SEMINAR

ON
FUNCTIONAL MATRIX
HYPOTHESIS & REVISITED

BY,
Dr. Hiten Kalra

Department of Orthodontics and Dentofacial Orthopedics,

S.D.M. College of Dental Sciences and Hospital, Dharwad.

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 FUNCTIONAL MATRIX HYPOTHESIS
Introduction

The concept of functional matrix hypothesis was introduced

Melvin Moss. This hypothesis did not came into existence in one
day but has a long history.

It was in the years 1948-51 when Moss studied in the

department of Anatomy at Columbia University. Where he learnt
that it was possible observing bones as a whole but rather by
considering th em as being composed of a series of ‘segments’, a
term which later became refined to ‘skeletal units’. It was during
this time when Moss started working on the concepts proposed by
‘Van der Klauvw’ that all bones in reality are composed of several
functional cranial components in his (original terminology) and
the growth of these components were relatively independent of
each other. Thereafter Moss carried out various experimental
procedures and laboratory studies and ten years o f work was
culminated and for the first time functional approach to the
problems of craniology was published in 1960  by Moss and
Young in AJ of Physical Anthropology.

After that several paper were published in the subsequent

years and it was finally in the year 1981 Moss gave the classical
statement ‘THE ORIGIN, GROWTH AND MAINTAINANCE OF
ALL SKELETAL TISSUES AND ORGANS ARE ALWAYS
SECONDARY, COMPENSATORY AND OBLIGATORY TO
TERMPORALLY AND OPERATIONAL PRIOR EVENTS OR
PROCESS THAT OCCUR IN SPECIFICALLY RELATED NON-
SKELETAL TISSUES ORGANS OR FUNCTIONAL SPACES
(FUNCTIONAL MATRICES)’

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 Moss – said that head and neck consists of a number of
relatively independent and yet integrate functions – i.e.

o Digestion

o Respiration

o Speech

o Olfaction

o Balance

o Vision

o Neural integration

Each of this function is completely carried out by a 

FUNCTIONAL CRANIAL COMPONENT.

Each such component is composed of two parts.

1) Functional matrix – which actually carries out the function.

2) Skeletal unit – whose biomechanical role is to protect and /

or support its specific functional matrix.

Before going into the details of these 2 components it is

important to understand the basis concept of how growth is
brought about. This are 2 mechanism i.e. transformation and
translation.

Transformation refers to change in the shape and size of

skeletal unit which is brought about by osseous deposition and
resorption which is a direct response to primary morphogenic
demand of specifically related function matrix.

Translation – refers to change in the spatial relation of the

skeletal unit which is brought about by the movement in space,
occurring passively and without necessity of osseous deposition or
resorption.

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 Skeletal unit – co mposed variably of bone, cartilage or
tendinous tissue.

- Macroskeletal un it

- Microskeletal unit

Macroskeletal – when adjoining portion of a number of a number

of neighbouring bone are united to function as a single cranial
component.

Example – endocranial su rface of calvaria.

Microskeletal units:

Wh en a bone consists of a number of small skeletal units.

Example – Both maxillary and mandibular are formed by

number of such contiguous microskeletal units.

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Maxilla – Orbital

- Pneumatic

- Palatal

- Basal

Mandible

 Coronoid – temporalis

 Angular – Masseter and medial pterygoid

 Alveolar – presence and position of teeth.

 Basal – Inferior alveolar neuromuscular triad

matrix.

FUNCTIONAL MATRICES

This not only includes “soft tissue” i.e. muscles, gland, nerves,
vesicle fat etc but teeth which are also considered as functional
matrix.

 Periosteal matrices

 Capsular matrices

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Periosteal matrices

All the non-skeletal functional units adjacent to skeletal unit

form the periosteal matrices. These may be attached indirectly to
the outer fibrous layer of the periosteum or sometimes by insertion
into skeletal tissue itself. This is best explained by the taking the
example of temporalis much and coronoid process. The process
first arises within the earlier formed analage of temporalis muscle
whose contractile ability are well developed in prenatal stages. It
has been proved experimentally – that whenever there is removal,
denervation  decrease in size post infectively / post
traumatically or total disappearance.

