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The capsular matrix

Melvin 1. Moss, D.D.S., Ph.D., and Letty Salentijn, D.D.S.


New Pork, N. P.

I t is commonly agreed that a comprehensive knowledge of cranial


growth is a sine qua non for any rational approach to orthodontic therapy.
Quite apart from the intrinsic value of such information, many aspects of ortho-
dontic diagnosis, therapy, and prognosis are based firmly on this aspect of
cranial biology. Far from being an academic matter, it is easily demonstrated
that the funda.mental conceptualizations of the orthodontist concerning the pro-
cess of cranial growth strongly influence his therapeutic techniques. Some years
ilgo it appeared that the major problems of cranial growth processes had been
solved; sutures were thought to be sites of primary expansive growth of the
fla.t hones; the nasal, mandibular, condylar, and spheno-occipital cartilages were
thought to be primary sites of expansive growth, while the processes of osseous
deposition and resorption completed the roster of growt,h agencies. Indeed, it
is no exaggeration to say that cranial growth was considered to be the semantic
(and conceptual) equivalent of skull growth. The processes by which the skeletal
tissues effect any alteration of their size and shape were mistakenly considered
to be identical with the factors which cause these same alterations. Unfortu-
nately, it is true that the classic approaches of anatomy and anthropology in
general, and of craniology in particular, served to reinforce this erroneous view-
point and to embed it even deeper in each succeeding professional generation.
For something over a decade a reaction to these traditional teachings has been
developing, brought about in no small degree by the infusion into dental educa-
tion of a more profoundly based group of investigators. After several years of
scientific apprenticeship, evidenced by an increasing number of publications,
t,he “visibility” of a number of these newer dissident’schools of craniologic
theory became apparent in orthodontic circles. Among these the English-
language publications of Scott, 23 KoskP Enlow,* and Mossle probably are most
commonly known to the readers of this article. This incomplete listing is not

From the Department of Anatomy, College of Physicians and Surgeons, and the
Division of Oral Biology, School of Dental and Oral Surgery, Columbia University.
This study was aided, in part, by Grant NB-00965 from the National Institutes of
Health.

474
Volume 56 Capsular matrix 475
Number 5

intended as a denial of the great significance of the work of essentially zoologi-


tally oriented European workers, of whom Delattre and Fenart,, Kummer,12 and
StarckZ4 are excellent examples.
The net result of this vertiable renaissance of craniologic thought has been
to set aside almost all of the older theories and to erect in their place a variety
of newer ideas. At the present time a new conceptual equilibrium has not yet
been attained. Several variant ideas currently are in the intellectual market
place. Behind what appears superficially to be great divergence, there lies
an essential agreement; indeed, it is possible to perceive that almost all the
thoughts of the several workers form a coherent and mutually complementary
whole. It is the purpose of this article to present such an eclectic viewpoint.
This is accomplished, within the encompassing theory of functional cranial
analysis, by clarification of the role played by capsular funct.ional matrices in
cranial morphogenesis.
Functional cranial analysis

The basic principles and supporting data of this analytic methodology have
been published extensively elsewhere.16’ IT, 21 The essence of the method is simply
expressed. The head, not the skull, is a region of the body where a number of
operations are carried out. Every activity is completely executed by an individual
functional cranial component which may be resolved into two parts: a skeletal
unit and a functional matrix. Any function, per se, actually is performed by a
matrix, while the necessary biomechanical role of providing protection and sup-
port to this same matrix is assumed by the skeletal unit. In essence, the head
consists of a number of functional matrices (which are the operational eom-
ponents), all of which are rendered morphologically and spatially secure by
their specifically associated skeletal units.
Since all cranial bones are biomechanically implicated in a multiplicity of
functions, they all consist of several skeletal units. With the demonstration that
the size, shape, location, and structural integrity of every skeletal unit is always
and without exception a secondary, compensatory response to the prior require-
ments of a functional matrix, these same attributes of any “bone” are seen to
be a complex morphologic summation of several operational demands. The fur-
ther demonstration of the relative morphologic independence existing between
even neighboring cranial functional matrices leads directly to several conclu-
sions. First, it is no longer valid to study the “bones” of formal osteology, since
they do not possess a unitary biologic reality. Rather, we must direct our atten-
tion to their component, and equally independent, skeletal units, Second, since
functional matrices are morphologically dominant, cranial growth is viewed cor-
rectly as reflecting the prior growth of matrices and not of the skeletal units.
The final, and obvious, conclusion is that the processes of skeletal unit growth
(osseous deposition and resorption and cartilaginous and connective tissue multi-
plication and hypertrophy) are not sufficient causes to explain cranial growth.
However reluctantly, we must turn our attention away from the all-too-obvious
results of cranial growth and toward its causes; that is, away from the skeletal
units and toward the functional matrices. The recent demonstration that there
is no direct genetic determination of skeletal form or growth (be it osseous or
476 Jloss and Salentij~l

cartilaginous) but, rather, that this cncod~l informational content is displayctl


in their embedding matrices only reinforces this stat cmcnt.!’

