THEORIES OF GROWTH & DEVELOPMENT

Dr.Thameem Ahmed.R
1 st yr PG
CONTENTS
Introduction
Growth fields
Growth sites
Growth centers
Enlow V principle
The Remodelling theory (Brash 1930s)
The Genetic concept (Brodie 1940s)
The Sutural hypothesis ( Sicher 1941)
The Cartilaginous theory (Scott 1950s)
 The Functional Matrix theory (Moss 1962)
 Neurotrophism
 Functional Matrix revisited
 The Cybernetics theory
 Conclusion
 References
INTRODUCTION
Growth and development are two integral
process which defines the existence of life.

Growth of an organism is the interplay

between its genetic constitution and
environment in which it thrives.
Assessment of growth revels about the general health of
the individual and can be used for growth modification
treatments.

Growth is a complex process and is not supported by a

single theory but is based to a large extent on evolving
concepts concerning the biological mechanisms of
craniofacial development.
Definition of Growth
 “Growth refers to increase in size” - Todd

“Growth usually refers to an increase in size

and number” – Proffit

 “Self multiplication of living substance”-
J.S.Huxley.
Change in any morphological parameter which is
measurable”- Moss.

6
Definition of Development
Development is a progress towards
maturity” – Todd

“Development connotes a maturational

process involving progressive
differentiation at the cellular and tissue
levels” - Enlow

7
GROWTH FIELDS

 Bone growth is controlled

by growth fields.

 Distributed in a mosaic like

pattern across the surface of a
given bone.

 They have pacemaking

function.

 They are either resorptive or

depository activity.
About one half of the
bone is periosteal and
the other half
endosteal.

If endosteal surface is
resorptive then
periosteal surface
would be depository.
GROWTH SITES
 Growth fields having
special role in the
growth of the particular
bone ( grows fast) are
called growth sites
 e.g. mandibular condyle,
maxillary tuberosity,
synchondrosis of the
basicranium, sutures
and the alveolar
process.
Baume proposed
the term growth site
for “regions of
periosteal or sutural
bone formation and
modeling resorption
adaptive to
environmental
influences.”
Koski, K. : Cranial growth centres: Facts or fallices?
, AJO-DO : Aug 1968: 566-583
 GROWTH CENTERS

 Special areas which

are believed to
control the overall
growth of the bone
e.g.mandibular
condyle.
 Force, energy or
motor for a bone
resides primarily
within its growth
centre.
The term growth center is widely used
inconnection with skeletal growth
phenomena.

Baume proposed that the term skeletal

growth center be used to describe “places
of endochondral ossification with tissue
separation forces.”
Koski, K. : Cranial growth centers: Facts or fallices? ,
AJO-DO : Aug 1968; 566-583
ENLOW’S V PRINCIPLE

 Most useful and basic

concept in facial growth as
many facial and cranial
bones have a Vshaped
configuration.

 Bone deposition(+)
occurs on the inner side
and resorption (-) occurs
on the outer surface
EXAMPLE WITH V
ORIENTED VERTICALLY

 Bone deposition on
lingual side of coronoid
process , growth proceeds
and this part of the ramus
increases in vertical
dimension.
V ORIENTED
HORIZONTALLY

 Same deposits of bone

also bring about a posterior

direction of growth
movement.
 This produces a
backward movement of
coronoid processes even
though deposit is on the
lingual side
 Same deposits carry base of bone in medial
direction .
 So, the wider part undergoes relocation into a
more narrow part as the whole v moves towards
the wide part .
BONE REMODELLING THEORY
BY BRASH (1930)
This theory states that bone grows only by
interstitial growth.

The fundamental tenets of this theory are:

Bone grows only by apposition at the surface.
Growth of jaws takes place by deposition of bone
atthe posterior surfaces of the maxilla and
mandible.

This is described as Hunterian growth.

Carlson ,D.S.:Theories of craniofacial growrth in postgenomic era .

Seminorthod 2005;11:172-183
Calvarium grows through bone deposition on the
ectocranial suface of the cranial vault and resorption
of bone on the endocranial surface.


THE GENETIC THEORY( 1941)

Genetic theory was given by Stewart and Brodie.

The genetic theory simply stated that

genes determine and control the whole
process of craniofacial growth.

But the mechanism of action by the genetic

unit and the mechanism by which the traits
are transmitted were not understood.
Genetic concept suggests that the genes
supply all the information in growth and
development.