Similarly, Functional hypertrophy / hyperactivity  increase in

the

Size and change in shape.

Finally – by altering the much attachment to other

mandibular skeletal unit can produce compensating change in
muscle function itself.

Hence in simple terms – it can be stated that coronoid

process does not grow first and thus provide a “platfo rm” upon
which the temporalis can alter its function but it’s the opposite of
this, which is true. i.e. total growth changes in all aspect of
coronoid process form are at all times a direct and compensatory
response to morphogenetically and temporally prior demands of
the temporalis muscle function.

Similarly, the same effect can also serve to alter th e relative

proportions between contiguous skeletal units. For example the net
backward displacement of several contiguous microskeletal unit
forming the ramus durin g growth.

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CAPSULA MATRICES

An explained earlier all skeletal unit are totally embedded

within the functional periosteal matrices. At the same time this
functional cranial component are organized in the form of cranial
capsule.

4 cranial capsules –

- Neurocranial

- Orofacial

- Otic

- Orbital

Each of these capsule is an envelop containing functional

cranial component which as a whole are sandwiched in between
two covering layer with intervening spaces between the component
themselves and between them and limit of capsule are filled with
loose connective tissue.

The capsule expands in response to volumetric increase of

the capsular matrix. The embedded bones are passively carried
outward by process of growth.

NEUROCRANIAL CAPSULE

- Sandwiched between  skin and dura mater

- It consists of – 5 layer of scalp

- Bone

- Two layer dura mater

Bone consists of number of contiguous skeletal unit  outer

table, Inner table, diploic space.

2 Important points –

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 1) Volume o f the neural mars is import and it makes little
different whether or not this neural mars contains a normal
amount of brain tissue.

2) Expansion of this enclosed and protected capsular matrix

volume is the primary event in expansion of the capsule.

This volumetric increase causes compensatory expansion of

surroun ding capsule as a whole, which takes place by the mitotic
activity of connective tissue.

Later the calvarial functional cranial components as a whole

are passively and secondary translated without the process of
selective periosteal apposition and resorption.

In cases of hydrocephaly  passive, non-periosteal

translative growth is produced by capsular matrices.

OROFACIAL MATRIX

- Surrounds and protect – oronasopharyngeal spaces.

- These spaces constitute capsular matrices for oro-facial

cap sule.

- It is surrounded by skin and mucous membrane on either

side.

- The capsule originates by process of enclosure.

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 - The volumetric growth of these spaces in the primary
morphogenetic event in facial skull growth.

Growth of these functioning spaces causes increase in size of

capsule (by the process of mitosis).

This is followed by passive movement of functional cranial

component.

This brings about compensatory transformation of skeletal

unit in response to an alteration in periosteal matrices therefore

Growth = Transformation + Translation.

MANDIBULAR GROWTH

Can be used for demonstrating the integrated activity of

periosteal and capsular matrices of facial growth.

This is done by

- A longitudinal series of lateral cephalograms representing

mandibular growth is taken 2 assumptions are made.

1. Anterio r cranial base fossa has completed the growth

by end of 3 rd year, so that anterior cranial base is
constan t in shape size.

2. Position of mental foramen does not alter with time.

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First and last series of tracings are taken and superimposition are
done

1) 1 s t superimposition is done on anterior cranial base.

This represents the total growth changes of the mandibular

complex during this period. This is termed as interosseous
growth i.e. total growth, which is relative to fixed anterior
cranial base. (Capsular and periosteal matrix).

2) Second tracing – orient both mandibles so that anterior

cranial base outlines are perfectly parallel and registering
both mand ibular outline on mental foramen.

This represents change in size and shape independent to the

spatial translation termed as intra osseous growth occurring
due to osseous deposition and resorption.

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 Hence, cranial growth is combination of morphogenetically
primary activity of both types of matrix. Growth is accomplished
by both spatial translation and change inform.

In AJO, May 1972, Melvin Moss, stated that investigation

are still going to find out various means and process by which
morpho genetic stimuli of functional matrices are transmitted to
their skeletal units, nature of stimulus, mode of transmission, its
reception and translation and like all research has not true
completion but, rather, is only the beginning of yet another cycle
of work.