Functional matrices

It is a basic postulate of the method of func.tional cranial analysis that, with


the exception of embryonic tissue differentiation, all other aspects of cranial
skeletal tissue growth (osseous, cartilaginous, ant1 fibrous) arc secondary anti
compensatory reflections of t,hc morphogenctically ant1 temporally prior growth
and operation of the cranial functional matrices. The primary growth of all 01’
these matrices, taken together with the secondary xrowth of all of the skeletal
units (the skull), comprises the totality oU head growth. This being so, the fnr-
thcr analytical analysis of the functional rnat,rix (Joricrpt is warranted.
There arc two distinct types of fun&onal ntatriss: t.he periostcal ant1 tlr+.
capsular. Their differentiation is both CWJ- and fun~lamcntal. While the activit:.
of both types of matrix is essential for cranial gVowtli to o(~cur, it is only 1’~
cently that capsular matrices wcrc rccognizcd auf1 thc4r significance clarifietl.lt;
Pcriostcal functional matrices are virtually self-defining. They are best
exemplified by the effect of mnsclcs upon the skclcttal units which subscryc their
contractile function. Whether these mns~los attach their tcndinous portions
directly into thr skeletal tissue or indire&ly by fusion with only the outer, fibrous
layer of the pcriosteum (pcrichondrinm) , they all produce a direct. (a,lthough
secaondary ) , rapitl and complcmcntary response 1)~.their specific skeletal units.
All ptriosteal matrices act homologousl~, usually by mcans of the processes oi
osscnus deposition ant1 rc3orption. ‘I’hcy acat to all (>r cGthc>ror b0t.h the size anti
shape of skelc+al units per se. I N (I z*cq r~rrl s( HW, when wf’ stdy the trclioa
of pcriosteul mairiccs zcc really arc. obserL’i~yj Ihi illtcwctio,l bctwce?L the fwcj
I>urYs of alzl~ fu~ctiond cm7liaZ co)rcj~o)rf71.f.The II~I~I(~I’OLISexamples of actirit).
of periostcal mat.riccs arc rcviclT-cd cstcnsivcl>- cls(~\vh(~.~‘~
Almost all experimental methods for Cc study of skeletal tissue growth are
ca.pablc of demonstrating t.hc activity of only pcriosteal matrices. Vital stain-
ing, intraosseons implants, ant1 histologic csa.minat~ion depict the qualitativc~
and/or the quantitative aspects of the procc~es which effectirely alter o?~Zythe
size and/or shape of the skeletal tissutls. It is not rc>markable that when an in-
vestigator directs attention cxclnsircly towa.rd t,hc observation and mensuration
of a particular t.issue (and in this case a “hard” one)! a further assumption is
made by him that precisely what is observed and measured in growth is, in fact.
identical with bhc sufficient muse of that same growth. Accordingly, it is no
wonder that, t,here arose many thcorios of growth which postulated that thr:
totality of effective cranial bone growth processes could be encompassed by tht>
correlat.ed processes of osseous deposition and resorption. One of the better
known earlier representatives of this school was Brash,j while the more recent,
work so well summarized in Enlo\v‘s* new book is a true deri~~ative. However,
even the strongest proponents of the solitary activity of pcriosteal matrices are
faced with certain observable aspects of cranial growth which present conceptual
inconformities.
Faced with this dilemma, some workers apparently felt forced to accept an
Capsular matrix 477