 This originated with classical Mendelian

genetics.

Later with the blending of data from

vertebral paleontology created the neo-
Darwinian synthesis which is currently
accepted paradigm of phylogenetic
regulation.
The field of genetics consists of two
principle areas :

“Transmission genetics” is characterized

by statistical approach and involved only in
explaining possible method of transmission.

It is based on Mendelian laws and did not

explain about genes or its characteristics.
For

Certain types of malocclusion run in families like

HAPSBURG JAW – Prognathic mandible common in
Austrian royal families.
Stockard experiments-crossbred dogs showed that
dramatic malocclusions occurred in crossbred dogs.
Against

 Hawaiian melting pot : Hawaii had homogenous

Polynesian population. Large scale migrations by European,
Chinese & Japanese resulting in various tooth size, jaw size
and jaw proportions which were all different for the
Polynesian, Oriental & Europeans led to the Hawaiian
melting pot but incidence of malocclusion was same.
Twin studies: - Twins have different malocclusions.

Stockard experiments indicated that dramatic

malocclusions did occur in his crossbred dogs, more from jaw
discrepancies than from tooth- size imbalance.
THE SUTURAL HYPOTHESIS

Given by Sicher and Weinmann in 1947 .

According to this theory, sutures ,cartilages and

periosteum are responsible for facial growth and
were assumed to be under intrinsic genetic
control.

Sicher came to the conclusion that

sutures were causing most of the growth
based on the studies using vital dyes.
Essence of the Theory:
According to Sicher, the sutures are the
primary determinants of craniofacial growth.

The craniofacial skeleton enlarges due to

expansible forces exerted by the sutures as
they separate.
Theory:

He stated that all bone forming elements like

sutures ,cartilages and periosteum are growth
centers like the epiphysis of the long bone.

Sicher called this theory as the sutural

dominance theory because he believed that the
primary event in sutural growth is proliferation of
the connective tissue between the two bones.
Proliferation of the sutural connective
tissue creates the space for appositional
bone growth between the borders of two
bones.

 Increase in the size of the cranial vault

takes place via primary growth of the bone at
the sutures, which forces the bones of the
vault away from each other.
Growth of the midface takes place via intrinsically
determined sutural expansion of the
circummaxillary suture system, which forces the
midface downward and forward.

Mandibular growth takes place via intrinsically

determined growth of the cartilage of the
mandibular condyle ,which pushes the mandible
downward and forward.
Koski (1968) stated there are two different
views regarding the structure of sutures.

Thefirst school of thought (Sicher and

Weinmann ) considers sutures as a
three layered structure.

Koski, K. : Cranial growth centres: Facts or fallices? , AJO-DO :

Aug 1968: 566-583
It stated that the connective tissue between
the two bones plays the same role as the
cartilage at the bases of the skull and like
epiphysis of the long bone.

There is spreading of sutures due to

proliferation of middle layer of the sutural
tissue.

According to this concept tissue separating

force exists in the suture itself.
The second school of thought (Pritchard
,Scott and Girgis,1956) says the suture as a five layer
structure .

 Each bone at the suture has its own two layer of

periosteum on both sides and the intervening fifth
layer between these periosteal layers.

This layer plays a role in adjustment between the

bones during the growth .

while the active proliferating role is played by the

cambial layers of the periosteum of each bone.
It is very clear now from the histological evidenced
that the sutural structure is not identical to that of
the epiphyseal growth plate.

Sicher also perceived the mandible as a long bone

and the mandibular condylar cartilage as comparable
to epiphyseal plate of the long bone.
EVIDENCES AGAINST SUTURAL
THEORY

Trabecular pattern in the bones at the suture change with

age ,indicating the changes in the direction of growth it
cannot be accepted that suture will have the necessary
information for altering growth.

Extirpation of facial sutures has no appreciation

effects on the dimensional growth of the skeleton
(Sarnat 1963)
Shape of sutures have been found to depend on the
functional stimulus.( Moss &Salentejin, 1969)

Closure of sutures to be extrinsically determined.

( Moss ML, 1954)

Sutural growth can be halted by mechanical forces

like clips placed across the sutures.
The parallelism of circummaxillary suture so as
to effect a forward and downward growth of maxilla
is only superficial .

Growth at zygomaticomaxillary suture occurs

predominantly in lateral direction .

