Paper - The ossification of the human frontal bone with special reference to its presumed pre- and post-frontal elements: Difference between revisions

From Embryology
(Created page with "{{Header}} {{Ref-Inman1937}} {| class="wikitable mw-collapsible mw-collapsed" ! Online Editor   |- | 90px|left <br> Musculoskeletal - Skull Deve...")
 
mNo edit summary
Line 10: Line 10:
{{Historic Disclaimer}}
{{Historic Disclaimer}}
The Ossification of the Human Frontal Bone with Special Reference to its Presumed Pre- and Post-Frontal Elements
The Ossification of the Human Frontal Bone with Special Reference to its Presumed Pre- and Post-Frontal Elements
J Anatomy, 71(3):


By Verne T. Inman, M.D., Ph.D. And John B. De C. M. Saunders, M.B., F.R.C.S. (Edin.)
By Verne T. Inman, M.D., Ph.D. And John B. De C. M. Saunders, M.B., F.R.C.S. (Edin.)
Line 20: Line 16:
Division of Anatomy, University of California Medical School
Division of Anatomy, University of California Medical School


==Introduction==


INTRODUCTION
In a recent number of the Journal of Anatomy there has appeared an article by Ashley-Montagu on the presumed existence of a post-frontal element in the skull of a young gorilla. Ashley-Montagu’s conclusions are based largely on the view that the post-frontal bone "is present in the developing skull of probably all primates and certainly of all men. “ It is not Widely known among anatomists”, he says, “that the post-frontal bone is a constant element of the developing human skull. For some extraordinary reason most modern textbooks of anatomy and embryology make no mention of the post-frontal element of the human foetal skull in spite of the fact that a constant and independent centre of ossification of this bone is present in every case, and the bone itself may generally be observed up to the end of the third foetal month, and not rarely in the skulls of newborn children.” .
 
IN a recent number of the Journal of Anatomy there has appeared an article
by Ashley-Montagu on the presumed existence of a post-frontal element in
the skull of a young gorilla. Ashley-Montagu’s conclusions are based largely
on the view that the post-frontal bone "is present in the developing skull of
probably all primates and certainly of all men. “ It is not Widely known among
anatomists”, he says, “that the post-frontal bone is a constant element of
the developing human skull. For some extraordinary reason most modern
textbooks of anatomy and embryology make no mention of the post-frontal
element of the human foetal skull in spite of the fact that a constant and
independent centre of ossification of this bone is present in every case, and the
bone itself may generally be observed up to the end of the third foetal month,
and not rarely in the skulls of newborn children.” .
 
Having available a considerable collection of material specifically prepared
to demonstrate the ossification of the skull in man, we were prompted to seek
for this element. We were surprised, in the light of the above definite statement,
that we were unable to find the slightest trace of this element occurring as an
independent centre, nor were we at any time able to find such other secondary
centres of ossification as have been commonly described for the frontal bone.
We noted, furthermore, that there was some disagreement among the authors
of various standard textbooks, and it became increasingly apparent that in
view of the great morphological and phylogenetic conclusions which have
been based upon the supposed existence of such secondary centres, reinvestigation of the ossification of the frontal bone was necessary.


LITERATURE


Gabriel Fallopio appears to have been one of the first to state clearly that
Having available a considerable collection of material specifically prepared to demonstrate the ossification of the skull in man, we were prompted to seek for this element. We were surprised, in the light of the above definite statement, that we were unable to find the slightest trace of this element occurring as an independent centre, nor were we at any time able to find such other secondary centres of ossification as have been commonly described for the frontal bone. We noted, furthermore, that there was some disagreement among the authors of various standard textbooks, and it became increasingly apparent that in view of the great morphological and phylogenetic conclusions which have been based upon the supposed existence of such secondary centres, reinvestigation of the ossification of the frontal bone was necessary.
the frontal bone constantly develops from two symmetrical halves. In his
Observatimws Anatomicae (1561) he says: “Ita ut in omnibus puerulis qui
tenerimi sunt frontis os ex duabus constare partibus reperierim.” Vesalius
was also well aware of this mode of origin of the frontal bone, for not only does


Anatomy Lxxt 25
==Literature==
384 Verne T. Immm and John B. de 0. M. Saunders


he discuss Fallopio’s observations in his Observationes Gabrielis Falloppii
Gabriel Fallopio appears to have been one of the first to state clearly that the frontal bone constantly develops from two symmetrical halves. In his Observatimws Anatomicae (1561) he says: “Ita ut in omnibus puerulis qui tenerimi sunt frontis os ex duabus constare partibus reperierim.” Vesalius was also well aware of this mode of origin of the frontal bone, for not only does he discuss Fallopio’s observations in his Observationes Gabrielis Falloppii Ezvamen (1564), but the matter is mentioned in the Fabrica (1543). Jhering states that Galen was ignorant of this fact, but it would seem on examining his De usu partium and De admim'stratiom'bus anatomicis that he may have been aware of this in some instances.
Ezvamen (1564), but the matter is mentioned in the Fabrica (1543). Jhering
states that Galen was ignorant of this fact, but it would seem on examining
his De usu partium and De admim'stratiom'bus anatomicis that he may have
been aware of this in some instances.


Up to the time of Kerckring (1670) but few serious attempts had been made
Up to the time of Kerckring (1670) but few serious attempts had been made to examine the origin of the bones before birth. This author dissected and described specimens from approximately the second month of foetal life.
to examine the origin of the bones before birth. This author dissected and
described specimens from approximately the second month of foetal life.


Albinus illustrates in his Icones ossium foetus humani (1737) that area
Albinus illustrates in his Icones ossium foetus humani (1737) that area which has since been called the post-frontal element and records an instance in which each half of the frontal bone was divided into three parts. This single observation was made on an aborted foetus. He says: “In abortu juniore, cui totum os magnitudinis unguis mediocris, utramque illam partem inveni divisam in tres; superiorem, quae ad frontem pertinet; inferiorem, quae ad foramen oculi, tertiam quae ad caput supercilii.”
which has since been called the post-frontal element and records an instance
in which each half of the frontal bone was divided into three parts. This single
observation was made on an aborted foetus. He says: “In abortu juniore,
cui totum os magnitudinis unguis mediocris, utramque illam partem inveni
divisam in tres; superiorem, quae ad frontem pertinet; inferiorem, quae ad
foramen oculi, tertiam quae ad caput supercilii.”


Apart from the single observation quoted above from Albinus, the recognition of secondary or accessory centres of ossification dates from the work of
Apart from the single observation quoted above from Albinus, the recognition of secondary or accessory centres of ossification dates from the work of Serres (l829).1 This author described accessory centres, (1) for the part of the pars orbitalis at the trochlear fossa and (2) for the zygomatic process. Serres considered them, in association with his studies on the comparative anatomy of the skull, as representing the pre- and post-frontal elements. Rambaud & Renault (1864) accept the findings of Serres, but do not appear to have seen these centres as independent ossicles. They state that at the fourth month the centres have already been joined by a suture which leaves only a shallow groove and that the centres have always so appeared at this time. These authors added a third centre for the nasal spine which, they state, appears late, sometime after the eighth year.
Serres (l829).1 This author described accessory centres, (1) for the part of the
pars orbitalis at the trochlear fossa and (2) for the zygomatic process. Serres
considered them, in association with his studies on the comparative anatomy
of the skull, as representing the pre- and post-frontal elements. Rambaud
& Renault (1864) accept the findings of Serres, but do not appear to have seen
these centres as independent ossicles. They state that at the fourth month the
centres have already been joined by a suture which leaves only a shallow
groove and that the centres have always so appeared at this time. These
authors added a third centre for the nasal spine which, they state, appears
late, sometime after the eighth year.


A fourth centre, for the posterior part of the pars orbitalis, is given by
A fourth centre, for the posterior part of the pars orbitalis, is given by Von Spec (1896) in von Bardeleben’s H andbmth. The existence of such a centre appears to be based on an observation originally made by Hyrtl (1860) that a Wormian bone is sometimes found in this region. Von Spee also observed such a Wormian bone and accepts Hyrtl’s conclusion that this has arisen from an independent centre of ossification. It has frequently been pointed out since (Patten, 1912) that the presence of a Wormian bone cannot be accepted as evidence of a true secondary centre.
Von Spec (1896) in von Bardeleben’s H andbmth. The existence of such a centre
appears to be based on an observation originally made by Hyrtl (1860) that
a Wormian bone is sometimes found in this region. Von Spee also observed
such a Wormian bone and accepts Hyrtl’s conclusion that this has arisen from
an independent centre of ossification. It has frequently been pointed out since
(Patten, 1912) that the presence of a Wormian bone cannot be accepted as
evidence of a true secondary centre.


Since the publication of Rambaud & Renault’s monograph, a number of
Since the publication of Rambaud & Renault’s monograph, a number of special papers have appeared on the ossification of the frontal bone and on its pre- and post-frontal elements. Lecourtois (1869) published a study based upon foetal skulls of 5-6 months, in which he expresses agreement with the findings of Serres and of Rambaud & Renault.
special papers have appeared on the ossification of the frontal bone and on its
pre- and post-frontal elements. Lecourtois (1869) published a study based
upon foetal skulls of 5-6 months, in which he expresses agreement with the
findings of Serres and of Rambaud & Renault.


1 Many authors give as references to the work of Serres on accessory centres the C'.R. Acad. S’ci.,
1 Many authors give as references to the work of Serres on accessory centres the C'.R. Acad. S’ci., Paris, 1819 and 1838. These abstracts contain no mention of specific centres. Rambaud & Renault also give as reference Serres, Des Lois de l’oatéoge’m'e, Paris, 1829. This work is unavailable to us. It would seem that the majority of authors have quoted Serres through Rambaud & Renault.
Paris, 1819 and 1838. These abstracts contain no mention of specific centres. Rambaud & Renault
also give as reference Serres, Des Lois de l’oatéoge’m'e, Paris, 1829. This work is unavailable to us.
It would seem that the majority of authors have quoted Serres through Rambaud & Renault.
The Ossification of the Human Frontal Bone 385


In 1872, von Jhering published an important paper which has since been
In 1872, von Jhering published an important paper which has since been widely quoted. J hering, after reviewing the work of earlier authors, confirms the conclusions arrived at by Rambaud & Renault. He examined, however, no specimen earlier than the 4th month and at no time saw the assumed accessory centres as independent ossicles. Maggi (1897, 1898) published two extensive articles on the morphology of the cranial and cranio-facial bones in which he concludes that the existence of accessory centres for the frontal bone has been satisfactorily demonstrated, and that two of these centres are homologous with the pre- and post-frontal elements of other forms. His observations on man were based on the examination of foetal skulls from the 5th month and on the work of Rambaud & Renault. finally Von Bardeleben (1896), Augier (1912) and Ashley-Montagu (1931) have Written articles which, on analysis, present no original observations on the development of the frontal bone, but whose conclusions are either directly or indirectly based on the acceptance of the work of Serres and of Rambaud & Renault.
widely quoted. J hering, after reviewing the work of earlier authors, confirms
the conclusions arrived at by Rambaud & Renault. He examined, however,
no specimen earlier than the 4th month and at no time saw the assumed
accessory centres as independent ossicles. Maggi (1897, 1898) published two
extensive articles on the morphology of the cranial and cranio-facial bones in
which he concludes that the existence of accessory centres for the frontal bone
has been satisfactorily demonstrated, and that two of these centres are homologous with the pre- and post-frontal elements of other forms. His observations
on man were based on the examination of foetal skulls from the 5th month and
on the work of Rambaud & Renault. finally Von Bardeleben (1896), Augier
(1912) and Ashley-Montagu (1931) have Written articles which, on analysis,
present no original observations on the development of the frontal bone, but
whose conclusions are either directly or indirectly based on the acceptance of
the work of Serres and of Rambaud & Renault.