overly restrictive definition of the term growth; that is, to accept a definition
consistent with their unique reliance upon the sole activity of periosteal ma-
trices. A most recent example is that of Enlow,s who defines osseous growth as
follows: “Over-all enlargement in the size of any bone involves two basic pro-
cesses: growth and remodeling. Growth itself represents the sum of actual in-
creases in size,” while remodeling is defined as a process of “sequential, progres-
sive adjustment that functions to maintain the shape and proportions of the
bone. . . .” The hard and ineluctable fact that Enlow, and others so oriented,
face is that there is an additional phenomenon observable in cranial bone
growth-that of spatial relocation or translation, a motion termed “displace-
ment” by Enlow.
Any careful student of cranial growth, when observing the rapid volumetric
increase of first the neural skull and later the facial skull, was forced to accept
the fact that no possible combination of periosteal apposition and resorption
could account for either the rapidity or the magnitude of such growth. Addi-
tional nonosseous growth processes were then implicated in an attempt to pre-
serve the old conceptual unity. The expansive motion between intramembranous
bones was thought to be due to a primary expansive force generated by the
interstitial growth of the sutural connective tissues. Analogously, the spatial
translations of the facial bones and of the cranial base were thought to be the
result of expansive forces ultimately derived from the primary cellular prolif-
eration and subsequent hypertrophy of the nasal, mandibular condylar, and
spheno-occipital cartilages.
The “pax periostica” was irretrievably destroyed when the experimental and
clinical data of many independent workers explicitly and definit,ively demon-
strated that neither removal of the calvarial or splanchnocranial sutural tissues
nor removal of any of the cranial cartilages in any way inhibited the translative
expansive motions of either the neurocranial or the splanchnocranial bones.131I882o
The resolution of this seeming perplexity was achieved when the full implica-
Cons of the capsular functional matrices were realized.

Capsular matrices

All functional cranial components (skeletal units plus functional matrices)


arise, grow, operate, and are maintained within a series of cranial capsules. The
cranial components comprising the neural region exist within the neurocranial
capsule, while those forming the facial region lie homologously within the oro-
facial ca.psule. Similar statements can be made with respect to the orbital and
otic regions. Both the neurocranial and orofacial capsules as a whole (capsular
tissues per se together with totally embedded functional cranial components)
act to surround and protect their respective capsular matrices.
The neurocranial capsular matrix is formed by the brain, the leptomeninges,
and, most important, by the cerebrospinal fluid. Taken as a functioning whole,
the neurocranial capsular matrix is identical with the volume of this neural
mass, just as the orofacial capsular matrix is identical with the volume of the
functioning spaces of the oronasopharyngeal cavities. Most workers have little
difficulty in the conceptual visualization of the neurocrania.1 capsular matrix.
478 Moss and Mentijn

The “reality” of t,he neural mass is self-evident, in a sense, as indeed is the


“reality” of its cxpsulc. The orofacial capsular matrices, on t.he ot,her hand, re-
quire an operational approach t,o the funct.ioning of the respiratory (and digcs-
live) systems, an approach which appears at first glance to violate the ovcrlp
nai’ve definit,ion of biologic “reality” which some students of the subject adhcr(~
to al: present. The “reality,” and the sine qua non, of any respiratory system is
its patency. Operationally, the form (the size and the shape) as ~11 as thki
spatial location of the orofacial Czl~)SUlC, arid thcrcforc of any of its completcl~~
cmbrdded and included functional cranial components, is dctermincd primarily
b\, the operat~ional volumetric demantls of tlic enclosed patent functioning
spaces. This conclusion is supported intlrpendently l)y the \\.ork of Bosma,“,
who demonstrates the postural atljnstmenis of i he b~;2nc,l~io~nc~ricfuctiomti
cranial components, which he terms the “ailwily maiiltenancr mechanism.”
An abundant body of mutually confirmatory csperimcnt,al ilnd clinical tla t a
has established the morphogcnetic primacy of tlw nt’t>rill capsular matrix in
ncurnl skull growth beyond question- “’ To tlatt, but littlc cxpcriniental data
csist. to support the direct extension of this view to the orofaeial capsular ma-
ticcs; this is due in large part to the intrinsic difficulties of creating adeyuatc:
chronic cervical shun& (For an interesting example, see de Both.“) However,
by- an appropriate analytical nicthotl. it is possible to illustrate clearly ths
homologous morphogcnetia primacy of tlic orofacial functioning spaces in facial
growth. Before such a demonstration, it is well to illustrate first the gcnc~‘al
nature of the responses of all cranial capsnlrs to capsular matrices, while simu-
tancously establishing the sharp differentiation bc~twccn the activities of pcrios-
teal and capsular matrices.
The neurocranial capsule scrvcs as a valuable motlcl for this pnrpostt. The
developing fetal neural mass is surrounded by an essentially mesenchymal
neural capsule whose boundary lagtr externally is the skin and internally the
dura mater. It is within this capsule that the several primary o,ssification centers
of t,he membranous calvarial bones arise (as do the primary chondrific.ation
centers of the supraoccipital bone). There is a gcnetic.ally determined volumetric
growth of the entire neural mass. Vcctorially, the mapitudc of this growtli
force is not influenced by the form of the capsule, although the dircctim of this
primary volumetric expansion is so influenced by several cnl-ironmental (es.
trinsic) factors, among which arc the dcgrcc of flexion of the cranial base and
the mode of att,achmcnt of the several organized dural fiber tracts, which undcr-
lie the major calvarial suture systems.‘4. 22
This vectorially analyzable volumet~ric increase is the primary morphogenetic
fo~w, and it causes a secondary expansion of the surrounding capsule as a
wlwlc. The capsule responds secondarily to this capsular matrix growth by FS-
panding itself, a process brought about by the mitotic activity of the capsular
connective tissues. Jt is self-evident that all portions of the capsule arc similarl)-
aEcctcc1. Any a.nd all embedded functional cranial components are passi~uely
translatetl in space in such an event; that is, both neurocranial functional ma-
trices together with t,heir related skeletal units quite literally are moved in
space. Such spatial relocation takes place without involving the processes of
Volunre 56 Capsular* matrix 479
Number 5