The direction of growth of maxilla ranges from 0

to 82 degree in relation to SN plane .

It is practically impossible for the suture running

in same direction to push the maxilla parallel to the
reference plane. (Bjork 1966)

Bjork: Acta odont.scandinav.1966; 24:109-127

Conclusion:
Present evidences indicates suture as
adaptive growth sites.
Sutural tissues have no tissue separating
forces and they are not comparable to
growth centers.
NASAL SEPTUM THEORY/CARTILAGENOUS
THEORY/NASOCAPSULAR THEORY (1950)

James H Scott , an Irish anatomist in 1950

proposed the nasal septum theory as the
single and unified theory of craniofacial
growth.
Essence of theory:

According to this sutures play little role

or no direct role in the growth of the
craniofacial skeleton .

Sutures are considered as merely

passive secondary and compensatory
sites of bone formation and growth
After recognizing the importance of
cartilaginous parts of the head , nasal
capsule ,mandible and cranial base in prenatal
growth.

Scott felt that this cartilaginous development

was under genetic control and was of the
opinion that they continued to dominate
postnatal facial growth also.
Scott concluded that nasal septum is mostly active
and vital for craniofacial growth both prenatally and
postnatally.

The anterioinferior growth of the nasal septal

cartilage which is buttressed against the cranial base
“pushes” the midface downward and forward.

The cranial base synchondroses causes the growth

of the cranial base and Scott compared the condylar
cartilage to the cranial base cartilage
Numerous experimental studies were conducted to
address the validity of Scott’s hypothesis .

This theory is based on the fact that cartilage is a

pressure adapted tissue and expansion of cartilage
provides the force to displace downward and forwards.

According to Scott ,bone separation must precede

before the adaptive sutural bone growth occurs .

The bone separation is because of growth

of organs like brain , eyeball or cartilage.
Scott is of the opinion that there are two suture
system:

 Posterior suture system lies behind the

maxilla and separates it from palatine ,lateral
mass of ethmoid , lacrimal , zygomatic and
vomer bones.

 Anterior suture system separates premaxilla

nasal and vomer bone.
Scott said that the nasal cartilage is an
extension of the cranial base cartilage and
as it grows further, it separates the facial
bones from one another and also from the
cranial portion .

 Thus it allows bone growth to take place at

the sutures (frontomaxillary , frontonasal ,
frontozygomatic and frontozygomaticomaxillary)
by surface deposition.
Evidences supporting the theory:

Experimental research on rats by

Ohyama(1969)
removal of septal cartilage produces deficient
growth of snout.

Also supported by research of Burdi , Petrovic

Baume, Latham (1965,1967-1968,1968,1972)
respectively .
EXPERIMENT BY BURDI , PETROVIC ,
BAUME,
LATHAM (1965,1967-1968,1968,1972)
RESPECTIVELY
The importance of the septal growth was also
seen in impulse to maxillary growth in cleft
palate cases.

Failures of the underdeveloped maxillary

segment to unite with nasal septum in complete
unilateral clefts deprives it of the growth
impulse or energy.

The normal contra lateral side on the other

hand, attained normal growth.
Sarnat in (1988)from experiments on rabbit snout
concluded that deformity of snout after
resection of nasal septum was the result of
lack of growth.

 Latham (1974) described the role of

septomaxillary ligament passing from
anterioinferior border of nasal septum to anterior
nasal spine and inter maxillary
suture in premaxillary region.

He stated that the traction through the

ligament will exert downward and forward
growth of maxilla.
Koski(1968) after histological study of nasal
septal cartilage found that there is
endochondral ossification taking place at
septoethmoidal junction.

Hunter and Enlow(1968) –in their growth

equivalent theory –emphasizes on relatively
lesser response of the endochondral cranial
base as opposed to immediate response of the
intramembranous cranial vault to external
influences
Evidence against the theory:

Moss and Bloonberg(1968), Brigit

Thilander(1970) found only slight deformity
after extirpation of septal cartilage

They concluded that septal cartilage provides

only mechanical support for the nasal bones
and is not a primary growth center.
Two studies were carried out by Gilhus-Moe and
Lund in Scandinavia in 1960’s showed that
There are excellent chances that condylar
process would regenerate to approx. its
original size after trauma

In a few there was even a overgrowth of condyle.

