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Washington Academy of Sciences

Order Form for New Book

75 Years of Scientific Thought

The Washington Academy of Sciences has just published a book, 75 Years of Scientific Thought, containing 25 of the

best papers published in the Journal of the Washington Academy of Sciences during its first 75 years of existence

(1911-1986). Eight of the papers were written by Nobel laureates.

The book was compiled by the Academy’s Committee on Scholarly Activities over an intensive two-year period of

review. The Committee was chaired by Dr. Simon W. Strauss. Members of his committee were Drs. Robert F. Blount,

Randall M. Chambers, Lloyd E. Church, Robert A. Owens, and Bhakta B. Rath.

The papers, listed below, cover a wide variety of scientific fields, and provide a classic portrayal of scientific thought

over the past 75 years.

A limited number of copies are available for sale on a first come basis. The cost to Academy members is $15.00, and

for non-members is $30.00. The price includes postage and handling. Please complete the form below and return it with

your check, payable to Washington Academy of Sciences.

Chronological Title Index

1913. Recent Theories of Heat and Radiation. W. Wien*

1918. The Size and Shape of the Electron. Arthur H. Compton’

1918. Biology and War. Raymond Pearl

1921. Chemical Structure and Physiological Action. C.L. Alsberg

1929. Zoogenesis. Austin H. Clark

1932. Synthesis of a Human-Nucleus, An Important Constituent of Humus in Soils, Peats and Composts.

Selman A. Waksman’ and K.R.N. Iyer

1933. High Voltage. Karl T. Compton

1935. What is Electricity? Paul R. Heyl

1940. The Separation of Isotopes by Chemical Means. Harold C. Urey’

1941. Physical Reflections in A Chemical Mirror. R.E. Gibson

1942. Great Astronomical Treatises of the Past. Edgar W. Woolard

1944. Filed Indian Teeth from Illinois. T.D. Steward and P.F. Titterington

1945. Faster than Sound. Theodore von Karman

1946. An Application of the Theory of Themes in Culture. Morris Edward Opler

1947. Joseph Henry Builds an Institution. Paul H. Oehser

1947. Wind River Shoshone Literary Forms: An Introduction. D.B. Shimkin

1948. General Survey of Certain Results in the Field of High Pressure Physics. Percy W. Bridgman*

1948. The Chemical Nature of Enzymes. James Batcheller Sumner’

1954. Mesons and Nuclear Forces. Hans A. Bethe*

1956. Atomicity and Patterns. Sir George P. Thomson

1957. Cultural Implications of Scientific Research. R.E. Gibson

1958. Gravitation: An Enigma. R.H. Dicke

1959. Men and Electrons. L. Marton

1977. A Century of Cryogenics. R.P. Hudson

1978. The First Trans-Atlantic Cable. Walter D. Freezee

* signifies Nobel laureate

Order Form

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Journal of the Washington Academy of Sciences,

Volume 77, Number 2, Pages 47-54, June 1987

Remarks on Culture and the Self

George L. Farre

Department of Philosophy, Georgetown University, Washington,

D.C. 20057

Introduction

The objective in this essay is to look at

the basic mechanism involved in the for-

mation of the mind within the total con-

text in which human beings assume their

general ecological niche. Because the brain

of man develops largely ex utero, and

therefore primarily in a social context, the

formation of the mind is naturally domi-

nated, and indeed governed, by the re-

markable process of the acquisition of

language.

This process has two major aspects: the

transfer of culture from the community to

each of its members, and the resulting

formation of the self. These two sides of

acculturation provide the rationale for the

division of this essay into two main sub-

stantive sections, one an examination of

the notion of cultural transfer, the other

an analysis of the notion of the self and

of its correlates. These are preceded by a

few general remarks on particular aspects

of language which are relevant to the topic

under discussion.

A study of this sort may be approached

from a variety of angles. In the present

case, the approach is essentially Wittgen-

steinian, in that the notion of deep gram-

mar (i.e. the grammar of meaning) plays

a central role in showing how conceptual

attitudes are affected by cultural transfer,

both in the case of the acquisition of lan-

guage, and in the more general case of

the transfer of information from one cul-

tural area, such as science, to some Other

such as art, religion, policy, etc.