It is not surprising in consequence, that the standard textbooks of anatomy
It is not surprising in consequence, that the standard textbooks of anatomy and of embryology almost uniformly describe and illustrate, apart from the primary centres, three pairs of accessory centres: (1) for the frontal spine, (2) variously for the medial angular process or trochlear pit (pre-frontal element), and (3) for the region of the zygomatic process (post-frontal element). These accessory centres, apart from that for the frontal spine, are, in the general concensus of opinion, assumed to appear about the 4th month and to fuse at the 7th. The centre for the nasal spine is recognized as appearing late, at or before the 10th year. In addition Quain and Von Spee (von Bardeleben’s H andbuch) mention the occasional occurrence of a fourth pair for the posteromedial part of the orbital plate. The statements in Quain, Cunningham, Gray, Morris, Piersol, Gegenbaur, Merkel, von Bardeleben, Testut, and Poirier, are, as in the special articles noted above, a continuation of the opinion of Serres
and of embryology almost uniformly describe and illustrate, apart from the
primary centres, three pairs of accessory centres: (1) for the frontal spine,
(2) variously for the medial angular process or trochlear pit (pre-frontal
element), and (3) for the region of the zygomatic process (post-frontal element).
These accessory centres, apart from that for the frontal spine, are, in the
general concensus of opinion, assumed to appear about the 4th month and to
fuse at the 7th. The centre for the nasal spine is recognized as appearing late,
at or before the 10th year. In addition Quain and Von Spee (von Bardeleben’s
H andbuch) mention the occasional occurrence of a fourth pair for the posteromedial part of the orbital plate. The statements in Quain, Cunningham, Gray,
Morris, Piersol, Gegenbaur, Merkel, von Bardeleben, Testut, and Poirier, are,
as in the special articles noted above, a continuation of the opinion of Serres


and of Rambaud & Renault.
and of Rambaud & Renault.


The only dissenting opinions which we have been able to find in the
The only dissenting opinions which we have been able to find in the literature are those of Toldt and Le Double. Toldt, while recognizing the areas described by Serres and by Rambaud & Renault, is unable to accept these as indicating the existence of accessory centres. In Maschka’s Hcmdbuch d. gerichtlichen Medizin he says: “Ich habe etwas Derartiges beim Menschen nie gesehen und muss daher die Angaben von Serres, Rambaud u. Renault, welche durch J hering auch in die deutsche Literatur iibergegangen sind, wonach beim Menschen ein urspriinglich selbstandiges Os frontale posterius bestehe, in Abrede stellen. Ebenso wenig kann ich fiir den Menschen die Existenz eines zu irgend einer Zeit isolierten Os frontale anterius einraumen.”
literature are those of Toldt and Le Double. Toldt, while recognizing the areas
described by Serres and by Rambaud & Renault, is unable to accept these as
indicating the existence of accessory centres. In Maschka’s Hcmdbuch d.
gerichtlichen Medizin he says: “Ich habe etwas Derartiges beim Menschen
nie gesehen und muss daher die Angaben von Serres, Rambaud u. Renault,
welche durch J hering auch in die deutsche Literatur iibergegangen sind, wonach
beim Menschen ein urspriinglich selbstandiges Os frontale posterius bestehe,
in Abrede stellen. Ebenso wenig kann ich fiir den Menschen die Existenz
eines zu irgend einer Zeit isolierten Os frontale anterius einraumen.”


Le Double discusses extensively the ontogeny and phylogeny of the preand post-frontal elements and points out the lack of agreement among anatomists. The existence of accessory centres in man has arisen, in his opinion,
Le Double discusses extensively the ontogeny and phylogeny of the preand post-frontal elements and points out the lack of agreement among anatomists. The existence of accessory centres in man has arisen, in his opinion, from misinterpretation of the findings.
from misinterpretation of the findings.


This survey would indicate that the present belief that accessory centres for the frontal bone exist in man depends in large measure on the observations
This survey would indicate that the present belief that accessory centres for the frontal bone exist in man depends in large measure on the observations and interpretation of Serres and of Rambaud & Renault. In spite of the denial of their existence by Toldt, whose opinion is unfortunately buried in a textbook of forensic medicine, and the questioning of Le Double, all modern textbooks of anatomy continue to illustrate and describe such centres. Furthermore their existence forms the chief basis for their acceptance as pre— and post—frontal elements.
and interpretation of Serres and of Rambaud & Renault. In spite of the
denial of their existence by Toldt, whose opinion is unfortunately buried in
a textbook of forensic medicine, and the questioning of Le Double, all modern
textbooks of anatomy continue to illustrate and describe such centres. Furthermore their existence forms the chief basis for their acceptance as pre— and
post—frontal elements.


MATERIAL AND METHODS
==Material and Methods==


Our observations are based on a total of ninety-eight specimens ranging
Our observations are based on a total of ninety-eight specimens ranging from the 6th week of intra-uterine life to the 10th month, post-natal. All the material was Caucasian. The majority of the specimens were prepared to demonstrate ossification by a technique developed in this department. This method is much more satisfactory than that of Schultze in that it enables one to follow the progress of ossification to term. The method, published elsewhere by R. Skarda (1934), briefly consists in the rapid dissection of fresh foetal material which has previously been thoroughly freed from blood by washing via the umbilical vessels. The specimens are then hardened in 95 per cent alcohol for 48 hours and cleared in 1 per cent solution of NaOH and finally permanently mounted in 2 per cent formalin solution. The clearing process is carried out in darkness which improves the development of the final preparation. This technique results in preparations of great beauty. The bone is pure white, and its finest ramifications can be traced with case when viewed in good light under the dissecting microscope. The cartilage remains opalescent blue in colour. Shrinkage is approximately 2 per cent and distortion negligible. The specimens are mounted intact. This is a great advantage in that relations are maintained and there is little danger of creating artifacts. Furthermore the fact that the specimens are wet avoids cracking of the delicate bone which has given rise to so many of the erroneous observations of earlier workers on ossification.
from the 6th week of intra-uterine life to the 10th month, post-natal. All the
material was Caucasian. The majority of the specimens were prepared to
demonstrate ossification by a technique developed in this department. This
method is much more satisfactory than that of Schultze in that it enables one
to follow the progress of ossification to term. The method, published elsewhere
by R. Skarda (1934), briefly consists in the rapid dissection of fresh foetal
material which has previously been thoroughly freed from blood by washing
via the umbilical vessels. The specimens are then hardened in 95 per cent
alcohol for 48 hours and cleared in 1 per cent solution of NaOH and finally
permanently mounted in 2 per cent formalin solution. The clearing process
is carried out in darkness which improves the development of the final preparation. This technique results in preparations of great beauty. The bone is
pure white, and its finest ramifications can be traced with case when viewed
in good light under the dissecting microscope. The cartilage remains opalescent
blue in colour. Shrinkage is approximately 2 per cent and distortion negligible.
The specimens are mounted intact. This is a great advantage in that relations
are maintained and there is little danger of creating artifacts. Furthermore
the fact that the specimens are wet avoids cracking of the delicate bone which
has given rise to so many of the erroneous observations of earlier workers on
ossification.


The above technique demands absolutely fresh material. Foetuses of
The above technique demands absolutely fresh material. Foetuses of 6 months or over, if not fresh, tend to be opaque, and the ramifications of the delicate trabeculae cannot be made out with the same clarity. This can be remedied by carrying the preparation a stage further. The less satisfactory material is carefully dehydrated by carrying it successively from 50 per cent to absolute alcohol in steps of 5 degrees and then transferring to pure benzol. Specimens prepared in this second fashion show distinctly more shrinkage, and although the most delicate bone centres can be seen and readily followed, they do not have the same elegance as the others, nor do they have the same permanency. '
6 months or over, if not fresh, tend to be opaque, and the ramifications of the
delicate trabeculae cannot be made out with the same clarity. This can be
remedied by carrying the preparation a stage further. The less satisfactory
material is carefully dehydrated by carrying it successively from 50 per cent
to absolute alcohol in steps of 5 degrees and then transferring to pure benzol.
Specimens prepared in this second fashion show distinctly more shrinkage,
and although the most delicate bone centres can be seen and readily followed,
they do not have the same elegance as the others, nor do they have the same
permanency. '


In addition, dissections were made of formalinized specimens and a few
In addition, dissections were made of formalinized specimens and a few dried foetal skulls examined.
dried foetal skulls examined.
The Ossification of the Human Frontal Bone 387


OSSIfiCATION OF THE FRONTAL BONE
==Ossification of the Frontal Bone==


The primary centres
The primary centres


It is universally agreed that the frontal bone ossifies from two primary
It is universally agreed that the frontal bone ossifies from two primary centres which appear in membrane between the fortieth and fiftieth day. There is some disagreement however with regard to the precise position in which these centres arise. The majority of investigators and most standard textbooks describe ossification as beginning in the position of the future frontal eminences; the minority place the centres at the superciliary region.
centres which appear in membrane between the fortieth and fiftieth day.
 
There is some disagreement however with regard to the precise position in
We /have observed the frontal bone appearing as two finely reticulated areas of ossification in specimens of 32 and 36 mm. sitting height (s.H.). In these and in all other specimens of approximately the same age ossification involves an area whose long axis corresponds approximately to the position of the superciliary arch, and which forms the lower part of the frontal squame and the more anterior portion of the orbital plate and a. primitive medial angular process. In our younger specimens this ossific centre is about 2 by 3 mm. The ossific meshwork is of regular and even density throughout this area. except for a slight thickening and condensation which forms a line in the position of the primitive supraorbital margin. In slightly older specimens a few fine, less dense trabeculae project radially from the periphery of the centre. These trabeculae are best defined along the upper margin of the developing pars frontalis.
which these centres arise. The majority of investigators and most standard
textbooks describe ossification as beginning in the position of the future frontal
eminences; the minority place the centres at the superciliary region.


We /have observed the frontal bone appearing as two finely reticulated
After the appearance of the primary centres, ossification spreads more rapidly in the pars frontalis than in the pars orbitalis. Ossification proceeds in the pars frontalis by the formation of heavy islands of bone which constitute a crescentic area whose maximum depth corresponds to the future posterosuperior angle (see Pl. I, fig. 1). These islands fuse to form a series of primary radiating trabeculae (Pl. I, fig. 2). The appearance of secondary trabeculae serves to link the primary trabeculae, expanding the even meshwork of the primary centre. This expansion of the meshwork of the primary centre behind the advancing zone of ossification gives rise to the appearance of a centre in the position of the frontal eminence from which the primary trabeculae radiate, an effect which is often seen in specimens of 4 months and older. In fact, however, this false centre of radiation rises higher and higher up on the frontal squame with increasing age.
areas of ossification in specimens of 32 and 36 mm. sitting height (s.H.).
In these and in all other specimens of approximately the same age ossification
involves an area whose long axis corresponds approximately to the position
of the superciliary arch, and which forms the lower part of the frontal squame
and the more anterior portion of the orbital plate and a. primitive medial
angular process. In our younger specimens this ossific centre is about 2 by
3 mm. The ossific meshwork is of regular and even density throughout this
area. except for a slight thickening and condensation which forms a line in the
position of the primitive supraorbital margin. In slightly older specimens
a few fine, less dense trabeculae project radially from the periphery of the
centre. These trabeculae are best defined along the upper margin of the
developing pars frontalis.