In a few children there is a reduction in growth

after injury maybe due to the trauma to the soft
tissues / scarring

Therefore Scott’s hypothesis does not hold true

completely.
CONCLUSION:
At present ,nasal septum theory is still accepted
as a reasonable explanation for craniofacial
groawth.

 Nasal septum may be important for

anterioposterior growth of face because of
endochondral growth process occurring at its
posterior border.

It is not considered to be an active

contributor for vertical development of face.
HUNTER &ENLOW’S GROWTH EQUIVALENT

The Hunter-Enlow growth equivalents concepts is

an important principle covering the development of
the facial skeleton.

As the individual components of the skull develop

in different directions ,they must interact directly in
order to compensate for the various growth
activities.

This is achieved by growth equivalents

which act in opposing directions.
These growth equivalents coordinate the different
movements of the cranial base ,the nasomaxillary
complex and mandible , which are due to
development ,and thus determine the adaptive
changes in relation to individual parts of the skull.

For example, elongation of the anterior cranial base is

related with enlargement of the nasomaxillary
complex.
THE FUNCTIONAL MATRIX THEORY
INTRODUCTION

The concept of this theory was

introduced first by Vander Klaww(1948- 52).

Melvin L. Moss developed the form and

function concept into the functional
matrix hypothesis.

Introduced in 1962.
ESSENCE OF THE THEORY

The functional matrix hypothesis claims that the

origin , form , position, growth and maintenance of
all skeletal tissues and organ are always secondary
,compensatory and necessary responses to
chronologically and morphologically prior events or
processes that occur in specifically related
nonskeletal tissues ,organs or functioning spaces
(functional matrices).
The hypothesis as shown that change in size, shape,
and location (growth) of all craniofacial skeletal
entities are epigenetically( causally related series of
processes in external and internal environment)
regulated.

The epigenetic hypothesis suggests that the post

fertilization genome does not contain sufficient
information ,such as a blueprint, to regulate all
subsequent development.
Proponents of the functional matrix states
that the expansion of the soft tissue matrix is
primary and the bone growth is purely
secondary and compensatory event.

Translation of the various bones of the face is

due to volumetric expansion of the
encapsulated spaces or tissues.
PERIOSTEAL MATRIX

Relates the matrix to those tissues that influence

the bone directly through the periosteum.

Examples of periosteal matrices includes:

Muscles, Blood vessels and nerves lying in grooves or
entering or exiting through foramina of Teeth.
Lack of contraction leads to atrophy of the bone.

All the periosteal matrices act homogeneously by

means of osseous deposition and resorption.

The muscles are attached either into the skeletal

tissue or indirectly by fusion with the outer fibrous
layer of periosteum.

Functioning muscles influence developmental

changes in the form of skeletal tissue to which they
are attached.
The periosteal matrices stimulation causes growth
of the microskeletal units.

The growth process that occurs due to periosteal

matrix stimulation is called Transformation.
CAPSULAR MATRIX

Included in this matrix are those masses

and spaces that are surrounded by capsules.

Example:

Neural mass with scalp and dura.

Orbital mass with supporting tissues of

the eyes.
Capsules tend to influence macroskeletal units
which means portions of several bones are
simultaneously affected.

This sharing of reaction by several adjacent bones

constitutes a macroskeletal unit.

Each capsule is an envelope which contains a series

of functional cranial components ,skeletal units
and their related functional matrices and is
sandwiched between two covering layers.

Examples: neurocranial capsule and orofacial

capsule
Neurocranial capsule:

In this cover consists of skin and duramater ,the

neurocapsular matrix consists of the brain , leptomeninges
and CSF.

The expansion of the enclosed and protected capsular matrix

volume is the primary event in the expansion of
the neurocranial capsule.
The expansion of the neurocranial capsule is
proportional to the increase in neural mass. This can
be shown by considering hydrocephaly as an example.

This suggests that the neural skull does not grow first
and thus provide space for the expansion of the neural
mass rather the growth of neural mass is primary and
causes secondary compensatory growth of the skull.
Thus the point is simple the
neural skull does not grow
first and provide space for
the secondary expansion of
the neural mass rather the
expansion of the neural
mass is primary event
causing growth of the neural
skull.
Orofacial capsule:

All the functional cranial components of the facial

skull arise, grow and maintained within the
orofacial capsule.

This surrounds and protects the oropharyngeal

functioning spaces, and the volumetric expansion of
these spaces serves as a primary morphogenetic
extent in facial skull growth.

