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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
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 First, it is no longer valid to study the “bones” of formal osteology, since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
Functional matrices
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
Spatial translaCon
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.”
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.
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. 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.
- 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
REFERENCES
1. Badoux, D. M.: Framed structure in the mammalian skull, Acta morph. neerlands-
scanclinav. 6: 239-250, 1966.
2. Bosma, J. F.: Oral and pharyngeal development and function, J. D. Res. 42: 375-380,
1963.
3. Bosma, J. F.: Maturation of functions of the oral and pharyngeal region, Am. .I.
Orthodontics 49: 94-104, 1963.
4. de Both, N. J.: Transplantation of Axoloth heads, Science 162: 460-461, 1968.
5. Brash, 5. C.: Some problems on the growth and developmental mechanics of bone,,
Edinburgh M. J. 41: 305-319, 363-387, 1934.
G. Davis, C. L.: Description of a human embryo having twenty paired somites, Carnegie Inst.
Contrib. Embryol. 15: l-51, 1923.
7. Delattre, A., and Fenart, F.: Analyse morphologique du splanchnocrane chez les primates
et ses rapports anec le prognathisme, Mammalia 20: 169-325, 1956.
8. Enlow, D. H.: The human fact, New York, 1968, Hoeber Medical Division, Harper & Roiv.
9. Griineberg, H.: The pathology of skeletal development, New York, 1963, John Wiley &
Sons, Inc.
10. Humphrey, T.: The development of mouth opening and related reflexes involving the oral
area of human fetuses, Alabama J. M. SC. 5: 126-157, 1968.
11. &ski, K.: Cranial growth centers: Facts or fallacies, Anr. J. ORTIIODONTICS 54: 566-583,
1968.
12. Kummer, B. : Untersuchungen iibcr die ontogenetische Entwicklung des mensehlichen
Schidelbasiswinkels, Ztschr. Morphol. u. Anthropol. 43: 331-360, 1952.
13. Moss, M. L.: The growth of the calvaria in the rat, Am. J. Anat. 94: 333-362, 1954.
14. Moss, M. L.: The pathogenesis of premature cranial synostosis in man, Acta anat. 37:
3X-370, 1959.
15. Moss, M. L.: A functional analysis of human mandibular growth, J. Pros. Dent. 10:
1149-1160, 1960.
16. Moss, M. L.: The primacy of functional matrices in orofacial growth, D. Practitioner
19: 65-73, 1968.
17. Moss, M. L.: A theoretical analysis of the functional matrix, Acta biotheoret. 18: 193.
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.
19. Moss, M. L., and Greenberg, S. N.: Functional cranial analysis of the human maxillary
bone. I. Basal bone, Angle Orthodontist 37: 151-164, 1967.
20. Moss, M. L., and Rankow, R.: The role of the functional matrix in mandibular growth,
Angle Orthodontist 38: 95.103, 1968.
21. Moss, M. L., and Salentijn, L.: The primary role of functional matrices in facial growt.h,
AXE. J. ORTHODONTICS 55: 566-577, 1969.
22. Moss, M. L., and Young, R. W.: A functional approach to craniology, Am. J. Phys.
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.