After the appearance of the primary centres, ossification spreads more
In the pars orbitalis the progress of ossification is less rapid. The radiating trabeculae so characteristic of the pars frontalis are very much smaller and less obvious and an expanding meshwork of trabeculae is maintained.
rapidly in the pars frontalis than in the pars orbitalis. Ossification proceeds
in the pars frontalis by the formation of heavy islands of bone which constitute
a crescentic area whose maximum depth corresponds to the future posterosuperior angle (see Pl. I, fig. 1). These islands fuse to form a series of primary
radiating trabeculae (Pl. I, fig. 2). The appearance of secondary trabeculae
serves to link the primary trabeculae, expanding the even meshwork of the
primary centre. This expansion of the meshwork of the primary centre behind
the advancing zone of ossification gives rise to the appearance of a centre in
the position of the frontal eminence from which the primary trabeculae radiate,
an effect which is often seen in specimens of 4 months and older. In fact,
however, this false centre of radiation rises higher and higher up on the frontal
squame with increasing age.


In the pars orbitalis the progress of ossification is less rapid. The radiating
These observations show that ossification begins in the region of the superciliary arch and simultaneously gives rise to a portion of both the pars frontalis and the pars orbitalis and to the primitive medial angular process. The common statement that ossification commences at the site of the future frontal eminence is erroneous, and is undoubtedly the outcome of observations made on older specimens.  
trabeculae so characteristic of the pars frontalis are very much smaller and
less obvious and an expanding meshwork of trabeculae is maintained.


These observations show that ossification begins in the region of the
superciliary arch and simultaneously gives rise to a portion of both the pars
frontalis and the pars orbitalis and to the primitive medial angular process.
The common statement that ossification commences at the site of the future
frontal eminence is erroneous, and is undoubtedly the outcome of observations
made on older specimens.
388 Verne T. Inman and John B. de 0. M. Saunders


The secondary centres
The secondary centres


In the ninety-eight specimens examined we have at no time been able to
In the ninety-eight specimens examined we have at no time been able to find any true accessory centres of ossification. No centres appear in those regions in which they have been stated to occur. In all instances the bony trabeculae were found to be continuous and unbroken. Furthermore, the regions in which such accessory centres are presumed to arise do not correspond to those in which secondary island formation is found. The specimens on which we base our observations are of such clarity that there is little possibility that such centres, had they existed, could have been overlooked. None the less, if dried preparations of foetal skulls between the third and seventh months, and in some at term, are examined, there are appearances which suggest the possible existence of secondary centres. It is these appearances which have, no doubt, been accepted by many authors as indicating the existence of accessory centres. We shall therefore consider the changes which occur in those areas which have given rise to error in interpretation.
find any true accessory centres of ossification. No centres appear in those
regions in which they have been stated to occur. In all instances the bony
trabeculae were found to be continuous and unbroken. Furthermore, the
regions in which such accessory centres are presumed to arise do not correspond
to those in which secondary island formation is found. The specimens on which
we base our observations are of such clarity that there is little possibility
that such centres, had they existed, could have been overlooked. None the
less, if dried preparations of foetal skulls between the third and seventh months,
and in some at term, are examined, there are appearances which suggest the
possible existence of secondary centres. It is these appearances which have,
no doubt, been accepted by many authors as indicating the existence of accessory
centres. We shall therefore consider the changes which occur in those areas
which have given rise to error in interpretation.


(1) The presumed centre for the zygomatic process or post-frontal element.
(1) The presumed centre for the zygomatic process or post-frontal element.


The primary centre of ossification gives rise to a part of the frontal and
The primary centre of ossification gives rise to a part of the frontal and orbital plates and a primitive medial angular process. The bony superior orbital margin seen at this period does not correspond to that of the adult. The lateral two-thirds from the position of the future supra-orbital notch to the zygomatic process is due to an entirely new formation. At 42 mm. s.H. a linear thickening of osseous material appears in the lateral two-thirds of the orbital margin at the junction between the pars orbitalis and pars frontalis. This thickening becomes more and more apparent as it is followed laterally, and has resulted in the formation at birth of almost the entire supero-lateral wall of the orbit as well as giving rise to the zygomatic process, and thereby completes the separation of the orbital cavity from the temporal fossa. At 49 mm. S.H. it has spread to form a recognizable zygomatic process which is more apparent at 59 mm. s.H. (see Pl. I, fig. 2). A considerable interval exists between this process and the frontal process of the zygomatic bone; an interval which persists until the foetus has attained a length of 250 mm. The closure of this interval. is due to a further extension of ossification in the frontal process of the zygomatic bone rather than to any further ossification in the zygomatic process of the frontal, for at 155 mm. s.H. the proportions of the zygomatic process of the frontal have almost attained those seen at birth.
orbital plates and a primitive medial angular process. The bony superior orbital
margin seen at this period does not correspond to that of the adult. The lateral
two-thirds from the position of the future supra-orbital notch to the zygomatic
process is due to an entirely new formation. At 42 mm. s.H. a linear thickening
of osseous material appears in the lateral two-thirds of the orbital margin at
the junction between the pars orbitalis and pars frontalis. This thickening
becomes more and more apparent as it is followed laterally, and has resulted
in the formation at birth of almost the entire supero-lateral wall of the orbit
as well as giving rise to the zygomatic process, and thereby completes the
separation of the orbital cavity from the temporal fossa. At 49 mm. S.H. it
has spread to form a recognizable zygomatic process which is more apparent
at 59 mm. s.H. (see Pl. I, fig. 2). A considerable interval exists between this
process and the frontal process of the zygomatic bone; an interval which persists
until the foetus has attained a length of 250 mm. The closure of this interval.
is due to a further extension of ossification in the frontal process of the zygomatic
bone rather than to any further ossification in the zygomatic process of the
frontal, for at 155 mm. s.H. the proportions of the zygomatic process of the
frontal have almost attained those seen at birth.
 
From 49 mm. s.H. onwards the base of the zygomatic process of the frontal,
including the lateral two-thirds of the orbital margin, progressively thickens.
This thickening extends dorsally on to the lateral surface of the pars frontalis
but does not involve its postero-lateral angle. This angle is, as a result of this
thickening, bounded antero-superiorly by a ridge. The ridge and the area
bounded by it becomes, with further growth of the skull, more and more
apparent until, in any specimen from the third month on, a pseudo-suture
The Ossification of the Human Frontal Bone 389
 
appears to separate off this angle, giving rise to the appearance of an autonomous
centre. In a macerated skull at 7 months, the ridge extends upwards and
dorsally from the posterior border of the zygomatic process (lateral angular
process) to a point some 5 mm. above the pterion (Text-fig. 2, l.t.). This ridge
delimits a quadrilateral area of bone, approximately 5mm. in height and
15 mm. in length, which includes the postero-lateral angle (Text-fig. 2, p. f.).
Dissections and histological sections demonstrate that this ridge gives attachment ‘to the temporal muscle and is none other than the linea temporalis.
This is confirmed by the finding, in a few specimens, of a continuation of the
ridge on to the antero-inferior angle of the parietal bone.
 
In the disarticulated frontal bone, the new process which springs from
the margin between the pars frontalis and orbitalis, and which completes the
 
 
 
'7


~ 1
>1


Text-fig. l. Text-fig. 2.
From 49 mm. s.H. onwards the base of the zygomatic process of the frontal, including the lateral two-thirds of the orbital margin, progressively thickens. This thickening extends dorsally on to the lateral surface of the pars frontalis but does not involve its postero-lateral angle. This angle is, as a result of this thickening, bounded antero-superiorly by a ridge. The ridge and the area bounded by it becomes, with further growth of the skull, more and more apparent until, in any specimen from the third month on, a pseudo-suture appears to separate off this angle, giving rise to the appearance of an autonomous centre. In a macerated skull at 7 months, the ridge extends upwards and dorsally from the posterior border of the zygomatic process (lateral angular process) to a point some 5 mm. above the pterion (Text-fig. 2, l.t.). This ridge delimits a quadrilateral area of bone, approximately 5mm. in height and 15 mm. in length, which includes the postero-lateral angle (Text-fig. 2, p. f.). Dissections and histological sections demonstrate that this ridge gives attachment ‘to the temporal muscle and is none other than the linea temporalis. This is confirmed by the finding, in a few specimens, of a continuation of the ridge on to the antero-inferior angle of the parietal bone.


Text-fig. 1. Postero-inferior view of the left disarticulated frontal bone of 7 months’ old human
foetus. P.F. pars frontalis; P.0. pars orbitalis; l.t. linea temporalis; p. “presumed”
post-frontal element; f. superior limb of fissure separating supero-lateral process from the
pars orbitalis.


Text-fig. 2. Lateral View of the facial region of a full-term human foetal skull. p. “presumed”
Text-fig. 1. Postero-inferior view of the left disarticulated frontal bone of 7 months’ old human foetus. P.F. pars frontalis; P.0. pars orbitalis; l.t. linea temporalis; p. “presumed” post-frontal element; f. superior limb of fissure separating supero-lateral process from the pars orbitalis.


post-frontal element; l.t. linea temporalis; pr. “presumed” pre-frontal element; 7'. ridge
Text-fig. 2. Lateral View of the facial region of a full-term human foetal skull. p. “presumed” post-frontal element; l.t. linea temporalis; pr. “presumed” pre-frontal element; 7'. ridge separating the medial angular process from the pars frontalis.
separating the medial angular process from the pars frontalis.


supero-lateral wall of the orbit and gives rise to the zygomatic process, is
In the disarticulated frontal bone, the new process which springs from the margin between the pars frontalis and orbitalis, and which completes the supero-lateral wall of the orbit and gives rise to the zygomatic process, is separated postero-laterally, from the rest of the pars orbitalis, by a deep fissure (Text-fig. 1, f.). This fissure gives attachment to the membrane filling the lateral fontanelle. The inferior limb of the fissure extends on to the roof of the orbital cavity, in the floor of the lachrymal fossa. The inferior limb of the fissure is obliterated, medio-laterally, at birth, but usually persists as a linear series of small foramina and is often evident in the adult. The presence of this fissure in the later months of intra-uterine life further falsely accentuates the appearance of an independent centre of ossification in this area.
separated postero-laterally, from the rest of the pars orbitalis, by a deep
fissure (Text-fig. 1, f.). This fissure gives attachment to the membrane filling
the lateral fontanelle. The inferior limb of the fissure extends on to the roof of
the orbital cavity, in the floor of the lachrymal fossa. The inferior limb of the
fissure is obliterated, medio-laterally, at birth, but usually persists as a linear
series of small foramina and is often evident in the adult. The presence of this
fissure in the later months of intra-uterine life further falsely accentuates the
appearance of an independent centre of ossification in this area.