Moss ML,SalentijinL.:The primary role of functional matrices in facial

growth: Am J Orthod june 1969; 55;566-77
These spaces are left over as it were, when facial
bones, muscles blood vessels and nerves complete
their growth.
The patency of these spaces are vital in the metabolic
demands of the body.
E.g. airway passage (accomplished by a dynamic
musculoskeletal postural balance the mechanism)
open masticatory cavity
SKELETAL UNITS

May be composed of bone, cartilage or tendinous

tissue. Each bone is composed of several micro
skeletal units.

The possible interaction between periosteal matrix

and microskeletal units includes pterygoid –
Gonial angle, temporalis-coronoid Process.
When the adjoining portions of a number of
neighboring bones are united to function as a
single cranial component it is termed as
macroskeletal unit .

e.g. Endo cranial surface of the calvaria, maxilla,

Mandible

The overall skeletal growth is a combination of

changes in microskeletal and macroskeletalunits
due to stimulation of periosteal and capsular
matrices respectively.
In the mandible we distinguish the following
micro skeletal units.

 Coronoid micro skeletal unit – related to

functional demands of temporalis.

 Angular micro skeletal unit – related to activity of

both masseter and medial pterygoid.

 Alveolar unit – related to presence of teeth.

 Basal micro skeletal unit – related to inferior

alveolar neurovascular bundle.
These micro skeletal units are relatively
independent of each other.

The term functional matrix is by no means implies

only to soft tissues but also includes muscles,
glands, nerves, vessels, fat, teeth etc.

Most of the orthodontic therapy is firmly based

on the fact that when this functional matrix
grows or is moved, the related skeletal unit
responds.
AGAINST

• Spheno-occipital synchondrosis - Demonstrates

autonomous growth

• Nasal cartilage
- Scott- midfacial growth not responsive to
external Influence
- Removal - deficient growth
- Destruction of cell proliferation potential
without cicatrization – Deficient growth
• Craniostenosis – premature stenosis of sutures
inhibits growth – sutures have some capacity to
regulate the activity of functional matrix
TRANSMISSION OF FUNCTIONAL STIMULUS TO
THE BONE-NEUROTROPHISM

Neurotrophism is a non impulsive transmittive

neurofunction involving axoplasmic transport
providing for long term interaction between neurons
and innervated tissue , which homeostatically regulate
the morphological compositional and functional
integrity of those tissues.
Types of neurotrophism:
1.Neuromuscular
2.Neuroepithelial
3.Neurovisceral

Neuromuscular trophism:
Neural innervations are established at myoblast stage.
The genetic control cannot reside solely in the
functional matrices alone and there is neurotropically
regulated homeostatic control of genome and similar
neurotrophic mechanism exist for capsular matrix
which passively regulate the functional cranial
component.
Muscle denervation-reinnervation: muscle
denervation and reinnervation enable us to
diffrentiate effect on muscle tissue associated with
loss of impulse conduction and contraction from
those due to loss of neurotrophic factors.

If motor neurons are sectioned and the muscle

subsequently becomes reinnervated there is
reformation of neuronal conductive function, this
demonstrates neuromuscular trophism.
Neuroepithelialtrophism:
The neurological work of neurotrophism
first began in dermatology.

The factors which contribute to

neuroepithelialtrophism are:

1. local mechanism operating in areas of

high mitotic activity

2.Epithelial growth factors.

Neurotrophic control of genetic activity:
demonstrated in many tissues under
experimental conditions.

Protein synthesis in oral epithelial cells

and specific enzymatic sysnthesis in
taste buds epithelium appear to be
neurotrophically regulated.
 FUNCTIONAL MATRIX THEORY REVISITED

Functional matrix hypothesis could not describe how extrinsic ,

epigenetic functional matrix stimuli are transduced in to regulatory
signals by individual bone cells or how they communicate to co –
ordinate multicellular response.

CONSTRAINTS OF FUNCTIONAL MATRIX THEORY

Methodical Hierachial
Macroscopic measurements Did no explain the process at the
point mechanics cellular level, subcellular or molecular
- Arbitary reference frames. level

87
FMH revisited 1- mechanotransduction
FMH revisited 2- the role of an osseous connected
network
FMH revisited 3- Genomic thesis
FMH revisited 4-The epigenetic antithesis and the
resolving synthesis
CURRENT CONCEPTS OF FMH:

THE MOLECULAR BASIS

The fmh failed to explain the sequences of events
through which the extrinsic stimuli caused adaptive
responses in the skeletal structures i.e. the flow of
the signals that generated required response.