A—On Language

The importance of language to philo-

sophical inquiry derives from the fact that

all of our knowledge about the world and

about our selves is given form and expres-

sion in language. Thus all of our claims

to know or to believe are claims about

what we say so that the analysis of what

we say is the analysis of what we mean.

Malebranche put it rather well when he

said that “language is the locus of thought,

as space is the locus of body’’. Language

viewed in this light is the principal carrier

of intelligence, for it is by its means that

we make ourselves understood.

I—Remarks on Language

How we understand ourselves and our

world is determined in large part by the

tools we use to do it. The principal ones,

language and the conceptual framework

embedded in it are both inherited. In the

48 GEORGE L. FARRE

case of man, where most of the brain de-

velops ex utero, the cultural “wiring” takes

place in a social environment. It is marked

in particular by the acquisition of lan-

guage.

Language is formative of the mind: it

gives it a preferential attitude, as well as

a point of reference from which to ob-

serve nature and learn from it. In this

sense no two languages can translate ac-

curately into each other: for in addition

to the fact that the brain’s “‘wiring”’ differs

in each case, a particular sensitivity and

a way of perceiving are assumed with the

acquisition of a new language. Further-

more, the perspective inherent in any view

of nature does not itself admit of trans-

lation but requires something akin to a

conversion, a sort of “‘Gestalt shift” in

order to be got right. It is the sense in

which the limits of my language are truly

the limits of my world, the sense in which

language embodies a form of life. For lan-

guage is symbolic, and a symbolism pre-

supposes a whole conceptual stage set-

ting, as well as an extrinsic context in

which symbols have their point.

In some cases, a specialised and more

perspicuous language can be constructed

effectively whenever it is thought that the

structure of the natural language lacks the

kind of precision that perspicuity of rep-

resentation alone makes possible. For ex-

ample classical musicians have developed

a written symbolism that represents with

sufficient precision the structure of the

musical phrase; in similar fashion math-

ematical physicists have devised several

languages having the multiplicity required

for the analytical representation of nat-

ural phenomena by means of their struc-

tural properties. Indeed the present cen-

tury has been marked inter alia by the

development of many such specialised

languages, even in areas where the need

was not originally recognized. The point

of these remarks is that, hidden by the

grammatical structure of the natural lan-

guage lies the deeper grammar of the con-

ceptual framework. In principle, this deep

grammar (the grammar of meaning) can

be made overt by using a suitable sym-

bolism.

II—Remarks on Concepts

A key notion of the theory of descrip-

tion is that of concept: it functions as the

primary referring expression. Concepts

play a dual role in description: a refer-

ential role whereby an expression is linked

with elements of a domain of interpre-

tation (e.g. when we ask, of an object,

what it is); and a grammatical (formal)

role, it being the manner in which such

an expression is related to others in the

language (as when we ask of a term what

it means). These two aspects are inti-

mately linked, the former being depend-

ent on the latter.

The grammatical aspect of a concept is

given by the way in which the term used

to represent it relates to other expressions

in the language. This aspect is a direct

consequence of the partial interpretation

of an underlying network of relations, the

“syntactical mesh’’. The interpreted mesh

defines the conceptual framework within

which reference can be made to some ele-

ments of the domain of interpretation. In

principle, it is always possible to formalize

the conceptual mesh, though this is sel-

dom useful outside of the theoretical dis-

ciplines, such as mechanics. The formal

character of a concept is thus nothing other

than the grammatical aspect of its mean-

ing, i.e. its deep grammar. While the deep

grammar of an expression is seldom evi-

dent from its apparent structure the sit-

uation can be remedied (as pointed out

earlier), and the deep grammar made

overt in descriptive contexts by the use of

a notation designed specifically for that

purpose, should the need for one be

found.

The referential aspect of a concept typ-

ically underdetermines its formal aspects,

so that it is not rare for the latter to undergo

evolutionary developments as the cultural

framework changes while its reference is

REMARKS ON CULTURE AND THE SELF 49

left materially unchanged. A given term,

while referring to a particular entity, may

impute to it vastly different properties,

depending on which conceptual frame-

work it is enmeshed in. For example, what

is meant by “atom” differs considerably

to day from what was meant by it a hundred

years ago, even though its denotation has

not changed markedly since then.