The area outlined by the linea temporalis and the fissure described above
The area outlined by the linea temporalis and the fissure described above constitutes what has been designated as the post-frontal element of the human 390 Verne T. Inman and frontal bone, and has been regarded as arising from an autonomous secondary centre of ossification.
constitutes what has been designated as the post-frontal element of the human
390 Verne T. Inman and John B. de 0. M. Saunders
 
frontal bone, and has been regarded as arising from an autonomous secondary
centre of ossification.


(2) The presumed centre for the medial angular process (trochlear pit) or prefrontal element.
(2) The presumed centre for the medial angular process (trochlear pit) or prefrontal element.


A primitive medial angular process is ossified simultaneously with the
A primitive medial angular process is ossified simultaneously with the appearance of the primary ossific centre. This process appears to be a continuation of the superciliary margin medially and forms a small portion of the anterior part of the medial wall of the orbit. At 49 mm. S.H. only the more anterior portion of the pars orbitalis is osseous. At this time ossification is relatively more extensive in the medial angular process.
appearance of the primary ossific centre. This process appears to be a continuation of the superciliary margin medially and forms a small portion of the
anterior part of the medial wall of the orbit. At 49 mm. S.H. only the more
anterior portion of the pars orbitalis is osseous. At this time ossification is
relatively more extensive in the medial angular process.


Ossification spreads much more slowly in the orbital roof and in consequence an unossified interval persists, until the 4th month, between that part
Ossification spreads much more slowly in the orbital roof and in consequence an unossified interval persists, until the 4th month, between that part of the medial angular process which lies in the medial wall of the orbit and the roof.
of the medial angular process which lies in the medial wall of the orbit and
the roof.


At 59 mm. S.H. the medial angular process commences to show some
At 59 mm. S.H. the medial angular process commences to show some thickening of its reticular network, Whereas at this time scarcely a third of the orbital plate is ossified.
thickening of its reticular network, Whereas at this time scarcely a third of
the orbital plate is ossified.


At 80 mm. S.H. ossification has spread to complete the major part of the
At 80 mm. S.H. ossification has spread to complete the major part of the orbital plate and it commences to fuse with the medial angular process, obliterating the interval between these structures. This line of fusion is indicated at birth either by a groove, or by a series of foramina resembling a suture, which rums obliquely across the trochlear pit to the anterior end of the fronto-ethmoidal suture (Text-fig. 2).
orbital plate and it commences to fuse with the medial angular process,
obliterating the interval between these structures. This line of fusion is indicated at birth either by a groove, or by a series of foramina resembling a suture,
which rums obliquely across the trochlear pit to the anterior end of the fronto—
ethmoidal suture (Text-fig. 2).


At 59 mm. S.H. a slight thickening extends medially from the position of the
At 59 mm. S.H. a slight thickening extends medially from the position of the future supra-orbital notch across the base of the medial angular process to the metopic suture (Pl. I, fig. 2). This thickening indicates for the first time the boundary of the permanent orbital rim. At 99 mm. S.I-I. the trabeculae have thickened to such an extent as to produce a ridge which separates the medial angular process from the pars frontalis, a ridge which is very apparent at term (Text-fig. 2, r.). As a result, the superior orbital margin of the foetus is Well defined medially. A new margin is therefore formed anterior to the primitive margin which, if traced downwards, becomes continuous with the crista lacrimalis anterior. Between these two margins lies the fossa sacci lacrimalis and the lacrimal sac.
future supra-orbital notch across the base of the medial angular process to the
metopic suture (Pl. I, fig. 2). This thickening indicates for the first time the
boundary of the permanent orbital rim. At 99 mm. S.I-I. the trabeculae have
thickened to such an extent as to produce a ridge which separates the
medial angular process from the pars frontalis, a ridge which is very apparent
at term (Text-fig. 2, r.). As a result, the superior orbital margin of the foetus
is Well defined medially. A new margin is therefore formed anterior to the
primitive margin which, if traced downwards, becomes continuous with the
crista lacrimalis anterior. Between these two margins lies the fossa sacci
lacrimalis and the lacrimal sac.


In the later months of foetal life a quadrilateral area of bone is defined in
In the later months of foetal life a quadrilateral area of bone is defined in the trochlear region of the macerated frontal bone. This area is bounded anteriorly by the ridge of bone which crosses the base of the medial angular process, superiorly by a groove or series of foramina, inferiorly by the frontomaxillary, fronto-lacrimal and anterior part of the fronto-ethmoidal sutures (Text-fig. 2, pr.f.). Its surface is rough and irregular. It is this area which has been almost universally regarded as the accessory centre for the trochlear
the trochlear region of the macerated frontal bone. This area is bounded
anteriorly by the ridge of bone which crosses the base of the medial angular
process, superiorly by a groove or series of foramina, inferiorly by the frontomaxillary, fronto-lacrimal and anterior part of the fronto-ethmoidal sutures
(Text-fig. 2, pr.f.). Its surface is rough and irregular. It is this area which
has been almost universally regarded as the accessory centre for the trochlear


pit or medial angular process and which has been designated as the pre-frontal
pit or medial angular process and which has been designated as the pre-frontal element of man.
element of man.




The centre for the nasal  
The centre for the nasal


At no time prior to birth have we ever observed the presence of a secondary
At no time prior to birth have we ever observed the presence of a secondary centre for the nasal spine. This process remains unossified for some time after birth. Whether the spine ossifies from secondary centres at or about the 12th year, as is usually stated, or whether it ossifies as an extension from the frontal bone, we are not in a position to say.
centre for the nasal spine. This process remains unossified for some time after
birth. Whether the spine ossifies from secondary centres at or about the 12th
year, as is usually stated, or whether it ossifies as an extension from the frontal
bone, we are not in a position to say.


Radiograms, both flat plates and stereograms, of children up to the age of
Radiograms, both flat plates and stereograms, of children up to the age of puberty, have yielded little information. We have seen the nasal spine in both male and female subject at 10 years, when it appears, so far as We can tell, to be in continuity with the rest of the frontal bone. Earlier than this We have been unable to make out the process and have not seen‘ it as a separate centre.
puberty, have yielded little information. We have seen the nasal spine in both
male and female subject at 10 years, when it appears, so far as We can tell,
to be in continuity with the rest of the frontal bone. Earlier than this We have
been unable to make out the process and have not seen‘ it as a separate centre.


Centre for the posterior part of the orbital plate
Centre for the posterior part of the orbital plate


As already mentioned, the orbital plate is ossified in the anterior third of
As already mentioned, the orbital plate is ossified in the anterior third of its extent simultaneously with the appearance of the frontal centre. The further spread of ossification to complete the formation of this plate is delayed for some time, excepting that part which abuts against thefuture cribiform plate. At 78 mm. S.H. ossification of the pars orbitalis is almost complete, but for a short time an inconspicuous incisura exists between the main lateral portion of this plate and the earlier formed medial part. These soon fuse. As at the periphery of the superior angle of the pars frontalis, a few islands of osseous material may precede the formation of the reticulum. None of these can be regarded as a true secondary centre, but their independent development may give rise to a Wormian bone such as that described by Von Spee. The manner in which ossification proceeds in the pars orbitalis, with the production for a time of an incisura, might give rise, unless carefully followed in many specimens, to the appearance of what we must call a pseudo-centre.
its extent simultaneously with the appearance of the frontal centre. The
further spread of ossification to complete the formation of this plate is delayed
for some time, excepting that part which abuts against thefuture cribiform
plate. At 78 mm. S.H. ossification of the pars orbitalis is almost complete,
but for a short time an inconspicuous incisura exists between the main lateral
portion of this plate and the earlier formed medial part. These soon fuse.
As at the periphery of the superior angle of the pars frontalis, a few islands of
osseous material may precede the formation of the reticulum. None of these
can be regarded as a true secondary centre, but their independent development
may give rise to a Wormian bone such as that described by Von Spee. The
manner in which ossification proceeds in the pars orbitalis, with the production
for a time of an incisura, might give rise, unless carefully followed in many
specimens, to the appearance of what we must call a pseudo-centre.
 
DISCUSSION
 
It is perfectly apparent that the conception of the development and
ossification of the frontal bone found in almost all modern textbooks is
fundamentally a repetition of the views originally expressed by Serres, and
by Rambaud & Renault, which have been widely disseminated by Jhering’s
publication. The illustrations which continue to promulgate these misconceptions appear to be, for the most part, based upon those which accompany the
treatise of Rambaud & Renault. The observations of Toldt and the opinion
of Le Double seem to have had no influence on, and to have affected in no way,
the classical description.
 
It is universally recognized that the frontal bone ossifies from two primary
centres. Mall gives the 56th day as the time of their appearance, a little later
than that given by Toldt. Our findings are essentially in agreement with those
of Mall. The statement that the primary centre of ossification appears at the
position of the future frontal eminence is, although exceedingly common,
392 Verne T. Inmcm and John B. de 0. M. Saunders
 
incorrect. Almost all original observers recognize that ossification commences
in the superciliary region, forming simultaneously part of the frontal squame
and orbital plate.


We have been unable to find any evidence, at any time in the prenatal
==Discussion==
period, of true accessory centres of ossification for the frontal bone.


It has been stated that a secondary centre appears in the region of the
It is perfectly apparent that the conception of the development and ossification of the frontal bone found in almost all modern textbooks is fundamentally a repetition of the views originally expressed by Serres, and by Rambaud & Renault, which have been widely disseminated by Jhering’s publication. The illustrations which continue to promulgate these misconceptions appear to be, for the most part, based upon those which accompany the treatise of Rambaud & Renault. The observations of Toldt and the opinion of Le Double seem to have had no influence on, and to have affected in no way, the classical description.
zygomatic process or in the neighbourhood of the pterion. This centre is
regarded by many observers as representing the post-frontal bone of premammalian skulls. We have shown that there is no such centre. What has
been accepted as such is an area ossified by extension from the primary centre
and modified by the development of the lateral two-thirds of the orbital margin
and bounded externally by the temporal line, and internally by a fissure or
series of foramina. This fissure extends on to the roof of the orbit and lies in
the floor of the lacrimal fossa. It may persist as a series of more or less extensive
foramina which occur in the region where the cribra orbitalia of Welcker (1887)
are most commonly found. While Welcker denies that cribra are to be found in
the newborn, he finds them in the skulls of young children. Both Ahrens (1905)
and Koganei (1912) state that they are found most frequently in the young.


We have noted in young skulls with weakly developed cribra that the cribra
It is universally recognized that the frontal bone ossifies from two primary centres. Mall gives the 56th day as the time of their appearance, a little later than that given by Toldt. Our findings are essentially in agreement with those of Mall. The statement that the primary centre of ossification appears at the position of the future frontal eminence is, although exceedingly common, incorrect. Almost all original observers recognize that ossification commences in the superciliary region, forming simultaneously part of the frontal squame and orbital plate.
lie in the precise region of the fissure and we suggest that there is some relationship between the development of the cribra and the blood vessels which enter
this region, as has been demonstrated for the cribra cranii.


The lateral two-thirds of the permanent supra-orbital margin arise, as
We have been unable to find any evidence, at any time in the prenatal period, of true accessory centres of ossification for the frontal bone.
detailed above, as a separate process which serves to close the orbit from the
temporal fossa. It also completes the fossa lacrimalis and, appearing late, is
not completely fused with the rest of the bone until after birth.