The new researches focused on intercellular

signaling, communication and signal
transduction from the molecular matrix to
micromolecular matrix.
CONCEPT OF MECHANOTRANSDUCTION

Mechanotransduction signifies cellular

signal transduction.

It is the process by which macromolecular

extrinsic stimuli are converted into cellular
signals, which can be internalized by a cell and
processed so that a suitable adaptive response can
be generated.
OSSEOUS MECHANOTRANSDUCTION

It is a highly specialized and unique mechanism by

which bone cells respond to extrinsic stimuli.

It occurs in single bone cells and bone cells are

computational elements that function
multicellularity as a connected cellular
network.
The unique nature is highlighted by the following
facts:

1. Unlike other mechanosensory cells ,bone cells

are not specialized for such stimuli

2.These cells shows aneural transmission of signals

3. Bone cells show multiple adaptational responses to

a single force ,in contrast to singular response by
other tissue cells

4. The changes brought about by stimuli are confined

to a single bone to which the signal is transduced.
Osseous mechanotransduction translates the periosteal
functional stimulus into a skeletal unit cell signal by two
skeletal cellular mechanotasductive process:
1. Ionic
2.Mechanical.

Ionic or electrical processes involves some form of

ionic transport through the bone cell plasma
membrane. The possible process includes stretch
activated ion channel ,electrokinetic and electric field
strengths .
The flow of these ions is thought to occur through the
voltage gated channels or gap junctions between the
adjacent osteocytes .
The passage of K+ ,Ca+ ,Na+, ions across the strained
osteocytes have been proved.

Electric field strength may also be a significant

parameter.

Electrokinetic stimuli in the range of +- 2mv can initiate

both osteogenesis and osteocytic action potential
Mechanical process directly, without the intracellular
transductive process may itself be a strong stimulus
altering the cellular responses through the
transmembrane molecule integrin ,which may
transduct the stimuli directly into the nuclear
membrane.

This cytoskeletal lever chain ,connecting to the

nuclear membrane may have the potential to
activate the osteocytic genome.
BONE AS AN OSSEOUS CONNECTED CELLULAR
NETWORK
•The term ccn implies a network exists between the
adjacent cells of a tissue through specialised
structures in the cell membranes.

•The specialized structures includes the tight

junction ,gap junctions in cell membrane. These
junctions spread stimuli very fast across the
connected cell.

•Extensive ccn exists in the bone and that the

principle component is the gap junction.
•Connexin 43 ,a cytoskeleton protein is the
major constituent this network.

•Gap junction not only connects the osteocytes to the

neighbouring osteocytes but the superficial osteocytes to
periosteal and endosteal osteoblasts too.

•Gap junction allows passage of ionic currents molecules

signals.
• All osteoblast are also interconnected laterally.
•Vertically they connect periosteal osteoblast with
preosteoblastic cells and this in turn is
interconnected ,thus each ccn is like a true syncytium and
are electrically active.
THE FMH AND EPIGENETICS

This concept of moss aims to find a middle path

to solve the controversy of genomic versus
epigenetic control of biologic processes.

Epigenetics is a term which includes : the sum

of all the biochemical, bioelectrical and
biophysical parameters-instantaneously present
intra ,inter and extracellularly- all of which are
produced by the functioning of the cell, tissue,
organ or organism itself.
It should be noted that these same epigenetic factors
serve as an internal environment and must be
considered in addition to the classical external
environment of genetics.

Moss M.L.:The functional matrix hypothesis and

epigenetics:GraberT.M.:Physiologic principles of functional appliances,STLouis;CV
Mosby, 1985;3-4
It is postulated that epigenetic factors act upon the
products of the genome to regulate all developmental
processes leading to the production ,increase and
maintenance of biological structural complexity and
provides feedback regulation of genome itself.

The fmh denies that the genome of skeletogenic cells

contain in and of itself sufficient information to regulate
the type, site ,rate, direction and duration of skeletal
tissue growth.