The referent of a concept is given in

terms of some general property repre-

sented by a predicate function. Individual

instances of the concept are singled out

through the circumstances attendant to the

actual use of the predicate function, there

being no reason, philosophical or other-

wise why there should be a single instan-

tiation rather than many, or none.

Therefore to say of something that it is

an object of a particular kind is to embed

it in a conceptual framework; in this sense,

it is appropriate to say that the world we

know, that is the one we describe, is con-

ceptually laden [1]. Therein are to be found

the philosophical roots of conventional-

ism.

A referring expression, considered by

itself, does not reveal its meaning in use,

and may in fact be associated with differ-

ent conceptual frameworks on different

occasions, a condition typical of natural

languages. In such cases, the use of the

same expression to refer to different enti-

ties helps to hide the differences in mean-

ing, an effect compounded whenever the

referent picked out is the same object in

each case. Ambiguities such as these are

inevitable in ordinary discourse where a

number of cultural attitudes come to-

gether.

IiI—The Argument Against

Private Language

The argument against the possibility of

there being a so-called “private language”’

[2], presented by Wittgenstein in the

“Philosophical Investigations”, includes

two main points that are made in criticism

of the view that language can be private

in a nontrivial, philosophically interesting

way.

The first point has to do with the wide-

spread assumption, more or less explic-

itely endorsed by philosophical writers

since antiquity, that the meaning of de-

scriptive expressions is in some essential

way dependent on their referents, rather

than the other way around (referential

theories of meaning).

Wittgenstein argues that, even in the

paradigmatic case of names, referential

theories fail to account for the meaning

of referring expressions primarily, though

not exclusively, because the existence of

their referents is immaterial to their sense

whether these be construed in the classical

manner as public objects (such as Julius

Ceasar or the circle) or as private objects

in the manner of the Moderns (i.e. as states

of consciousness) [3].

The second point has to do with the

observation that it is not possible to use

language in unheard-of ways in order to

effect a publicly defined task, such as re-

ferring, describing, promising, and so

forth.

The upshot of this is an argument for

the inescapably public character of all

meaningful expressions, including those

reporting on states of consciousness,

whether to oneself or to others.

B—On Culture

This part is divided into two sets of re-

marks. The first group is designed to high-

light various aspects of human culture,

with language as its principal distinguish-

ing feature. A number of important no-

tions are introduced here; these in turn

set the stage for a discussion of the phe-

nomenon of cultural transfer.

The second set of remarks is given to

various aspects of cultural transfer. The

frequent emergence of cultural gradients

within societies undergoing this process

reveals the existence of stresses associated

with information transfers across cultural

boundaries; it also points to the impor-

tance of having a /ingua franca to cut across

50 GEORGE L. FARRE

ideological lines and to help in the emer-

gence of new syncretic cultures within so-

cieties marked by a measure of cultural

heterogeneity (including world society).

I—Remarks on Culture

The culture of a people results from the

activities they engage in. The more varied

these activities, the richer the culture is

likely to be.

Human activities are modes of adap-

tation to the world which man inherits at

birth. Some of these are clearly intended

to improve personal survival (e.g. farm-

ing), while others affect the survival of

the social group as a whole (e.g. organ-

izing). Others are designed to improve the

quality of life (e.g. medicine), or one’s

understanding of nature (e.g. the natural

sciences). Some activities are meant to

achieve a significant mode of expression

for one’s aspirations (e.g. the fine arts),

while still others probe beyond the limits

of nature and reach for the transcendent

(e.g. religion).

To talk of the culture of a people is not

to say that all of the individuals in society

engage in all of the activities constitutive

of that culture. For aside from the obvious

case of the elderly and the young whose

scope of activities is limited, people tend

to specialize, a process which adds to the

survival value of society and to the quality

of its culture. The division of labors leads

inevitably to an uneven distribution of

cultural activities and, by way of conse-

quence, to an uneven distribution of the

benefits such activities bestow. So we may

ask, what does it mean to talk about the

culture of a people if it is the culture of

no one in particular?

In general, human societies are cultur-

ally identifiable in terms: both of a com-

mon language and of social institutions

(government, laws, etc). These provide

the stage upon which the development of

culture and the differentiation of its ele-

ments can take place.