This process leads to the formation in the adult skull of what has been
It has been stated that a secondary centre appears in the region of the zygomatic process or in the neighbourhood of the pterion. This centre is regarded by many observers as representing the post-frontal bone of premammalian skulls. We have shown that there is no such centre. What has been accepted as such is an area ossified by extension from the primary centre and modified by the development of the lateral two-thirds of the orbital margin and bounded externally by the temporal line, and internally by a fissure or series of foramina. This fissure extends on to the roof of the orbit and lies in the floor of the lacrimal fossa. It may persist as a series of more or less extensive foramina which occur in the region where the cribra orbitalia of Welcker (1887) are most commonly found. While Welcker denies that cribra are to be found in the newborn, he finds them in the skulls of young children. Both Ahrens (1905) and Koganei (1912) state that they are found most frequently in the young.
described by Schwalbe (1906) as the planum or trigonum supra—orbitale. An
oblique groove separating the trigonum supra-orbitale from the rest of the
frontal bone is chiefly characteristic of recent man and occurs in some individuals
of all primates. This groove extends from the supra-orbital notch upwards
and laterally towards the temporal ridge, and has been named the sulcus
supra-orbitalis.


In those forms such as the gorilla, chimpanzee, and the Neanderthal race
We have noted in young skulls with weakly developed cribra that the cribra lie in the precise region of the fissure and we suggest that there is some relationship between the development of the cribra and the blood vessels which enter this region, as has been demonstrated for the cribra cranii.
in which the various supra-orbital elements are blended to form a distinctive
torus supra-orbitalis, the sulcus supra-orbitalis may be obliterated, but when
found is regarded by Cunningham (1908) as a juvenile character. Although
the sulcus is most frequently related to the supra-orbital notch and may contain
a branch of the supra-orbital nerve in part or in its entire extent, it cannot in
any way be regarded primarily as a nerve pathway.


It has not been recognized that the sulcus supra-orbitalis defines developmentally the level of the primitive orbital margin and that variations in the
The lateral two-thirds of the permanent supra-orbital margin arise, as detailed above, as a separate process which serves to close the orbit from the temporal fossa. It also completes the fossa lacrimalis and, appearing late, is not completely fused with the rest of the bone until after birth.
dimensions of the orbital rim are dependent upon the degree of development
of the supero-lateral process.
The Ossification of the H umcm Frontal Bone 393


In the hypertelerotic skull described by Greig (1924-), under-development
This process leads to the formation in the adult skull of what has been described by Schwalbe (1906) as the planum or trigonum supra—orbitale. An oblique groove separating the trigonum supra-orbitale from the rest of the frontal bone is chiefly characteristic of recent man and occurs in some individuals of all primates. This groove extends from the supra-orbital notch upwards and laterally towards the temporal ridge, and has been named the sulcus supra-orbitalis.
of this process has given to the orbital cavity an exaggerated lateral inclination,
and suppression of its medial half has profoundly modified the orbital margin
which is megaseme. Greig has pointed out that in hypertelorism there is
retention of many foetal characteristics and among them we note retention
of the primitive supra-orbital margin.


A study of the development of the eyebrow region and of the supra-orbital
In those forms such as the gorilla, chimpanzee, and the Neanderthal race in which the various supra-orbital elements are blended to form a distinctive torus supra-orbitalis, the sulcus supra-orbitalis may be obliterated, but when found is regarded by Cunningham (1908) as a juvenile character. Although the sulcus is most frequently related to the supra-orbital notch and may contain a branch of the supra-orbital nerve in part or in its entire extent, it cannot in any way be regarded primarily as a nerve pathway.
margin indicates that the essential primordia in man and the primates are
identical. The differences which exist between adult forms are the outcome of
differences in developmental direction alone.


We have demonstrated that there is no accessory-centre for the medial
It has not been recognized that the sulcus supra-orbitalis defines developmentally the level of the primitive orbital margin and that variations in the dimensions of the orbital rim are dependent upon the degree of development of the supero-lateral process.  
angular process or trochlear pit and we have described the area which has
been erroneously considered as the pre-frontal element.


Opinions differ as to whether the anterior or posterior lacrimal crest should
In the hypertelerotic skull described by Greig (1924-), under-development of this process has given to the orbital cavity an exaggerated lateral inclination, and suppression of its medial half has profoundly modified the orbital margin which is megaseme. Greig has pointed out that in hypertelorism there is retention of many foetal characteristics and among them we note retention of the primitive supra-orbital margin.
be considered as forming, for purposes of mensuration, the exact medial margin
of the orbit. Some have given the posterior crest as the true limit of the
orbital contents on the ground that it gives attachment to the septum orbitale.
Others have taken the anterior crest. It would appear that the crista lacrimalis
posterior is a continuation of the primitive orbital margin on the medial side.
The lacrimal sac is therefore at first an extra-orbital structure and only with
the formation of the new rim margin does it become potentially intra-orbital.
This change in relationship is of ontological and phylogenetic interest. Flower
(1907) has shown that in most mammals and of the primates in the lemur
alone, the lacrimal fossa is extra-orbital. In all other primates the sac is
considered to lie within the orbit.


SUMMARY
A study of the development of the eyebrow region and of the supra-orbital margin indicates that the essential primordia in man and the primates are identical. The differences which exist between adult forms are the outcome of differences in developmental direction alone.


1. An historical review of the literature on the ossification of the frontal
We have demonstrated that there is no accessory-centre for the medial angular process or trochlear pit and we have described the area which has been erroneously considered as the pre-frontal element.
bone and its accessory centres is given.


2. Observations were made upon a total of ninety-eight foetal skulls ranging
Opinions differ as to whether the anterior or posterior lacrimal crest should be considered as forming, for purposes of mensuration, the exact medial margin of the orbit. Some have given the posterior crest as the true limit of the orbital contents on the ground that it gives attachment to the septum orbitale. Others have taken the anterior crest. It would appear that the crista lacrimalis posterior is a continuation of the primitive orbital margin on the medial side. The lacrimal sac is therefore at first an extra-orbital structure and only with the formation of the new rim margin does it become potentially intra-orbital. This change in relationship is of ontological and phylogenetic interest. Flower (1907) has shown that in most mammals and of the primates in the lemur alone, the lacrimal fossa is extra-orbital. In all other primates the sac is considered to lie within the orbit.
in age from the 6th week of intrauterine life to the 10th month postnatal.


3. Primary centres of the frontal bone make their appearance in the
==Summary==
superciliary region and the details of ossifications are given.


4-. No secondary centres of ossifications have been found at any time in
# An historical review of the literature on the ossification of the frontal bone and its accessory centres is given.
the frontal bone. The changes which have given rise to the error that such
# Observations were made upon a total of ninety-eight foetal skulls ranging in age from the 6th week of intrauterine life to the 10th month postnatal.
centres exist are considered.
# Primary centres of the frontal bone make their appearance in the superciliary region and the details of ossifications are given.
# No secondary centres of ossifications have been found at any time in the frontal bone. The changes which have given rise to the error that such centres exist are considered.
# Attempts which have been made to homologize portions of the frontal bone with the pre- and post-frontal elements of premammalian skulls have been based upon the presumed existence of secondary centres. As no such centres exist in the human skull, such conclusions as have been made, on this basis, must be discarded.
# The significance of the trigonum supra-orbitalis of Schwalbe and of the sulcus supra-orbitalis as well as of other details about the orbit are discussed. 394 Verne T. Inman and John B. de 0; M. Saunders


5. Attempts which have been made to homologize portions of the frontal
bone with the pre- and post-frontal elements of premammalian skulls have
been based upon the presumed existence of secondary centres. As no such
centres exist in the human skull, such conclusions as have been made, on this
basis, must be discarded.


6. The significance of the trigonum supra-orbitalis of Schwalbe and of
In conclusion it is with great pleasure We acknowledge our indebtedness to Mrs L. G. Philippe for secretarial assistance, and to Mr Rudolph Skarda for his skill in the preparation of material
the sulcus supra-orbitalis as well as of other details about the orbit are discussed.
394 Verne T. Inman and John B. de 0; M. Saunders


In conclusion it is with great pleasure We acknowledge our indebtedness
==References==
to Mrs L. G. Philippe for secretarial assistance, and to Mr Rudolph Skarda
for his skill in the preparation of material


REFERENCES
AHRENS, E. (1905). Inaug. Diss. Géttingen (1904), abst. in Jber. Leist. 0phthal., 1905, p. 39.


AHRENS, E. (1905). Inaug. Diss. Géttingen (1904), abst. in Jber. Leist. 0phthal., 1905, p. 39.
ALBINUS, BERNARD S. (1737). Ioones ossium foetus humani accedit osteogeniae bremls historia. Leidae Batavorum, p. 10.
ALBINUS, BERNARD S. (1737). Ioones ossium foetus humani accedit osteogeniae bremls historia.
Leidae Batavorum, p. 10.


ASHLEY-MONTAGU, M. F. (1931). J. Anat., Lond., vol. Lxv, p. 446.
ASHLEY-MONTAGU, M. F. (1931). J. Anat., Lond., vol. Lxv, p. 446.
Line 571: Line 185:
GALEN (1542). Opera Omnia. Froben. Basileae.
GALEN (1542). Opera Omnia. Froben. Basileae.


GEGENBAUR, C. (1899). Lehrbuch der Anatomic des Menschen. Bd. 1, S. 220. Leipzig.
GEGENBAUR, C. (1899). Lehrbuch der Anatomic des Menschen. Bd. 1, S. 220. Leipzig. GRAY, H. (1930). Anatomy of the Human Body (22nd ed.), p. 134. Philadelphia.
GRAY, H. (1930). Anatomy of the Human Body (22nd ed.), p. 134. Philadelphia.


GREIG, D. M. (1924). Edinb. med. J. (N .S.), vol. XXXI, p. 560.
GREIG, D. M. (1924). Edinb. med. J. (N .S.), vol. XXXI, p. 560.
Line 586: Line 199:
KOGANEI, Y. (1912). Mitt. med. Falc. Tokio, Bd. x, S. 113.
KOGANEI, Y. (1912). Mitt. med. Falc. Tokio, Bd. x, S. 113.


KOLLIKER, ALBERT (1879). Entwickelungsgeschichte des M enschen, S. 455. Leipzig.
KOLLIKER, ALBERT (1879). Entwickelungsgeschichte des M enschen, S. 455. Leipzig. LECOURTOIS, M. (1869). Bull. Soc. Anat. Paris, 1;. xLrv, p. 478.
LECOURTOIS, M. (1869). Bull. Soc. Anat. Paris, 1;. xLrv, p. 478.


LE DOUBLE, A. P. (1903). Traité des variations des 03 dn crane de l’homme, pp. 201-15. Paris.
LE DOUBLE, A. P. (1903). Traité des variations des 03 dn crane de l’homme, pp. 201-15. Paris. MAGGI, L. (1897). Arch. ital. Biol. t. xxvm, p. 329.
MAGGI, L. (1897). Arch. ital. Biol. t. xxvm, p. 329.