osseous deposition or resorption (or, indeed, of cartilaginous growth either). It


is quite true that the embedded skeletal units are simultaneously altering their
form by active transformative osseous growth processes. However, the observ-
able periosteal apposition and resorption represent the response of these skeletal
units to the constantly altering demands of their respective functional perios-
teal matrices, which are themselves also undergoing passive spatial translation.
An analogy may help to clarify this point.

Spatial translaCon

Let us consider a flat platform, as in a freight elevator car suspended inside


an elevator shaft. Leaning against the sides of this car are a large number of
boards which a passenger may lay on the floor of this car. By laying these
boards down successively, one layer upon another, the passenger most certainly

B. C. D.

T
------_ -----
_------ _----.
Resorb. Depos.
A

TRANSFORMATION TRANSLATION
Fig. 1. Diagrammatic presentation of differences between transformation and translation
as processes involved in cranial bone growth. Transformation is brought about by osseous
deposition and resorption and is a direct response to the primary morphogenetic demands
of specifically related periosteal functional matrices. In this instance, picking up floor boards
in an elevator car (representing a cranial capsule) (A) or laying them down (6) would move
a passenger (representing a skeletal unit) within the car either down or up relative to the
car. Should the car itself move within the shaft (the capsule), the passenger (skeletal unit)
would be translated in space, passively and without the necessity of osseous deposition
or resorption. Obviously, process A or B can occur in situations C or D. Accordingly, the ex-
pansion of cranial capsules in response to the volumetric alterations of enclosed capsular
matrices can be distinguished from transformative changes which, in turn, may be either
complementary or antagonistic to the direction of such translations.
480 Moss and S*lentij,z

will elevate himself vertically within the car. However, the car may bc made to
ascend the shaft simultaneously with the deposition of these boards on its floor.
Let us make the following analogies. Let the car as a whole represent a capsule.
Let the passenger represent, a periostcal matrix, and let the boards rcprcsent il

skeletal unit. Finally, let the elevator sha,ft rcprcscnt a c*apsnlar matrix (that is.
a functioning space). The vertical upward mot,ion of the car as a whole in thtl
shaft, is analogous to the motion of a cranial ca~~sulc as a whole in response tct
a primary altwation in the fnnctSional tlcmantl of a wpsnlar nlntris.
The totally enclosed .func*tional (~rani:11 componc>nt, passcngw plus boartls
(functional matrix plus slielrtal unit), is passively translatctl in space within
this same capsule. 8iirlultaneonsl!-, of wnsecutivcl~-, the passively translated
funct,ional crania1 component may itself alter its o\vn size and shape. The pas-
senger increasing the height of’ the -Iloor 11(1stantls on 1)~ laying tlown the boards
is an example of such a tlirwl. tl,aiisfoi,niatirc, prowss. JYlwn the passeng~~r I~J-s
down boards while the clcvator car is asccndin~, t Iris elcsrrly is ii case in which
the direction of lwth the c~apsular ant1 pwiosttwl gro\z.tli prowsscs is identical.
However, it. is cqaally possibl(~ for this silnlc l~ssengc~r t,o pick up boar*~ls al-
ready laid down while the c>ltlvator ear is ascrntlinp. In this case, although 11~~
is descending within the ('ill' (tlic> c~~y)snloJ! hc is ncvc~rthrless moving upward
within the capsule.
The necessity for sharp dist,inction bctwten these two t,k-pcs of motion (01
of growth proce~sw) now is swn to be very critical (Fig. 1). It is as if craniol-
ogists for many years have obscrvc~l only the changes within the elevator cam
and never compr&cndetl fully the signifiranw of lh(l motion of the car withill
111~shaft,. Nor is this too snrprising. While all rnw acknomlrtlg~e the daily rota-
t,ion of the earth upon its own axis, ant1 less distinctly “fwl” the effect of thus
motion of their planet, about) our sun, still less arc they consciously an-are ol
the motion of 0111’entire solar systcwi through OIII* galaxy, and almost no one
considers the reality of the galactic motion as a whole ilhOUt cithcr its own asis
or through the immensitg of int,crgalactic spaw. Y(>t all of these motions a~‘(’
equally “t,ruc” anct “real.”