FUNCTIONAL CRANIAL COMPONENTS

Skeletal Units Functional matrices

Micro Micro Periosteal Capsular

Eg. Coronoid Eg. Endocranial - Muscle - Orofacial

Angular Surface of calvarium - Teeth -
Neurocranial

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THE FMH REVISITED
Revised Statement

The developmental origin of all cranial skeletal elements

(eg. Skeletal units) and all their subsequent change in size and
shape (eg. Form) and location, as well as their maintenance in
being, are always without exception, secondary, compensatory and
mechanically obligatory responses to the temporally and
operationally prior demands of their related cephalic non-skeletal
cells, tissue, organs an d operational volumes (eg. Functional
matrix).

ROLE OF MECHANOTRASDUCTION

Incorporation of advances in biomedical, bioengineering and

computer sciences have lead to creation of comprehensive
explanatory FMH version.

This revision should indicate

1) Those portions that are retained extended or discarded.

2) Wh ich prior deficiencies are now resolved.

CONSTRAINTS OF FMH

Initially FMH gave only qualitative narrative description of

biologic dynamics of cephalic growth at the gross anatomic level,
and had 2 explanatory constraints.

1) Methodological – Earlier macroscopic measurement

techniques using point mechanics and arbitrary refers frames
like cephalometric were used, those are now replaced by
continuous mechanics technique of Finite Element Method –
FEM which gives a more quantitative view of localized

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 cephalic growth Kinematics to earlier qualitative description
of growth dynamics.

2) Hierarchical constraint

In the earlier version description did not extend downward to

process at cellular, sub-cellular or molecular or extend
upward to multicellular process by which bone tissu e
responded to lower level signals, i.e. it was suspended
between two hierarchical levels.

Moreover, almost all studies of bone adaptation considered

only unicellular, unimolecular level, which could not be
extended to higher, multicellular levels.

MECHANOTRANSDUCTION

It is a general opinion that all vital cells are irritable, i.e. they
respond to alterations in their external environment.

This is brought about by mechanosensing, which includes

Mechanoreception - transmition of an extracellular physical
stimuli into a receptor cell.

Mechanotransduction  transforming of stimulus energetic /

informational content in to intracellular signals.

Osseous Mechanotransduction

Wh enever a load or a stimuli is applied on a bone tissue it,

tends to deform both extracellular matrix and the bone cell, but
only when it exceeds threshold value loaded tissue responds by
trial of bone, cell adaptation process.

OM is unique in 4 ways.

1) Most of mechanosensing cells are cytologically specialized,

but bone cells are not.

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 2) One loading stimulus can evoke 3 adaptational response,
others produce only one.

3) Osseous signal transmission is aneural whereas all other

have some afferent neural pathway.

4) Evoked bone adaptational responses are confined within each

bone organ indep endently. So there is no necessary interbone or
organisal involvement.

There are 2 skeletal cellular mechanotransductive process –

ionic, mechanical.

Ionic / Electrical process

This involves ionic transport th rough the plasma membrane

of bone cell which results in creatio n of ionic or electric signal
which are taken up by connected cellular network which evokes
bone cell response. Other possible, parallel cellular pro cess
include.

Stretch – Activated channels

These (S-A) ion channels are present on the plasma

membrane of bone cells. whenever th e bone is loaded, in strained
osteoblast S-A channels get activated and permit the passage of
certain sized ions like K+, Ca2+, N this ionic flow inturn initiate
intracellular electrical events and modulate membrane potential
and Ca++ influx.

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Electrical

Electromechanical and electrokinetic

Voltage activated ions channels transmit action potential

through gap junction.

Mechanical process

This is explained by the presence of macromolecular

mechanical lnflux, capable of transmitting information from
strained matrix to bone cell nuclear membrane.

The basis of this mechanism is physical continuity of a

transmembrane molecule integrin

Collagen of organic matrix

Extracellular

Integrin

Intracellular

Actin cytoskeletal

Nuclear membrane

Cfosgenes

Intranuclear  processes which regulating genomic activity.