—— (1898). Arch. ital. Biol. 1;. xxx, p. 161.
—— (1898). Arch. ital. Biol. 1;. xxx, p. 161.
Line 622: Line 233:
TESTUT, L. (1921). Traitef d’Anatomie Hurmzine, t. I, p. 126. Paris.
TESTUT, L. (1921). Traitef d’Anatomie Hurmzine, t. I, p. 126. Paris.


TOLDT, C. (1881-82). Maschka’s Handbnch der gerichtlichen M edizin, Bd. III, S. 524.
TOLDT, C. (1881-82). Maschka’s Handbnch der gerichtlichen M edizin, Bd. III, S. 524. VEsAL1Us, ANDREAS (1564). Observationes Gabrielia Fallopii Exam. Venitiis.
VEsAL1Us, ANDREAS (1564). Observationes Gabrielia Fallopii Exam. Venitiis.


WELCKER, H. (1887). Arch. Anthrop., Braunschw., Bd. xvII, S. 1.
WELCKER, H. (1887). Arch. Anthrop., Braunschw., Bd. xvII, S. 1.




==Explanation of Plate I==


EXPLANATION OF PLATE I
fig. 1. Untouched photograph (infra-red, Wratten filter No. 25) of norms. lateralis of an ossifying human foetal skull, sitting height 80 mm. (13-5 weeks, Streeter’s tables).


fig. 1. Untouched photograph (infra-red, Wratten filter No. 25) of norms. lateralis of an ossifying
fig. 2. Untouched photograph (infra-red, Wratten filter No. 25) of norma facialis of an ossifying human foetal skull, sitting height 126 mm. (16-5 weeks, Streeter’s tables).
human foetal skull, sitting height 80 mm. (13-5 weeks, Streeter’s tables).


fig. 2. Untouched photograph (infra-red, Wratten filter No. 25) of norma facialis of an ossifying
{{Historic Disclaimer}}
human foetal skull, sitting height 126 mm. (16-5 weeks, Streeter’s tables).






{{Historic Disclaimer}}
{{Footer}}
{{Footer}}
[[Category:Skull]][[Category:Bone]][[Category:1930's]][[Category:Draft]]
[[Category:Skull]][[Category:Bone]][[Category:1930's]][[Category:Draft]]

Revision as of 10:02, 6 January 2017

Embryology - 19 Apr 2024    Facebook link Pinterest link Twitter link  Expand to Translate  
Google Translate - select your language from the list shown below (this will open a new external page)

العربية | català | 中文 | 中國傳統的 | français | Deutsche | עִברִית | हिंदी | bahasa Indonesia | italiano | 日本語 | 한국어 | မြန်မာ | Pilipino | Polskie | português | ਪੰਜਾਬੀ ਦੇ | Română | русский | Español | Swahili | Svensk | ไทย | Türkçe | اردو | ייִדיש | Tiếng Việt    These external translations are automated and may not be accurate. (More? About Translations)

Inman VT. and Saunders JBdeCM. The ossification of the human frontal bone with special reference to its presumed pre- and post-frontal elements. (1937) J. Anat., 71(3): 384-394.

Online Editor  
Mark Hill.jpg


Musculoskeletal - Skull Development | Head Development | Bone Development

Historic Disclaimer - information about historic embryology pages 
Mark Hill.jpg
Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)

The Ossification of the Human Frontal Bone with Special Reference to its Presumed Pre- and Post-Frontal Elements

By Verne T. Inman, M.D., Ph.D. And John B. De C. M. Saunders, M.B., F.R.C.S. (Edin.)


Division of Anatomy, University of California Medical School

Introduction

In a recent number of the Journal of Anatomy there has appeared an article by Ashley-Montagu on the presumed existence of a post-frontal element in the skull of a young gorilla. Ashley-Montagu’s conclusions are based largely on the view that the post-frontal bone "is present in the developing skull of probably all primates and certainly of all men. “ It is not Widely known among anatomists”, he says, “that the post-frontal bone is a constant element of the developing human skull. For some extraordinary reason most modern textbooks of anatomy and embryology make no mention of the post-frontal element of the human foetal skull in spite of the fact that a constant and independent centre of ossification of this bone is present in every case, and the bone itself may generally be observed up to the end of the third foetal month, and not rarely in the skulls of newborn children.” .


Having available a considerable collection of material specifically prepared to demonstrate the ossification of the skull in man, we were prompted to seek for this element. We were surprised, in the light of the above definite statement, that we were unable to find the slightest trace of this element occurring as an independent centre, nor were we at any time able to find such other secondary centres of ossification as have been commonly described for the frontal bone. We noted, furthermore, that there was some disagreement among the authors of various standard textbooks, and it became increasingly apparent that in view of the great morphological and phylogenetic conclusions which have been based upon the supposed existence of such secondary centres, reinvestigation of the ossification of the frontal bone was necessary.

Literature

Gabriel Fallopio appears to have been one of the first to state clearly that the frontal bone constantly develops from two symmetrical halves. In his Observatimws Anatomicae (1561) he says: “Ita ut in omnibus puerulis qui tenerimi sunt frontis os ex duabus constare partibus reperierim.” Vesalius was also well aware of this mode of origin of the frontal bone, for not only does he discuss Fallopio’s observations in his Observationes Gabrielis Falloppii Ezvamen (1564), but the matter is mentioned in the Fabrica (1543). Jhering states that Galen was ignorant of this fact, but it would seem on examining his De usu partium and De admim'stratiom'bus anatomicis that he may have been aware of this in some instances.

Up to the time of Kerckring (1670) but few serious attempts had been made to examine the origin of the bones before birth. This author dissected and described specimens from approximately the second month of foetal life.

Albinus illustrates in his Icones ossium foetus humani (1737) that area which has since been called the post-frontal element and records an instance in which each half of the frontal bone was divided into three parts. This single observation was made on an aborted foetus. He says: “In abortu juniore, cui totum os magnitudinis unguis mediocris, utramque illam partem inveni divisam in tres; superiorem, quae ad frontem pertinet; inferiorem, quae ad foramen oculi, tertiam quae ad caput supercilii.”

Apart from the single observation quoted above from Albinus, the recognition of secondary or accessory centres of ossification dates from the work of Serres (l829).1 This author described accessory centres, (1) for the part of the pars orbitalis at the trochlear fossa and (2) for the zygomatic process. Serres considered them, in association with his studies on the comparative anatomy of the skull, as representing the pre- and post-frontal elements. Rambaud & Renault (1864) accept the findings of Serres, but do not appear to have seen these centres as independent ossicles. They state that at the fourth month the centres have already been joined by a suture which leaves only a shallow groove and that the centres have always so appeared at this time. These authors added a third centre for the nasal spine which, they state, appears late, sometime after the eighth year.

A fourth centre, for the posterior part of the pars orbitalis, is given by Von Spec (1896) in von Bardeleben’s H andbmth. The existence of such a centre appears to be based on an observation originally made by Hyrtl (1860) that a Wormian bone is sometimes found in this region. Von Spee also observed such a Wormian bone and accepts Hyrtl’s conclusion that this has arisen from an independent centre of ossification. It has frequently been pointed out since (Patten, 1912) that the presence of a Wormian bone cannot be accepted as evidence of a true secondary centre.

Since the publication of Rambaud & Renault’s monograph, a number of special papers have appeared on the ossification of the frontal bone and on its pre- and post-frontal elements. Lecourtois (1869) published a study based upon foetal skulls of 5-6 months, in which he expresses agreement with the findings of Serres and of Rambaud & Renault.

1 Many authors give as references to the work of Serres on accessory centres the C'.R. Acad. S’ci., Paris, 1819 and 1838. These abstracts contain no mention of specific centres. Rambaud & Renault also give as reference Serres, Des Lois de l’oatéoge’m'e, Paris, 1829. This work is unavailable to us. It would seem that the majority of authors have quoted Serres through Rambaud & Renault.

In 1872, von Jhering published an important paper which has since been widely quoted. J hering, after reviewing the work of earlier authors, confirms the conclusions arrived at by Rambaud & Renault. He examined, however, no specimen earlier than the 4th month and at no time saw the assumed accessory centres as independent ossicles. Maggi (1897, 1898) published two extensive articles on the morphology of the cranial and cranio-facial bones in which he concludes that the existence of accessory centres for the frontal bone has been satisfactorily demonstrated, and that two of these centres are homologous with the pre- and post-frontal elements of other forms. His observations on man were based on the examination of foetal skulls from the 5th month and on the work of Rambaud & Renault. finally Von Bardeleben (1896), Augier (1912) and Ashley-Montagu (1931) have Written articles which, on analysis, present no original observations on the development of the frontal bone, but whose conclusions are either directly or indirectly based on the acceptance of the work of Serres and of Rambaud & Renault.

It is not surprising in consequence, that the standard textbooks of anatomy and of embryology almost uniformly describe and illustrate, apart from the primary centres, three pairs of accessory centres: (1) for the frontal spine, (2) variously for the medial angular process or trochlear pit (pre-frontal element), and (3) for the region of the zygomatic process (post-frontal element). These accessory centres, apart from that for the frontal spine, are, in the general concensus of opinion, assumed to appear about the 4th month and to fuse at the 7th. The centre for the nasal spine is recognized as appearing late, at or before the 10th year. In addition Quain and Von Spee (von Bardeleben’s H andbuch) mention the occasional occurrence of a fourth pair for the posteromedial part of the orbital plate. The statements in Quain, Cunningham, Gray, Morris, Piersol, Gegenbaur, Merkel, von Bardeleben, Testut, and Poirier, are, as in the special articles noted above, a continuation of the opinion of Serres

and of Rambaud & Renault.

The only dissenting opinions which we have been able to find in the literature are those of Toldt and Le Double. Toldt, while recognizing the areas described by Serres and by Rambaud & Renault, is unable to accept these as indicating the existence of accessory centres. In Maschka’s Hcmdbuch d. gerichtlichen Medizin he says: “Ich habe etwas Derartiges beim Menschen nie gesehen und muss daher die Angaben von Serres, Rambaud u. Renault, welche durch J hering auch in die deutsche Literatur iibergegangen sind, wonach beim Menschen ein urspriinglich selbstandiges Os frontale posterius bestehe, in Abrede stellen. Ebenso wenig kann ich fiir den Menschen die Existenz eines zu irgend einer Zeit isolierten Os frontale anterius einraumen.”

Le Double discusses extensively the ontogeny and phylogeny of the preand post-frontal elements and points out the lack of agreement among anatomists. The existence of accessory centres in man has arisen, in his opinion, from misinterpretation of the findings.

This survey would indicate that the present belief that accessory centres for the frontal bone exist in man depends in large measure on the observations and interpretation of Serres and of Rambaud & Renault. In spite of the denial of their existence by Toldt, whose opinion is unfortunately buried in a textbook of forensic medicine, and the questioning of Le Double, all modern textbooks of anatomy continue to illustrate and describe such centres. Furthermore their existence forms the chief basis for their acceptance as pre— and post—frontal elements.