The Orofacial capsule

The orofacial capsule originatrs by the process of enclosure. In the embryonic


st,age of cephalogenesis we distinguish a phase prior to formation of t,hc several
facial processes, a phase in which there is, in fact, no oronasal functioning space
y-et in being. At about the twenty-fifth clay after fertilization (t,wenty somites)
there is “a deep cleft between the heart and the overhanging forebrain. The
maxillary process springs from t.he proximal border of the (first) arch. . . Wit11
the mandibular process it completes the lateral boundary of the oral pit.“” The
anterior end of the gut is effectively closed by the buccopharyngeal membrane,
which at this sta.gc is directly on the surfare of the embryo. At about the
twenty-first to the twenty-second day the swcral facial processes begin to
develop ; this development, while presumably guided by genetic information
encoded in the ectoderm, effectively is one of mesodermal proliferation. In effect,
the first arch swellings produced by the bilateral mesodermal proliferation quite
Volume 56 Capsuhr matrix 4%1
Number 5

Fig. 2. The earliest stages of enclosure of the potential common oronasal cavity in a
twenty-somite embryo. At left, in the frontal view, the now perforating buccopharyngeal
membrane is seen at the base of the oral pit which lies between the forebrain above
and the heart below. In the central figure the mandibular arch, to the right of the pit, is
seen to have just begun its anteriorward and mesial movement, caused by mesodermal
multiplication. The dorsally derived maxillary process is not yet in evidence. At the right,
in sagittal section, the buccopharyngeal membrane is seen still separating the pharynx
from the oral pit. Subsequent enclosure of this pit by the developing mandibular and
maxillary processes will form the oronasal cavity.

Fig. 3. Growth subsequent to that shown in Fig. 2 is illustrated. On the left, the maxillary
process of the first arch is now beginning to fill out as a result of proliferation of the
underlying mesoderm. The continued growth of the maxillary process and its subsequent
participation in the formation of the external nares are shown at the right. This, together
with mandibular process growth, completes the genetically determined process of en-
closure of the oronasal cavity.
482 Moss and Salentij?z

literally enclose, and thus form, the primordial oronasal cavity. The subsequent
terminal fusions of the maxillary and nasal processes, well described clsewherc\,
are completed at, about the thirty-fifth to the thirty-seventh day. The buc-
copharyngeal membrane rupturc>s on approximately the twenty-sisth day, join-
ing the ectodermally lined oronasal functioning space to the endodcrmaIly linclcI
primitive phar;vnx and thus creating the totality of the oronasopharyngeal Cum-
tioning space. Although this functioning space comes into being under gcnclic:
i,intrinsic) control, it,s subsequent growth and maintenance in being arc func:-
I ionally (environment,ally, extrinsically) drtermined (Figs. 2 and 3) .
The onset of ossification of the skeletal tissues which protect, and support t,his
l’lmctioning spaccl dots not begin until the sixth week (15 mm. crown-rump
length), while the re?fl~srs of month opening and swallowing are started at about
sl/ff wecks.l”
awhile t,hc primitive palate (extending posteriorly to the region of the futurr
incisive foramen) is formed when the maxillary and nasal processes join, mosi
of the primitive oronasal functioning space remains a common volume. It is OlllJ-
when the bilateral palatal processes form (at about. the fortieth day), eleratc,
and fuse (fort,y-seventh to fift,ieth clay) that, the functional differentiation b(l-
twccn the oral and nasal functioning spaces occurs.

Growth of orofacial functional cranial components

The orofacial capsular matrix (the oronasopharyngeal functioning space j


is surrounded by the orofscial capsule. The limiting layers of this cavity are skin
(generally) externally and mucous membranes internally. Totally embedded
within this CikpSUle lit a, number of orofacial functional cranial components,
both the indivitlual functional periosteal matrices and their respective skeletal
units. (For example, the tcmporalis muscle/coronoid proces8scomprises one such
cranial component ; the massetcr and medial pterygoid muscles/angular process
forms another unit.) The growth of all orofacial skeletal units is a combination
of the two types of growth process discussed above-periosteal and capsular,
transformative and translativc, changes in size and shape, and changes in spatial
position.
The establishment of the morphogenetic primacy of orofacial functioning
spaces as causing the branslation of all of the skeletal units embedded within the
orofacial capsule requires the demonstration that the volumetric expansion of
these spaces is not the result. of prior skeletal tissue growth. Specifically, it is
held currently by someZ3 that the interstitial expansive growth of the nasal
capsular cartilagr, and particularly the scptal cartilage, is the primary cause of
the translative growth of the middle face, while the expansive growth of the
mandibular condylar cartilages is held homologously to be the primary cause
of the translative growth of t.hc lower face. In other words, the nasal cavity
volume expands because of, and secondary to, the growth of the nasal septal
cartilage, while t,he oral cavity volume expands because of, and secondary to,
the growth of the condylar cartilages.
The available experimental and clinical data deny these older concepts and
furnish strong support for our view of the morphogenetic primacy of the
Volume 56 Capsular matrix 483
Number 5