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BONE AS AN OSSEOUS CONNECTED CELLULAR
NETWORK (CCM)

All the bone cells expect osteoclasts, are extensively inter

connected by gap junction, all osteocyte are enclosed within
mineralized lacuna. They have the cytoplasmic processes which
are oriented there dimensionally and interconnect with the similar
processes of upto 12 neighboring.

Gap junction are – found where the plasma membranes of

pain o f markedly overlapping canalicular process material.

These gap junctions are connected with each other to

superficially osteocytes, laterally and vertically forming a CCN.

These type junction are electrical synapsis and permit bi-

directional signal traffic. Once the bone cells are activated
mechanotranductively  the initiate membrane action potentials
which are capable of transmission through those interconnected
gap junction.

Hence, this can be compared to a “Artificial neural network,

parallel or parallel distributed signal processing occur.

The network which is formed  is such the cell are

organized in to layers – an initial output, a final output,
intermediate / hidden layer.

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 Wh en a stimuli is applied  it results in what is called
loading in the initial layer. Within each cell call the weighted
inputs – (stimuli) are summed and compared to against some
threshold value. If this value exceed, intracellular signal is
generated. This signal is then transmitted identically to all the
hidden layers to which the each initial layer to connected by gap
junction. Next, similar processes of weighted signal summation,
comparison and transmission occur in these intermediate layer unit
the final layer all are reached – this output finally determines rate,
direction, magnitude, duration of specific adaptation.

Hence CCN  shows oscillation i.e. reciprocal signaling

feedback between layers and at the same time the presence of
sharp histological discontinuities between different groups of
osteoblasts which discretely stored in a CCN – preventing the flow
of information.

Hence it can be summarized that CCN –

1) Devlopmentally – unstrained, self organized, self adapting,

epigenetically regulated system.

2) Operationally – stable, dynamic system capable of

oscillatory behaviour.

3) Structurally - non modular, variation in organization permit

discrete processing of different signals.

It is the strain  which is important in all bone adaptation

or remodeling respon se and a significant strain attribute may vary
with specific co ndition. These are

1) Loading – bone responds best to dynamic rather that static

loading.

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 2) Frequency – osteocytes can physiologically tuned to
frequencies of muscle junction.

3) Magnitude – relatively small i.e. microstrain (me) 10-6 mm/

mm and magnitude of 200 ± 100 me are morphogenetically
competent.

• Initial version – epigenetic factors

• Current – genomic regulation of growth

• Emergence- appearance of complex higher

levels by self org anization at lower levels.

This can be explained by the concept of

CAUSATION

There is a contin uing controversy concerning the role of

genomic and non geno mic processes in the regulation of growth –
this can be resolved by several types of causation.

Causation – A classical categorization of causation.

Materia Forial Efficient Final

(What is acted (Set of rules) (Immeidate Some

upon cell and geno mic preceding event) consequent
extracellular regulation Epigenetic event use.
substance)

Intrinsic prior necessary

AN ODOTOGENIC EXAMPLE:

Rigid genomic control of odontogenesis

 a. Temporally sequential

b. Spatially restricted

 Epigenetic regulation of odontogenesis

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  Various experiments on polyphydont chiclid fish.

THE GENOMIC THESIS

In simple term states that it the genomic of an in dividual i.e.

(DNA sequence) which determines the overall phenotype.

BIOLOGICAL BASES

It is believed that only 10% of the geno mic is related to

phenotypic ontogenesis.

The enconding of 10% DNA show it exists in 2 families.

It is in conditions where there is absence of normal types, or

presence of structurally abnormal type – a patholo gical state
appear.

Application of Genomic thesis in orofacial biology

It is claimed that craniofacial development is controlled by 2

inter related, temporally sequential processes.

1) Initial regulatory (homeobox) gene activity.

2) Subsequent activity of regulating molecular (group 

growth factors families  steroid / thyroid super family).

1. Homeobox gene – coordinate in the development of

cranio facial structures.

2. Regulating molecules – alter the manner in which

homeobox gene co-ordinate hence  contributing to
abnormal development.

Similar – orthodontic implication of this thesis – claims

that – poor co-ordinatio n of form and size of structures (teeth and

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jaw) by regulating genes  malocclusion and dentofacial
deformities.