Material and Methods

Our observations are based on a total of ninety-eight specimens ranging from the 6th week of intra-uterine life to the 10th month, post-natal. All the material was Caucasian. The majority of the specimens were prepared to demonstrate ossification by a technique developed in this department. This method is much more satisfactory than that of Schultze in that it enables one to follow the progress of ossification to term. The method, published elsewhere by R. Skarda (1934), briefly consists in the rapid dissection of fresh foetal material which has previously been thoroughly freed from blood by washing via the umbilical vessels. The specimens are then hardened in 95 per cent alcohol for 48 hours and cleared in 1 per cent solution of NaOH and finally permanently mounted in 2 per cent formalin solution. The clearing process is carried out in darkness which improves the development of the final preparation. This technique results in preparations of great beauty. The bone is pure white, and its finest ramifications can be traced with case when viewed in good light under the dissecting microscope. The cartilage remains opalescent blue in colour. Shrinkage is approximately 2 per cent and distortion negligible. The specimens are mounted intact. This is a great advantage in that relations are maintained and there is little danger of creating artifacts. Furthermore the fact that the specimens are wet avoids cracking of the delicate bone which has given rise to so many of the erroneous observations of earlier workers on ossification.

The above technique demands absolutely fresh material. Foetuses of 6 months or over, if not fresh, tend to be opaque, and the ramifications of the delicate trabeculae cannot be made out with the same clarity. This can be remedied by carrying the preparation a stage further. The less satisfactory material is carefully dehydrated by carrying it successively from 50 per cent to absolute alcohol in steps of 5 degrees and then transferring to pure benzol. Specimens prepared in this second fashion show distinctly more shrinkage, and although the most delicate bone centres can be seen and readily followed, they do not have the same elegance as the others, nor do they have the same permanency. '

In addition, dissections were made of formalinized specimens and a few dried foetal skulls examined.

Ossification of the Frontal Bone

The primary centres

It is universally agreed that the frontal bone ossifies from two primary centres which appear in membrane between the fortieth and fiftieth day. There is some disagreement however with regard to the precise position in which these centres arise. The majority of investigators and most standard textbooks describe ossification as beginning in the position of the future frontal eminences; the minority place the centres at the superciliary region.

We /have observed the frontal bone appearing as two finely reticulated areas of ossification in specimens of 32 and 36 mm. sitting height (s.H.). In these and in all other specimens of approximately the same age ossification involves an area whose long axis corresponds approximately to the position of the superciliary arch, and which forms the lower part of the frontal squame and the more anterior portion of the orbital plate and a. primitive medial angular process. In our younger specimens this ossific centre is about 2 by 3 mm. The ossific meshwork is of regular and even density throughout this area. except for a slight thickening and condensation which forms a line in the position of the primitive supraorbital margin. In slightly older specimens a few fine, less dense trabeculae project radially from the periphery of the centre. These trabeculae are best defined along the upper margin of the developing pars frontalis.

After the appearance of the primary centres, ossification spreads more rapidly in the pars frontalis than in the pars orbitalis. Ossification proceeds in the pars frontalis by the formation of heavy islands of bone which constitute a crescentic area whose maximum depth corresponds to the future posterosuperior angle (see Pl. I, fig. 1). These islands fuse to form a series of primary radiating trabeculae (Pl. I, fig. 2). The appearance of secondary trabeculae serves to link the primary trabeculae, expanding the even meshwork of the primary centre. This expansion of the meshwork of the primary centre behind the advancing zone of ossification gives rise to the appearance of a centre in the position of the frontal eminence from which the primary trabeculae radiate, an effect which is often seen in specimens of 4 months and older. In fact, however, this false centre of radiation rises higher and higher up on the frontal squame with increasing age.

In the pars orbitalis the progress of ossification is less rapid. The radiating trabeculae so characteristic of the pars frontalis are very much smaller and less obvious and an expanding meshwork of trabeculae is maintained.

These observations show that ossification begins in the region of the superciliary arch and simultaneously gives rise to a portion of both the pars frontalis and the pars orbitalis and to the primitive medial angular process. The common statement that ossification commences at the site of the future frontal eminence is erroneous, and is undoubtedly the outcome of observations made on older specimens.


The secondary centres

In the ninety-eight specimens examined we have at no time been able to find any true accessory centres of ossification. No centres appear in those regions in which they have been stated to occur. In all instances the bony trabeculae were found to be continuous and unbroken. Furthermore, the regions in which such accessory centres are presumed to arise do not correspond to those in which secondary island formation is found. The specimens on which we base our observations are of such clarity that there is little possibility that such centres, had they existed, could have been overlooked. None the less, if dried preparations of foetal skulls between the third and seventh months, and in some at term, are examined, there are appearances which suggest the possible existence of secondary centres. It is these appearances which have, no doubt, been accepted by many authors as indicating the existence of accessory centres. We shall therefore consider the changes which occur in those areas which have given rise to error in interpretation.

(1) The presumed centre for the zygomatic process or post-frontal element.

The primary centre of ossification gives rise to a part of the frontal and orbital plates and a primitive medial angular process. The bony superior orbital margin seen at this period does not correspond to that of the adult. The lateral two-thirds from the position of the future supra-orbital notch to the zygomatic process is due to an entirely new formation. At 42 mm. s.H. a linear thickening of osseous material appears in the lateral two-thirds of the orbital margin at the junction between the pars orbitalis and pars frontalis. This thickening becomes more and more apparent as it is followed laterally, and has resulted in the formation at birth of almost the entire supero-lateral wall of the orbit as well as giving rise to the zygomatic process, and thereby completes the separation of the orbital cavity from the temporal fossa. At 49 mm. S.H. it has spread to form a recognizable zygomatic process which is more apparent at 59 mm. s.H. (see Pl. I, fig. 2). A considerable interval exists between this process and the frontal process of the zygomatic bone; an interval which persists until the foetus has attained a length of 250 mm. The closure of this interval. is due to a further extension of ossification in the frontal process of the zygomatic bone rather than to any further ossification in the zygomatic process of the frontal, for at 155 mm. s.H. the proportions of the zygomatic process of the frontal have almost attained those seen at birth.


From 49 mm. s.H. onwards the base of the zygomatic process of the frontal, including the lateral two-thirds of the orbital margin, progressively thickens. This thickening extends dorsally on to the lateral surface of the pars frontalis but does not involve its postero-lateral angle. This angle is, as a result of this thickening, bounded antero-superiorly by a ridge. The ridge and the area bounded by it becomes, with further growth of the skull, more and more apparent until, in any specimen from the third month on, a pseudo-suture appears to separate off this angle, giving rise to the appearance of an autonomous centre. In a macerated skull at 7 months, the ridge extends upwards and dorsally from the posterior border of the zygomatic process (lateral angular process) to a point some 5 mm. above the pterion (Text-fig. 2, l.t.). This ridge delimits a quadrilateral area of bone, approximately 5mm. in height and 15 mm. in length, which includes the postero-lateral angle (Text-fig. 2, p. f.). Dissections and histological sections demonstrate that this ridge gives attachment ‘to the temporal muscle and is none other than the linea temporalis. This is confirmed by the finding, in a few specimens, of a continuation of the ridge on to the antero-inferior angle of the parietal bone.


Text-fig. 1. Postero-inferior view of the left disarticulated frontal bone of 7 months’ old human foetus. P.F. pars frontalis; P.0. pars orbitalis; l.t. linea temporalis; p. “presumed” post-frontal element; f. superior limb of fissure separating supero-lateral process from the pars orbitalis.

Text-fig. 2. Lateral View of the facial region of a full-term human foetal skull. p. “presumed” post-frontal element; l.t. linea temporalis; pr. “presumed” pre-frontal element; 7'. ridge separating the medial angular process from the pars frontalis.

In the disarticulated frontal bone, the new process which springs from the margin between the pars frontalis and orbitalis, and which completes the supero-lateral wall of the orbit and gives rise to the zygomatic process, is separated postero-laterally, from the rest of the pars orbitalis, by a deep fissure (Text-fig. 1, f.). This fissure gives attachment to the membrane filling the lateral fontanelle. The inferior limb of the fissure extends on to the roof of the orbital cavity, in the floor of the lachrymal fossa. The inferior limb of the fissure is obliterated, medio-laterally, at birth, but usually persists as a linear series of small foramina and is often evident in the adult. The presence of this fissure in the later months of intra-uterine life further falsely accentuates the appearance of an independent centre of ossification in this area.

The area outlined by the linea temporalis and the fissure described above constitutes what has been designated as the post-frontal element of the human 390 Verne T. Inman and frontal bone, and has been regarded as arising from an autonomous secondary centre of ossification.

(2) The presumed centre for the medial angular process (trochlear pit) or prefrontal element.

A primitive medial angular process is ossified simultaneously with the appearance of the primary ossific centre. This process appears to be a continuation of the superciliary margin medially and forms a small portion of the anterior part of the medial wall of the orbit. At 49 mm. S.H. only the more anterior portion of the pars orbitalis is osseous. At this time ossification is relatively more extensive in the medial angular process.

Ossification spreads much more slowly in the orbital roof and in consequence an unossified interval persists, until the 4th month, between that part of the medial angular process which lies in the medial wall of the orbit and the roof.

At 59 mm. S.H. the medial angular process commences to show some thickening of its reticular network, Whereas at this time scarcely a third of the orbital plate is ossified.

At 80 mm. S.H. ossification has spread to complete the major part of the orbital plate and it commences to fuse with the medial angular process, obliterating the interval between these structures. This line of fusion is indicated at birth either by a groove, or by a series of foramina resembling a suture, which rums obliquely across the trochlear pit to the anterior end of the fronto-ethmoidal suture (Text-fig. 2).

At 59 mm. S.H. a slight thickening extends medially from the position of the future supra-orbital notch across the base of the medial angular process to the metopic suture (Pl. I, fig. 2). This thickening indicates for the first time the boundary of the permanent orbital rim. At 99 mm. S.I-I. the trabeculae have thickened to such an extent as to produce a ridge which separates the medial angular process from the pars frontalis, a ridge which is very apparent at term (Text-fig. 2, r.). As a result, the superior orbital margin of the foetus is Well defined medially. A new margin is therefore formed anterior to the primitive margin which, if traced downwards, becomes continuous with the crista lacrimalis anterior. Between these two margins lies the fossa sacci lacrimalis and the lacrimal sac.

In the later months of foetal life a quadrilateral area of bone is defined in the trochlear region of the macerated frontal bone. This area is bounded anteriorly by the ridge of bone which crosses the base of the medial angular process, superiorly by a groove or series of foramina, inferiorly by the frontomaxillary, fronto-lacrimal and anterior part of the fronto-ethmoidal sutures (Text-fig. 2, pr.f.). Its surface is rough and irregular. It is this area which has been almost universally regarded as the accessory centre for the trochlear

pit or medial angular process and which has been designated as the pre-frontal element of man.


The centre for the nasal

At no time prior to birth have we ever observed the presence of a secondary centre for the nasal spine. This process remains unossified for some time after birth. Whether the spine ossifies from secondary centres at or about the 12th year, as is usually stated, or whether it ossifies as an extension from the frontal bone, we are not in a position to say.

Radiograms, both flat plates and stereograms, of children up to the age of puberty, have yielded little information. We have seen the nasal spine in both male and female subject at 10 years, when it appears, so far as We can tell, to be in continuity with the rest of the frontal bone. Earlier than this We have been unable to make out the process and have not seen‘ it as a separate centre.