volumetric expansion of the orofacial functioning spaces. Complete bilateral


extirpation of the mandibular condylar cartilage does not inhibit the translative
growth of the mandible.20~ 21 This statement is supported completely by a wide
variety of experimental data.ll Similar statements can be made concerning the
nasal septal cartilage.1s Indeed, the nasal septum has been shown to play an
important biomechanical supportive role, rather than acting as a primary source
of gr0wth.l
A semantic distinction made by Koski I1 helps to clarify the matter. By
carefully differentiating between a skeletal tissue growth center and a skeletal
tissue growth locus, we can distinguish between the regions which primarily
cause translative growth from those which secondarily respond to this same
spatial relocation. It is our contention that there are no growth centers in
skeletal tissues at all but, rather, that all such regions as the nasal septal cartilage
and the mandibular condylar cartilages are loci at which secondary and com-
pensatory periosteal growth changes occur in the size and shape of these skeletal
units, compensatory to both the spatial translations produced by the primary
expansion of the orofacial capsular matrices as well as to certain alterations in
the demands of periosteal matrices. Before demonstrating these differences quan-
titatively, let us examine the processes involved more closely.
We will use lower jaw growth as our present example. Ongoing studies in
both rodents and man, to be published elsewhere, enable us to classify this
growth as essentially logarithmatic in nature and to demonstrate certain highly
significant quantitative parameters. For the moment, we wish to state only that
the human oronasopharyngeal functioning space alters in size alone, not in
shape, after the beginning of the third month of pregnancy. This morphogeneti-
tally primary volumetric increase causes a compensatory increase in the size of
the orofacial capsule. Growth of the enveloping capsule is produced by mitosis
of both its epithelial and mesenchymal cellular elements and the consequent in-
crease in intercellular materials, which results in an expa.nsion of the capsule as
a whole. Within the capsule are situated a number of mandibular functional
cranial components, periosteal matrices together with their respective skeletal
units.
Because the mandibular cranial components arise and exist completely
embedded within this capsule, they all necessarily are passively and secondarily
translated in space to a new position as the capsule expands. Such passive trans-
lations affect both the periosteal matrices and their skeletal units. Accordingly,
simultaneously with such passive, indirect translations of the mandibular func-
tional cranial components as a whole, the individual periosteal matrices may also
alter their functional demands (muscle growth). These matrix changes will then
cause direct growth changes in the size and/or shape of their several skeletal
units. It remains only for us to see clearly that these later transformative
growth changes are not the cause of, or even the direct result of, the passive
translations of these same functional components.
Specifically, the change in size and/or shape of the mandibular eondylar
cartilage is not evidence that a primary growth center exists here. Rather, as
the mandibular complex of skeletal units as a whole is passively moved in the
484 Moss und Xulentijrt