•But to be sure the modern epigenticist accepts both the

data and fundamental concepts of modern molecular
biology.
VAN LIMBORGH’S COMPROMISE THEORY

Three major viewpoints considered:

Sicher’s
Scott’s
Moss’s
CONTROLLING FACTORS IN
CRANIOFACIAL
GROWTH
Sicher’s view :
• Cartilage
• Sutures
• Periosteum
Are all growth centers
Scott postulates
Intrinsic genetic factors affect:
• Cartilage
• Periosteum
while sutures are passive and reactory.
Moss is felt to have erred in denying any
intrinsic genetic factors in the control of
chondrocranial growth and… restricting the
control of sutural growth to local epigenetic
and environmental factors.
VAN LIMBORG’S COMPROMISE

Chondrocranial growth is controlled by intrinsic

genetic factors

Desmocranial growth is controlled mainly by local

epigenetic factors

Desmocranial factors is also controlled by local

environmental factors

General epigenetic and general environmental

factors have very little role to play.
SERVO SYSTEM THEORY/CYBERNETICS
THEORY(1972)

AlexandrePetrovic, explained that the growth of

various craniofacial regions is the result of
interaction of a series of causal changes and
feedback mechanisms.

 Based on a series of experiments , Petrovic

and coworkers have formulated a cybernetic
model for the control of mandibular growth.
Essence of the theory:
According to the theory ,control of primary
cartilages (mid face) takes a cybernetic form of
“command” whereas control of secondary
cartilages like condyle is comprised of both
direct effect of cell multiplication and also
indirect effects.
Simply stated, the servo system theory is characterized by
the following two principal factors:

 (1) The horizontally regulated growth of the midface and

anterior cranial base, which provide a constantly
changing reference input via the occlusion
(2) the rate limiting effect of this midfacial growth on the
growth of the mandible.

While growth of the mandibular condyle and of the

sutures may be affected directly and indirectly by systemic
hormones, growth of these structures is clearly more
compensatory and adaptive to the action of extrinsic
factors, including local function as well as the growth of
other areas of the craniofacial complex.
Theory:
According to this the midface grows downward
and forward under the primary influence of the
cartilaginous cranial base and nasal septum ,
influenced principally by the intrinsic cell tissue
related properties common to all primary
cartilages and mediated by the endocrine system.

Carlson D.S.:Growthmodification:from molecules to mandible: reprinted from:

McNamara J.A.:Growthmodification:Whatworks,whatdosen,t and
why?,Craniofacial growth series 35,The University of Michigan,Ann Arbor,1999
This theory starts with the explanation of
cybernetics.

 cybernetics is the science of control and

communication in the animal and machine.

The theory postulates that every thing affects

everything and therefore organized living systems
never operate in an open loop manner.

Open loop is a type of feedback mechanism. The other

type of feedback is closed loop mechanism.
The feedback closes the regulation loop of
a given system in the following way..
According to cybernetics theory ,the behaving organism is
not seen as a passive respondent called into action by the
changing environmental stimuli but as a dynamic system
which continuously generates intrinsic activity
for organized interaction with the environment.

Cybernetics in craniofacial growth:

cybernetics demonstrate the relationship between
observational and experimental findings.

Black box: The physiologic system under investigation is

represented by the block box .The contents of the black box
is usually not known.
Feedback signal:It is the function of controlled
variable that is compared to the reference input
 It is negative in regulator and servo system.

Closed loop system: If a physiologic system is

designed to maintain a specific correspondence
between inputs and outputs, in spite of disturbance .

 It is called as closed loop system .

 It is characterized by the presence of a

feedback loop and comparator state.
Closed loop has two variations namely
regulator and servo system.

Open loop system has no feedback

loop or comparator.

The regulator: The main input is a

constant feature in this system .The
comparator detects disturbances and
their effects.
The servo system : It is also called as follow up
system .
 The main input is not a constant in this system but
varies across in time.

Elements of the theory:

Command is a signal established independently of
the feedback system under scrutiny.
It affects the behavior of the controlled system
without being affected by the consequences of this
behavior.
Examples :
 secretion rates of growth hormones ,
testosterone ,estrogen , stomatomedin. They are not
modulated by variations of craniofacial growth.
References input elements :

establish the relationship between the command and

reference input. Includes septal cartilage, septo-
premaxillary ligament, premaxillary and maxillary
bones.

Reference input is the signal that establishes a

standard of comparison ,eg. Sagittal position of
maxilla. Ideally it should be independent of the
feedback.
The controller is located between the deviation signal
and the actuating signal.

The confrontation between the position of the upper

and lower dental arch is the comparator of the
servosystem.