Centre for the posterior part of the orbital plate

As already mentioned, the orbital plate is ossified in the anterior third of its extent simultaneously with the appearance of the frontal centre. The further spread of ossification to complete the formation of this plate is delayed for some time, excepting that part which abuts against thefuture cribiform plate. At 78 mm. S.H. ossification of the pars orbitalis is almost complete, but for a short time an inconspicuous incisura exists between the main lateral portion of this plate and the earlier formed medial part. These soon fuse. As at the periphery of the superior angle of the pars frontalis, a few islands of osseous material may precede the formation of the reticulum. None of these can be regarded as a true secondary centre, but their independent development may give rise to a Wormian bone such as that described by Von Spee. The manner in which ossification proceeds in the pars orbitalis, with the production for a time of an incisura, might give rise, unless carefully followed in many specimens, to the appearance of what we must call a pseudo-centre.

Discussion

It is perfectly apparent that the conception of the development and ossification of the frontal bone found in almost all modern textbooks is fundamentally a repetition of the views originally expressed by Serres, and by Rambaud & Renault, which have been widely disseminated by Jhering’s publication. The illustrations which continue to promulgate these misconceptions appear to be, for the most part, based upon those which accompany the treatise of Rambaud & Renault. The observations of Toldt and the opinion of Le Double seem to have had no influence on, and to have affected in no way, the classical description.

It is universally recognized that the frontal bone ossifies from two primary centres. Mall gives the 56th day as the time of their appearance, a little later than that given by Toldt. Our findings are essentially in agreement with those of Mall. The statement that the primary centre of ossification appears at the position of the future frontal eminence is, although exceedingly common, incorrect. Almost all original observers recognize that ossification commences in the superciliary region, forming simultaneously part of the frontal squame and orbital plate.

We have been unable to find any evidence, at any time in the prenatal period, of true accessory centres of ossification for the frontal bone.

It has been stated that a secondary centre appears in the region of the zygomatic process or in the neighbourhood of the pterion. This centre is regarded by many observers as representing the post-frontal bone of premammalian skulls. We have shown that there is no such centre. What has been accepted as such is an area ossified by extension from the primary centre and modified by the development of the lateral two-thirds of the orbital margin and bounded externally by the temporal line, and internally by a fissure or series of foramina. This fissure extends on to the roof of the orbit and lies in the floor of the lacrimal fossa. It may persist as a series of more or less extensive foramina which occur in the region where the cribra orbitalia of Welcker (1887) are most commonly found. While Welcker denies that cribra are to be found in the newborn, he finds them in the skulls of young children. Both Ahrens (1905) and Koganei (1912) state that they are found most frequently in the young.

We have noted in young skulls with weakly developed cribra that the cribra lie in the precise region of the fissure and we suggest that there is some relationship between the development of the cribra and the blood vessels which enter this region, as has been demonstrated for the cribra cranii.

The lateral two-thirds of the permanent supra-orbital margin arise, as detailed above, as a separate process which serves to close the orbit from the temporal fossa. It also completes the fossa lacrimalis and, appearing late, is not completely fused with the rest of the bone until after birth.

This process leads to the formation in the adult skull of what has been described by Schwalbe (1906) as the planum or trigonum supra—orbitale. An oblique groove separating the trigonum supra-orbitale from the rest of the frontal bone is chiefly characteristic of recent man and occurs in some individuals of all primates. This groove extends from the supra-orbital notch upwards and laterally towards the temporal ridge, and has been named the sulcus supra-orbitalis.

In those forms such as the gorilla, chimpanzee, and the Neanderthal race in which the various supra-orbital elements are blended to form a distinctive torus supra-orbitalis, the sulcus supra-orbitalis may be obliterated, but when found is regarded by Cunningham (1908) as a juvenile character. Although the sulcus is most frequently related to the supra-orbital notch and may contain a branch of the supra-orbital nerve in part or in its entire extent, it cannot in any way be regarded primarily as a nerve pathway.

It has not been recognized that the sulcus supra-orbitalis defines developmentally the level of the primitive orbital margin and that variations in the dimensions of the orbital rim are dependent upon the degree of development of the supero-lateral process.

In the hypertelerotic skull described by Greig (1924-), under-development of this process has given to the orbital cavity an exaggerated lateral inclination, and suppression of its medial half has profoundly modified the orbital margin which is megaseme. Greig has pointed out that in hypertelorism there is retention of many foetal characteristics and among them we note retention of the primitive supra-orbital margin.

A study of the development of the eyebrow region and of the supra-orbital margin indicates that the essential primordia in man and the primates are identical. The differences which exist between adult forms are the outcome of differences in developmental direction alone.

We have demonstrated that there is no accessory-centre for the medial angular process or trochlear pit and we have described the area which has been erroneously considered as the pre-frontal element.

Opinions differ as to whether the anterior or posterior lacrimal crest should be considered as forming, for purposes of mensuration, the exact medial margin of the orbit. Some have given the posterior crest as the true limit of the orbital contents on the ground that it gives attachment to the septum orbitale. Others have taken the anterior crest. It would appear that the crista lacrimalis posterior is a continuation of the primitive orbital margin on the medial side. The lacrimal sac is therefore at first an extra-orbital structure and only with the formation of the new rim margin does it become potentially intra-orbital. This change in relationship is of ontological and phylogenetic interest. Flower (1907) has shown that in most mammals and of the primates in the lemur alone, the lacrimal fossa is extra-orbital. In all other primates the sac is considered to lie within the orbit.

Summary

  1. An historical review of the literature on the ossification of the frontal bone and its accessory centres is given.
  2. Observations were made upon a total of ninety-eight foetal skulls ranging in age from the 6th week of intrauterine life to the 10th month postnatal.
  3. Primary centres of the frontal bone make their appearance in the superciliary region and the details of ossifications are given.
  4. No secondary centres of ossifications have been found at any time in the frontal bone. The changes which have given rise to the error that such centres exist are considered.
  5. Attempts which have been made to homologize portions of the frontal bone with the pre- and post-frontal elements of premammalian skulls have been based upon the presumed existence of secondary centres. As no such centres exist in the human skull, such conclusions as have been made, on this basis, must be discarded.
  6. The significance of the trigonum supra-orbitalis of Schwalbe and of the sulcus supra-orbitalis as well as of other details about the orbit are discussed. 394 Verne T. Inman and John B. de 0; M. Saunders


In conclusion it is with great pleasure We acknowledge our indebtedness to Mrs L. G. Philippe for secretarial assistance, and to Mr Rudolph Skarda for his skill in the preparation of material

References

AHRENS, E. (1905). Inaug. Diss. Géttingen (1904), abst. in Jber. Leist. 0phthal., 1905, p. 39.

ALBINUS, BERNARD S. (1737). Ioones ossium foetus humani accedit osteogeniae bremls historia. Leidae Batavorum, p. 10.

ASHLEY-MONTAGU, M. F. (1931). J. Anat., Lond., vol. Lxv, p. 446.

AUGIER, M. (1912). 0.12. Assoc. Anat. t. XIV, p. 22.

BARDELEBEN, K. v. (1896). Anat. Anz. (Ergiinzungsheft), Bd. XII, S. 153-4.

CUNNINGHAM, D. J. (1908). Tr. Roy. Soc. Erlin. vol. xLvI, p. 283.

CUNNINGHAM (1931). Textbook of Anatomy (6th ed.), p. 188.

FALLOPIO, GABRIEL (1561). Obaervationes Anatomioae.

FLOWER, T. (1907). (As quoted by Whitnall.) Catalogue of the Museum of the Royal College of

Surgeons (2nd ed.). London.

GALEN (1542). Opera Omnia. Froben. Basileae.

GEGENBAUR, C. (1899). Lehrbuch der Anatomic des Menschen. Bd. 1, S. 220. Leipzig. GRAY, H. (1930). Anatomy of the Human Body (22nd ed.), p. 134. Philadelphia.

GREIG, D. M. (1924). Edinb. med. J. (N .S.), vol. XXXI, p. 560.

HYRTL, J. (1860). Quoted by von Spee.

JHEBING, H. v. (1872). Arch. f. Amt. Phys. n. Wise. Med. von Reichert und Du Bois-Reymond,

S. 649-58.

KERCKRING, TH. (1670). Sptcilegium anatomicum, etc. Amstelodami.

KOGANEI, Y. (1912). Mitt. med. Falc. Tokio, Bd. x, S. 113.

KOLLIKER, ALBERT (1879). Entwickelungsgeschichte des M enschen, S. 455. Leipzig. LECOURTOIS, M. (1869). Bull. Soc. Anat. Paris, 1;. xLrv, p. 478.

LE DOUBLE, A. P. (1903). Traité des variations des 03 dn crane de l’homme, pp. 201-15. Paris. MAGGI, L. (1897). Arch. ital. Biol. t. xxvm, p. 329.

—— (1898). Arch. ital. Biol. 1;. xxx, p. 161.

MALL, FRANKLIN P. (1906). Amer. J. Anat. vol. v, p. 433.

MERKEL, F. (1913). Textbook of Anatomy (6th ed.), p. 188.

MORRIS, H. (1914). Human Anatomy, p. 62. Philadelphia.

PATTEN, C. J. (1912). Z. Morph. Anthr. Bd. xrv, S. 527.

PIERSOL (1930). Hmmm Anatomy (9th ed.). Philadelphia.

POIRIER, P., CHARPY, A. & CUNEO, B. (1908). Abrége’ d’Anatomie, t. I, p. 147. Paris.

QUAIN (1915). Anatomy, vol. Iv, pt. I, p. 61. London.

RAMBAUD & RENAULT (1864). Origine et de’veloppement des os. Paris.

SCHWALBE, G. (1906). (As quoted by Cunningham, 1908.)

SERRES, MARCEL DE (1829). (Quoted by Rambaud & Renault.)

SKARDA, R. (1934). J. tech. Meth. vol. XIII, pp. 38-40.

SPEE, F. G. v. (1896). Von Bax-deleben’s Handbuch der Anatormle, S. 123. Jena.

STREETER, G. I. (1920). Oontr. Embryol. Carney. I nstn, vol. XI, No; 55.

TESTUT, L. (1921). Traitef d’Anatomie Hurmzine, t. I, p. 126. Paris.

TOLDT, C. (1881-82). Maschka’s Handbnch der gerichtlichen M edizin, Bd. III, S. 524. VEsAL1Us, ANDREAS (1564). Observationes Gabrielia Fallopii Exam. Venitiis.

WELCKER, H. (1887). Arch. Anthrop., Braunschw., Bd. xvII, S. 1.


Explanation of Plate I

fig. 1. Untouched photograph (infra-red, Wratten filter No. 25) of norms. lateralis of an ossifying human foetal skull, sitting height 80 mm. (13-5 weeks, Streeter’s tables).

fig. 2. Untouched photograph (infra-red, Wratten filter No. 25) of norma facialis of an ossifying human foetal skull, sitting height 126 mm. (16-5 weeks, Streeter’s tables).

Historic Disclaimer - information about historic embryology pages 
Mark Hill.jpg
Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)



Cite this page: Hill, M.A. (2024, April 19) Embryology Paper - The ossification of the human frontal bone with special reference to its presumed pre- and post-frontal elements. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Paper_-_The_ossification_of_the_human_frontal_bone_with_special_reference_to_its_presumed_pre-_and_post-frontal_elements

What Links Here?
© Dr Mark Hill 2024, UNSW Embryology ISBN: 978 0 7334 2609 4 - UNSW CRICOS Provider Code No. 00098G