three planes of space within the expanding orofacial capsule, the condylar hea<
is passively carried away from its superior articulating surface. The obscrveti
and undoubted growth within these cartilages is a compensat,ion for wch potcn-
tial joint disa.rticulation and is brought about, in part, by 117~altered fnnctionai
demands of the lateral pteq-goid muscle. Similarly, thanges in the size and, (11.
shape of other mandibular skeletal units as indicated by sclectivc arras of rcsoq-
tion and deposition of skeletal (7~suall~ OSSCOIIS) tissncl ;~YPobsc~vcttl.
It is possible to demonstrate and differentiate t,he morphogrnetic effect oi
both capsular and periosteal matrices in clinical material. The technique is
simple.21 A longitudinal series of c~e~)halomc~tric;7Ll?- orientctl roentgenogram is
used and tracings are prepared iin this cast? in norma latcl*alis). For 0711
present purposes, it is suficient to trace the CClY’l)lXl stIrface oi’ the cranial base‘
and the external s77rfacc of the osseo77smandibular complcs (the mancliblc c~f
traditional osteology). On these tracings we include also the position of tllc*
mental and mandibular foramina as well as that of the inferior al\-eolar canz 1.
marking as they do that hasal skeletal unit response to the matrix formed I)>-
the inferior alveolar ncurorascular triad (Fig. 4). Taking the first and last o t
the srrics of t,racings, we ca,n nnlv l~~clucc~ a series of composites based on the
following assumptions : ( 1) that the neural mass overlying the anterior cercbr2 i
fossa has completed its growth by the end of tht: thirtl year so that the cel~rbral
surface of the anterior cranial base is constant in size, shape, and positiotl :
(2) t,hat the position of the Itlt~ntal fornmcn does not alter with timc’.‘i
When the two tracings arc supcrimposctl on the nntcrior cranial bast>, I\(.
observe the total growth changes of the mandibular complex dnring this pcrioci
(Fig. 4). This totality represents the response t,o both capsnlar and prriostcal
matrices. We term this a demonstration of in.fcrosseous growth, that is, the tota 1
growth relative to the fixed anterior cranial base. We map now prepare a seeontl
c*omposite t.racing, orienting both mandibles so that the anterior cranial hnsc
outlines are perfectly parallel and rcgistcring bot,li m:intlibalar 077tlines on tlic
mental foramena. We now obstrve the changes in size anal, ‘or shape of tlrtz
sctveral mandibular skeletal 7mits which occur irldr,l)c,ldcnt7!l of the changes ill
spatial position of these same nnits with timo. This is tcrmcd i~f~~~os,scousgrowth
i Figs. 4 and 5). This method has been applied prc‘viously to a preliminaq, &u(l).
of the maxillarv Y growth.‘” Finally! a thirtl composite is made in which wo t:lkts
both of the previous composite tracGngs ant1 superimpose them 071 thcl outlincss
of the old& (larger) mandibles.
We observe now two distinct positions of the earliest (smaller) mandittulal
outline. The distance between the two idcntical earlier tracings preciselg ant1
exactly represents the amount of passive, translative growth that would occur
if only capsular growth occurred. That is, if periosteal matrices did not alter
their functional demands, the expansion of the orofacial fnnctioning space (thr
capsular matrix) would have carried thcsc unchanging mandibular skeletal 7mits
to this new position in space passively, without invol\-ing the processes of osseo77s
deposition and resorption. ITowever, ossc~oustransformation did occur during
this period of passive translation. The net effect of t,hese changes in the size
and shape of skeletal units in response to the periostcal matrices is indicated b>
Volume 56 Capsular matrix 485
Number 5

-7yrs, 6
-- 15 yrs.

Fig. 4. TWO longitudinal tracings of a patient, registered on a common outline of the


anterior crania) base. The mandibular outline, the mental foramina, and the course of
the inferior alveolar canal are shown. This figure shows interosseous growth, a combina-
tion of both transformation and translation. This is the totality of mandibular growth rela-
tive to a biologically fixed neurocranial area.

Fig. 5. The tracings in Fig. 4 are now registered on the mental foramen, with the plane of
the anterior cranial base kept absolutely parallel. This now demonstrates intraosseous
growth. In effect, this demonstrates the amount and direction of transformation (osseous
deposition and resorption and cartilaginous proliferation) during this period of time.
486 Moss and Xalentij~~~ Am. J. Orthodontics
November 1969

Fig. 6. An overlay of the tracings of Fig. 4 and 5 produces this picture, which shows both
transformation and translation. In effect, the two positions of the younger mandible show
what would occur if the mandible were only translated (passively moved within the ex-
panding orofacial capsule), without transformation occurring simultaneously. The magnitude
and direction of this translation are shown by the arrows. However, transformation occurs
also and is shown as areas of resorption [black) and deposition (plus signs). This figure
is an approximate mean and clearly shows that the downward and forward motion of the
mandible primarily is passive translation, whereas active transformation produces minor
changes anteriorly and inferiorly while being entirely responsible for the posterior and
upward compensatory growth of the ramus.

the differences between the lowermost of the earlier mandibular outlines and
the outline of the older mandible (Fig. 6). As is seen, some of these changes are
additive and some are subtractive. In general, they account for the posterior and
upward growth of the ramal skeletal units, as well as for the slight adjustive
changes in the anterior and lower borders of the corpus. But the sum of all of
these direct periosteal changes, involving osseous and cartilaginous growth, does
not and cannot account for the translative growth. Indeed, it seems that passive
translation comprises by far the major portion of the totality of mandibular
growth in a downward and forward direction.
The vectors of mandibular growth have a variable range of both magnitude
and direction. Figs. 7, 8, and 9 illustrate a vertical growth case as well as two
extremes of the normal range of such vectorial growth as observed in a random
examination of a series of participants in a longitudinal growth study at this
dental school. The display of such a morphogenetic spectrum is to be expected,
rendering it impossible for us to present the clinician with any single idealized
“mean” figure which might in any way serve to establish an overly rigid concept
of mandibular growth, despite the repeated attempts of other workers to do so.
A final clinical example of the morphogenetic role of the orofacial function-
Volume 56 Capsular matrix 487
Number 5

- 7 yrs
- I5 yrs.