Activity of the retrodiscal pad and lateral pterygod

constitutes the actuating signals.

The elastic meniscotemporal and meniscomandibular

frenum of the condyle form the retrodiscal pad.
The controlled system is between the actuator and
controlled variable, i.e. growth of the condylar cartilage
through retro discal pad stimulation.

Controlled variable is the output signal of

the servosystem . Best example is a
sagittal position of mandible.
The gain: the gain of a system is the
output divided by input .

Gain value greater than one it is called amplification

and if its less than one it is called attenuation.

The pterygocondylar coupling is an example for gain

The disturbance: any input other than the
reference required is called a disturbance. It
results in deviation of the output signal .for
example: increase in hormone secretion
results in supplemental lengthening of
mandible.

The attractor: this is the final structurally

stable state in a dynamic system.

The repeller: this includes all unstable

equilibrium states like cusp to cusp occlusal
relationship.
The influence of somatotrophic hormones on the
growth of cartilages of nasal septum,
sphenooccipital synchondroses and other follows
that of a cybernetic form of command pattern.

•Related to this event the maxillary dental arch

is carried into a slightly more anterior position.
this is the first and primary event.

•This causes a minute discrepancy between the

upper and lower arches ,which Petrovic referred
to as the comparator that is the constantly
changing reference point between the positions
of the dental arches.
EVIDENCES AGAINST THE THEORY

•Goret-Nicaise, Awn (1983) found that the

resection of lateral pterygoid muscle fails to
diminish condylar growth.

•Das, Meyer, Sicher (1965) found that the

occlusion remained unaffected in
condylectomy studies.
The Aponeurotic Tension Model of
Craniofacial Growth in Man Richard G. Standerwick1,*
and W. Eugene Roberts2
The Open Dentistry Journal, 2009

Growth of the viscerocranium is believed to be

influenced by the superficial musculoaponeurotic
systems (SMAS) of the head through residual tension
in the occipitofrontalis muscle as a result of cephalad
brain growth and cranial rotation. The coordinated
effects of the regional SMAS develop a craniofacial
musculoaponeurotic system (CFMAS), which is
believed to affect maxillary and mandibular
development
This CFG model describes the affect of late cephalad growth
of the brain pushing on the occipitofrontalis muscl), which
places the muscle in tension, the peak in temporal and
occipital lobe gray matter being at 16-20 of years age . The
tension force is transmitted from the occipitofrontalis down
through the mask of muscles overlying the face due to
individual muscle fiber blending with adjacent muscles and
the associated superficial musculoaponeurotic systems
(SMAS; investing connective tissues)

This facial muscle mask and associated regional SMAS are

believed to be part of a CFMAS important in directing
craniofacial development and jaw rotation by acting as a
conduit for the brain derived force.
Cranial rotation is also believed to occur sagittally
around the atlantooccipital joint as a result of
allometric brain growth and progressive facial bone
pneumatization with sinus development.
 Brain extension consists of uprighting of the cerebral
portion of the brain, and therefore also the head,
relative to the body axis during growth and
development. This pattern is mimicked by cranial base
(basicranial) flexure and airorhynchy (posterior and
upper portions of the face rotate dorsally relative to
the posterior cranial base by extension of the ACB
relative to the posterior cranial base (PCB), analogous
with the “facial block”).
CONCLUSION
Craniofacial growth and development are based to
large extent on evolving concepts .
At the start these concepts were based on naïve
assumptions about the perceived competing roles
of heredity and environment ,often framed within
the context of the age-old “nature nurture”
controversy.

ATTEMPTS TO PROVE OR DISPROVE these theories have

been extensive however experimental models have
experienced flaws and opinion remains DIVIDED.
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•Contemporary orthodontics-William.R.Proffit
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Book Medical Publishers:1988.p.48-50
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Thomas Graber ,1stediton
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O.P.Kharbanda 2nd edition
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3rd ed. Philadelphia:Saunders;1992.p.133
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1954:94;333-62
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Angle Orthodontist:1963 July:vol 33(3);139-61
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Carlson DS:Growthmodification:from molecules
to mandible: reprinted from: McNamara
J.A.:Growthmodification:Whatworks,whatdosen,t
and why?,Craniofacial growth series 35,The
University of Michigan,Ann Arbor,1999

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THEORIES OF GROWTH & DEVELOPMENT

Dr.Thameem Ahmed.R
1 st yr PG