure which, however, usually disap-
pears or becomes reversed (180"+) by the time of birth
except in the primates and especially in man. I t would seem
therefore that if the face does not change its fetal position
relative to the cranium, the cranial base remains flexed.
311 J. H. SCOTT

Even if we can establish a correlation between facial growth

and the cranial base, it still remains for us to face the funda-
mental problem of why these changes began in the lower pri-
mates (Tarsius) and reached their full development in man.
We are back to Weidenreich's theory of a correlation between
skull form and the upright posture. I n animals which move
on 4 limbs and which carry the skull at the front end
of a horizontal vertebral column, the facial skeleton must
move outwards and forwards from its developmental position
below the skull to a position in front of the cranium. If this
does not happen the most satisfactory compensating mecha-
nism would be to sit up (as monkeys and apes, and certain
rodents, do) and then stand up (as men do). Both these
actions liberate the forelimbs to become organs of manipula-
tion and discriminative sensation, and this in relation with
the establishment of connections within the brain with the
centers for the sense organs of sight, smell, taste, etc., may
have been the biogenetic stimulus necessary to set the brain
on its dramatic human evolutionary expansion. The anatomy
of the Australopithecinae would seem to indicate that brain
development followed the acquisition of the upright posture
rather than determined it.
I n an interesting paper Du Brul ( '50) points out that lago-
morphs, rodents and marsupials, animals with general hori-
zontal body orientation, have skulls in which facial, cranial
and body axes are in the same general horizontal line as com-
pared with animals assuming relatively erect posture and loco-
motion habitudes, in which changes in orientation of these
axes take place in relation to the change in body posture.
Only in the primates, however, has there occurred any great
change in the size of the brain, and only among the primate6
does the cranial base angle fall much below 180".
This would seem to indicate that the growth of the primate
brain, although correlated with the upright posture, was not
a necessary consequence of the change in position of the body
axis. It is probably more directly correlated with the use of
the forelimbs which have developed a much greater repertory
 CRANIAL BASE 345

of skills among the primates than in any of the rodents or

marsupials. As Wood Jones ( ’16) puts it: “In the primates,
owing to the preponderant use of the forelimb, there is no
need for a mouth which reaches out for food, or for a mouth
which seizes food or kills it when seized, all these functions
being discharged by the mobile and grasping forelimb.”
We would, however, express it rather differently in suggesting
that the forelimb came into use to compensate for failure of
development of the facial skeleton. Reduction of the face is
often associated with reduction of the dentition, but the eden-
tates and other animals show loss of teeth without any reduc-
tion of the facial skeleton.
Elsewhere (Cole, ’54;Scott, ’54) it has been suggested that
a function of the nasal cavities in many animals is that of
regulating heat loss from the body. This is associated with
t.he complexity of the inferior turbinate processes, and espe-
cially the maxillary turbinates, which require a well developed
upper facial skeleton to house them. I n the primates there is
a progressive loss of turbinate complexity and this is especially
marked in man. It is possible that with the development of
the cutaneous heat-regulating mechanism which reaches its
fullest complexity in man, the nasal mechanism has fallen
progressively into disuse, and that this, rather than loss of
the teeth, is the main factor responsible for the reduction of
the facial skeleton among the higher primates.
It is interesting to notice that in the chimpanzee the facial
skeleton, which supports the alveolar processes and the rela-
tively large teeth, is about the same size as in man and the
reduction of the upper face in relation to the nasal cavities
is much the same as in man. This is partly hidden by the
development of the facial buttress system, including the supra-
orbital bars which are, however, related to the masticatory
apparatus. The alveolar prognathism of the living anthropoids
appears to be a secondary specialization and to have been less
extensive in fossils such as Proconsul. Among the anthropoid
apes alveolar prognathism developed in relation to an enlarge-
ment of a certain type of dentition (large incisors and canines
346 J. H. SCOTT

but moderate-sized cheek teeth). Among the Australopithe-

cinae it appears to have developed, probably independently, in
relation to a different kind of dentition (small incisors and ca-
nines but massive cheek teeth). Early man had a moderate
degree of alveolar prognathism which modern man is losing.
These dental changes, however, do not in themselves account
for the bending of the cranial base. They seem to be but
morphogenetic pulsations superimposed upon a deeper and
more continuous process underlying the progressive reduction
of facial skeleton among the primates, and the face of the ba-
boon and the gorilla are but secondary specializations super-
imposed upon this more basic theme. The secret beginnings of
the human brain may have begun in the structure and phys-
iology of its humble cousin the skin, and in nakedness the
master of creation may have been born.
SUMMARY

1. The cranial base in its growth and morphology has a

limited effect on the growth and form of the cranial vault and
has but little relationship to growth of the brain.
2. The prepituitary half of the cranial base is closely related
to the development and growth of the upper facial skeleton,
and development of the facial skeleton is closely related to the
flexure of the base.
3. I n man most of the bending process takes place during
fetal life and it probably occurs chiefly at the synchondrosis
between the postsphenoid and presphenoid elements. In ani-
mals other than man and the anthropoid apes, this synchon-
drosis persists until after birth permitting a continuation of
the bending processes.
4. As nasion bears a highly variable relationship to foramen
caecum, the true anterior end of the cranial base, many meas-
urements of the cranial base angle are unreliable for compara-
tive purposes.
5. It is suggested that failure of the human face to alter
its fetal developmental relationship to the cranium initiated
the establishment of the fully upright posture and that this,
 CBANIAL BASE 347

in association with sensory changes in the forelimbs and body

skin, was responsible for the later phenomenonal development
of the human brain.
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348 J. H. SCOTT

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