Fig. 7. In this and the next two figures the technique is that illustrated in Figs. 4, 5, and 6.
These three figures illustrate the range of growth observed. In this case a vertical direction
of the growth vector is shown.

- 6 yrs, I I mo.
-- 15yrs., 5 mo.

Fig. 8. A patient showing a marked horizontal vector of translative growth.


488 Moss and Xalentijx Am. J. 07euxkwLtics
November 1969

- Syrs, 6mo

-- 14yrs, 5mo

Fig. 9. Translative growth in this patient has a posterior and downward vector. Note in
this figure, as in Fig. 8, that resorption of the anterior border of the mandible is indepen-
dent of the direction of translative growth. [See also Fig. 1.)

ing spaces is shown in Fig. 10. The patient, a lo-year-old white girl, underwent
a two-stage condylectomy. The first date (July 15, 1968) was one week after
unilateral condylectomy; the second date (July 31, 1968) was 3 weeks after
the same operation; the last date (Nov. 14, 1968) was 5 weeks after the second
condylectomy. The magnitude of the spatial alterations depicted here are far in
excess of the transformations to be expected by a combination of osseous deposi-
tion and resorption ; rather, these represent translative growth. In cases such
as this the functional ankylosis of the tcmporomandibular articulation literally
acts as a mechanical blockade. In effect, the normal morphologic expression of
the increases in the volume of the functioning spaces was prevented. The
magnitude of these growth force vectors is not altered, but their direction is so
altered. Quite literally, in such ankyloses, the fixation of the condylar process
prevents passive translation from taking place. Following successful surgical
intervention, however, the orofacial capsule snd its embedded mandibular
skeletal units are permitted to respond to the still present divecthon of the oro-
facial matrix growth vectors. The short-term translative changes demonstrated
in this case are complementary to the long-term growth changes discussed
previously.
The potential significance of the concept of the capsular matrix in functional
craniology is great. Ongoing quantitative studies in our laboratory will soon
permit us to establish some of the parameters of totality of facial growth. The
absolute and relative amounts of active periosteal and passive capsular growth,
Volume 56
Number 5 Capsular matrix 489

R L

I I/ 14/ 68
---v--- 7/ 31/68
---.- 7/ 16/ 68
Fig. 10. Three longitudinal tracings superimposed on the anterior cranial base of a patient
with a two-stage bilateral condylectomy, with right (R) and left (1) sides shown individually.
The magnitude of spatial change here is far too great to be accounted for by active trans-
formation (osseous deposition and resorption) but is accounted for by passive translation.
In this case the deformed oral volume capsular matrix is now permitted to re-establish its
normal contours, and thus it rapidly translates the orofacial capsule together with the
embedded mandible.

while varying from age to age and from person to person, do in fact lend them-
selves satisfactorily to mathematical and graphic expression. Future publications
will present these data. For the moment, we are content to present the concept
that the growth of the orofacial functioning spaces plays a major role in facial
growth.
Summary

The morphogenetic role of two types of functional matrix-per&teal and


capsular-in craniofacial growth is examined. The term growth is defined
inclusively to include the changes in size and shape as well as changes in spatial
position in time. The periosteal matrices are responsible for transformative
growth, the changes in size and shape. The capsular matrices are responsible for
translative growth, the changes in position. The role of oronasopharyngeal func-
Am. J. Orthodontics
490 Moss and Salentij,~b November 19F9

tioning spaces as competent, primary, morphogenetic agencies is detailed. T11e


volumetric expansion of these capsular functional matrices is capable of accurate
quantitative description.

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scanclinav. 6: 239-250, 1966.
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3X-370, 1959.
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202, 1969.
18. Moss, M. L., and Bromberg, B.: The passive role of nasal septal cartilage in mid-facial
growth, Plast. & Reconstruct. Surg. 41: 536-542, 1968.
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bone. I. Basal bone, Angle Orthodontist 37: 151-164, 1967.
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Angle Orthodontist 38: 95.103, 1968.
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AXE. J. ORTHODONTICS 55: 566-577, 1969.
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Anthropol. 18: 281-292, 1960.
23. Scott, J. H.: Dento-facial development and growth, Oxford, 1967, Pergamon Press.
24. Starck, D.: Die endokraniale Morphologieder ratiten, Morphol. Jahrb. 96: 14-72, 1955.

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