Of the cultural traits that give cohesion

to human society, language is by far the

most significant. The natural language first

acquired by an infant (the mother’s tongue,

so aptly called), enmeshes it into the com-

mon social and cultural fabric by giving it

the means to understand itself and its

world. Eventually the language will ena-

ble the child to objectify its own self and

thereby to become a person in the ac-

cepted cultural sense of the word.

Each of the constitutive elements of a

culture (i.e. each of its constitutive activ-

ities) has its own conceptual framework,

and to that degree its own language. The

businessman and the politician, the artist

and the physicist, the farmer and the law-

yer, each lives in a different cultural world,

a fact normally hidden by the shared nat-

ural language. In this sense we may look

upon the activities imbricated in the gen-

eral culture as its subcultures.

Different as they may be, these sub-

cultures are obviously connected and in-

fluence each other. The over-all pattern

they form, and the relative importance of

each component within the whole, define

the cultural profile of a society, and thereby

provide it with a criterion of cultural iden-

tity. This in turn makes possible the com-

parison of different cultures. In general

the more complex the cultural fabric, the

richer it is, the wider is its scope and the

greater are the possibilities of its growth.

A cultural profile may evolve with time.

Changes in the cultural profile tend to

occur gradually. Sometimes however,

abrupt changes take place, which mark

the onset of cultural revolutions. The sci-

entific revolution, the French revolution,

the industrial revolution, the technologi-

cal revolution, among others, are in-

stances of this phenomenon.

In general, the rate of change in the

culture of a society tends to be uneven,

and this introduces a degree of cultural

stratification. The most influential factor

in this process of cultural change is un-

doubtedly the finite but appreciable time

it takes for new information to be ac-

quired, processed and distributed, those

REMARKS ON CULTURE AND THE SELF 51

at the leading edge, where the informa-

tion originates being different from most

of those who make use of it. Once re-

ceived, the new information is usually

processed further and re-emerges even-

tually in a degraded but more useful form

(i.e. one that is less general).

Ii—On Cultural Transfers

The net effect that the acquisition of

new information has on the recipient is a

modification of his conceptual frame-

work, which in turn affects his whole cul-

tural attitude. Therefore, it is to be ex-

pected that the transfer of information will

create a cultural gradient between its source

and its users. The time of diffusion is al-

most always substantial, and by the time

the new information has been absorbed

and used by those most distant from its

source, the leading group is likely to have

evolved conceptually beyond the stage at

which the information originated, further

steepening the gradient. As a result those

at the leading edge of change do not share

the same conceptual attitude with those

at the trailing edge, a situation familiar to

students of the sociology of science.

Ordinarily, the cultural gradient is

steeper across professional lines than it is

within the sub-culture itself: people en-

gaged in different pursuits find it harder

to communicate about what interests them

than those who are engaged in the same

enterprise and share the same conceptual

apparatus (e.g developments in the biol-

ogy of mind filter more rapidly through

the group working in the cognitive sci-

ences than it does across professional lines

and take even longer to reach the general

public).

If the cultural gradient becomes too

steep (a situation often encountered at

times of rapid developments, especially

across professional lines), discontinuities

are likely to appear in the general cultural

fabric. Depending on their nature and im-

portance, these in turn may elicit strong

reactions within society, perhaps rein-

forced by the perceived incompatibility of

assumed facts with values held (Cf e.g

fundamentalism, environmental extre-

mism, and so forth).

On the global scale, such discontinui-

ties are likely to generate enmity, and per-

haps even open hostility to the activities

of the society with the most advanced cul-

ture (Cf. for example, the attempts by

Third World countries to control both the

acquisition and the flow of information

through UNESCO).

As the specialized language of each

subculture differentiates out of the com-

mon matrix, the associated conceptual

frameworks begin to evolve in different

ways and eventually become incommen-

surable i.e the point is reached when there

is no longer any way to translate from one

language into another with any precision.

The best that can be hoped for in 4 sit-

uation like this is to paraphrase the orig-

inal text in the natural language shared

by all (for example, it is not possible to

translate from the language of plasma

physics into English, and to then proceed

with the physics in that language, any more

than it is possible to translate the poetry

of Dylan Thomas into French, and expect

to achieve the same poetic effect in that

language as in English. Nor is it possible

to translate Beethoven’s Triple Concerto

in any natural language).