THE EPIGENETIC ANTI THESIS

Broadly speaking 

Epigenetics refers to the entire series of interaction, among

cells and cell products which lead to morphogenesis and
differentiation.

Epigenetic factor includ e

1) All of the extrinsic, extraorg anismal macroenvironmetal

factors which impinge on vital structures (for eg – food,
light, temperature) including mechanical loadings and
Electromagnetic fields.

2) All intrinsic, in tra organismal, biophysic biomechanical,

bio-electrical microenvironmental events occurring on, in
and between individuals cells, extra cellular materials, cells
and extra cellular substances.

Epigenetic process and mechanism

In craniofacial morphogenesis  presently more is kn own

about the process than mechanism.

Process  Series of action or operation that lead towards a

particular result.

Mechanism  Fundamental physic or chemical process(es)

involved in, or respo nsible for an action, reaction or other nailed
phenomenon.

That is – Mechanisms underlie processes.

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EPIGENETIC PROCESS OF LOADING

Many different epigenetic processes can evoke mechanism

capable of modifying DNA.

2. Among those mechanical loading is very important.

3. Loads may be imposed at a microscopic level  molecular /

cellular level or at tissue level.

4. It may be dynamic (muscle contraction) or static

(gravitational).

To be effective  load may – increase; decrease or remain

constan t.

Epigenetic mechanism involved include

1) Extracellular matrix deformation – Regulating multicellular

tissue morphogenesis.

2) Altering cell shape  Controlling osteoblasts gene

expression.

Epigenetic regulation at higher structural level

Regulation of periosteal matrices. An inter relationship is

present between genotype and phenotype. Mechanical loads
regulate phenotype. Genotype is mainly expressed via the
processes of neurotropism.

DIALETICAL ANALYSIS

THESIS ANTITHESIS

(GENOMIC) (EPIGENETIC)

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 RESOLVING THESIS

RESOLVING THESIS

This required because it is clear that both g enomic and

epigenetic processes were necessary to exp lain growth and
development genomic factor are considered intrinsic and prior
cause. epigenetic factors are considered extrinsic and prox imate
cause.

COMPLEXITY THEORY

This theory gives a description of the behavior of complex

biologic system which exists as ensembles and not as luster of
individual cells and extracellular substances.

In the system functional cranial unit is termed as Complex

Adaptive System (CAS).

CT theory assumes that CAS processes bo th the genomic and

epigenetic information hence growth and development is not fully
predictable and ontogeny involves a non linear process.

Environmental factor thus play a decisive role in ontogenetic

processes. But it is the organism itself that as an integrated system
dictate the nature of every developmental response. The living
organism self organizes on the basis of its own internal
structuring, in continuous interaction with environmental in which
it find itself.

COMPLEX ADAPTIVE SYSTEM

 “en semble” of several tissue and organs.

 CAS processes genomic and epigenetic information in

parallel manner.

 Minor changes in epigenetic input result in huge fluctuation

in morphogenetic output.

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  Ontogeny is a nonlinear process.

 Spontaneo us self organizing ontogenic processes and

mechanism can create phenotypic variability under constant
genetic and other extra organismal epigenesis condition.

 “Enviromental factors play a decisive role in all ontogenic

processes. But it is the organ ism itself that, as an integrated
system ,dictates the nature of each and a very developmental
response.

 The living organism self organizes on the basis of its own

internal structuring, in continuou s interaction with the
environment in which it finds itself”

 Latham-’9 5

Complexity theory requires comprehensive presentation and

according to Moss will be reviewed subsequen tly.

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 BIBLIOGRAPHY

 Contemporary orthodontics-WILLIAM PROFFIT

 Principles and practice of Orthodontics - T.M.GRABER

 Primary role of functional matrix in facial growth-Moss

AJO-june 1969

 The doctrine of functional matrices-James Scott AJO

july1969

 The capsular matrix-Moss AJO nov 1969

 Twenty years of functional cranial analysis-Moss AJO may

1972

 Genetics, epigenetics and causation-Moss AJO october 1981

 Functional matrix hypothesis revisited-Moss AJO july-oct

1997

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