Yet in general it is possible to para-

phrase into a richer if less perspicuous

language and to invoke canons of cor-

rectness suitable to the process, thereby

transferring some information across cul-

tural lines. Even though the paraphrase

is not a true translation since the infor-

mation is being distorted in the process,

a transfer does takes place which is suf-

ficient to affect the cultural framework

inherent in the richer language.

In this manner, the general cultural

framework can be modified by the trans-

fer of some information across the lines

that divide its various sub-cultures. What

is required then is a suitable medium, a

lingua franca sufficiently plastic and rich

52 GEORGE L. FARRE

to encode changing notions having their

origins in specialised languages with es-

oteric conceptual grammars. The result

will be a modified cultural framework, and

therefore a different way of seeing things.

In this way general cultures evolve.

C—On the Self

A special case of cultural transfer is that

of the acculturation of infants, where the

effect of transferring the language is

marked by the initial formation of a self

and by its subsequent development into a

person.

What follows should not to be read as

the articulation of a particular view re-

garding the notion of the self, but simply

as indicative of the sort of points such a

view would include. Still the proposed

schema should be sufficiently clear to give

a reasonably fair idea of the theory here

suggested.

The underlying assumption in what fol-

lows is that a non-speaking organism has

no concept of self. This is meant to ex-

clude animals as weil as some higher

organisms deprived of the cultural envi-

ronment needed to acquire linguistic

competence at birth (for example “lEn-

fant Sauvage” of Aveyron, the wild boy

Ramu of India etc). This should not to be

construed to mean that a higher organism

with the capacity for self-consciousness

cannot have a perception of self. The only

claim made here is that such an organism

does not have a concept of self. This no-

tion can evolve only through the process

of adaptation of a suitably complex or-

ganism to its integral environment. This

last includes, in the case of human organ-

isms being discussed here, the cultural and

especially the linguistic dimension. For if

we do not take into account the degree

of anatomical complexity needed for self

consciousness and for its verbal expres-

sion, nor the obvious cultural dimension

of the environment in which man makes

his niche, the only way sense can be made

of the performance of a human organism

is to treat it as a “black box” 1.e. as a

purely mechanical system designed to

transform certain inputs into correspond-

ing outputs. The drawback of this ap-

proach is that black boxes have no selves!

That the approach proposed here is not

the only way to approach the self is evi-

dent, but it has the merit of presenting a

model that is human in the cultural sense

of the term. Man is admittedly a very

complex and sophisticated organism, but

it is not just that: it has emergent prop-

erties, chief among which is thinking. In

this sense Man is, first and foremost,

homo linguificans.

I—The Notion of the Self

The notion of the self that emerges from

these considerations is roughly as follows:

Selves are not born, they are made. The

self, although it may be genetically pre-

determined in important ways, is formed

mostly in the context of its cultural en-

vironment. The key to this process is the

acquisition of language, the importance

of which can hardly be underestimated.

It is the crucial event which provides the

infant with the means to express itself ver-

bally, and to map its world. This however

is a slow process, mostly because the brain

of the infant is not all there at the time

of birth. What may be called figuratively

“the wiring of the brain for language”’

cannot proceed before the Central Nerv-

ous System (or Central Information Sys-

tem: CIS) has developed sufficiently.

Therefore the actual wiring takes place in

an environment in which language is nor-

mally used. It is in this manner that the

infant learns how to speak whatever lan-

guage or languages are spoken in its im-

mediate cultural niche.

Once acquired, this verbal mode of

expression enables the infant to objectify

itself, i.e. to perceive itself as an object,

as a person in the culturally defined sense

of this term. (Cf in this context the inter-

esting though seldom noted contrast be-

REMARKS ON CULTURE AND THE SELF 53

tween the criteria used by Descartes to

define his own self, which are introspec-

tive, and those, purely linguistic, which

he used to ascertain that there are other

selves [4]).

Ii—The Self in Context

The self, having inherited its formal

properties through the language used to

map its features, may be expected to bear

the imprint of its linguistic matrix.

To the extent that the context in which

an infant is immersed is not culturally ho-

mogeneous, its emerging self will be com-

plex and reflect in its structure the variety

of conceptual frameworks associated with

each component of the culture. Such a

complexity is apparent for instance in the

multiplicity of culturally inherited traits,

ranging from the the diversity of the kind

of archetypes studied by C. G. Jung, to

other social traits of character (national,

tribal, family, etc).

The complexity of the self is also the

result of the multitude of conceptual

frameworks at work in any given human

culture, however unsophisticated. It is

common for these frameworks to overlap

in much of the domain of ordinary ex-

perience. Added to the fact that a person

understands himself only in terms of some

conceptual framework, this circumstance

leads naturally to the emergence of a

multitude of co-referential selves, each

bearing the imprint of a set of formal

characteristics derived from a different

grammatical (conceptual) matrix. This

unique cluster of selves will be referred

to as the Self, with a capital S.

In this manner the formation of the in-

dividual Self is often marked, perhaps

inevitably, by profound inconsistencies.

Whenever the circumstances are right,

these inconsistencies can manifest them-

selves and create perceptible tensions

within the individual, from which path-

ologies may in due course develop.

We may infer from this what the main

task of therapy could be: to first identify

the locus delictus, and then to undertake

whatever remedial action is appropriate.

Presumably, this would result in the

smoothing out of the kinks in the con-

ceptual fabric.

Iii—The Self as Person

A person may be defined as a physical

organism functioning as the referent of

multiple selves. Its principal feature is its

Central Information System (CIS): it en-

ables the organism to generate, to process

and to diffuse information, as well as pro-

vides it with a serviceable ground for iden-

tity.

Being a physical organism, a person

normally interacts with similar organisms,

as well as with the over-all materiai en-

vironment. Its mode of interaction with

the environment is naturally physical, that

is mediated through the body, but it is

also non physical in important respects,

especially as it relates to similar orga-

nisms, with whom the most important in-

teractions are mediated by language, the

essential carrier of intelligence.

The person-to-person interactions, and

to a lesser extent perhaps the person-to-

world interactions, are formative of the

person and may be expected to contribute

to its complexity. Taken together, these

interactions entwine the individual in a

complex network of relations which forms

the stage within which the person lives its

history. Such complex enmeshments make

for numerous interactions, leading to the

appearance of endogenous forces in the

Self. From their interplay, there emerges

a fine structure which is uniquely char-

acteristic of that individual.

IV—Egogeny, Identity and Privacy

Three points remain to be mentioned

in order to present a reasonable overview

of the notion of the Self.

54 GEORGE L. FARRE

Defined and understood in terms of

language the Self is naturally of a public

nature. Yet it is eminently private in an

important sense. The reasons for this are

to be found in the natural unity and

uniqueness of the person itself. Three key

elements are constitutive of this privacy,

namely the integrity of the person as a

single organism, the historical develop-

ment of the Self, and the priviledged mode

of access the person has to its own Self.

The internal integrity of the person is

derived from the principle of its organ-

ization: the existence of its Central In-

formation System. The CIS provides one

important criterion of identity, by being

relatively invariant through the vicissi-

tudes of the life of the organism. While

the person adapts to the changing envi-

ronment, and the Self develops and com-

plexifies, its growing sense of internal in-

tegrity gives form to the increasing

awareness of its own identity. In this way,

it provides the person with the grounds

for the co-referentiality of its different

constitutive selves. The lived experience

of this internal integrity may be said to

constitute the material basis for the pri-

vacy of the Self and of the person.

The Self is complex, a direct conse-

quence of the very complexity of the am-

bient culture. It is an ensemble of inter-

active selves, all referred to the same

organism or person. It is also a unique

succession of time-indexed worlds in which

the same person plays the central role.

This succession of worlds, as well as their

defining characteristics, combine to cre-

ate a unique history: that of the formation

of the Self (egogeny).

Given that each person has a privi-

ledged and immediate access to its own

consciousness, and therefore an incorri-

gible knowledge of its contents, the unique

configuration of its formal features may

be said to be private, notwithstanding the

fact that each of its constitutive elements

is essentially public in character: it is pri-

vate to the extent only that it is actualised

in a single person, and thus directly ac-

cessible by that person alone.

It is important to keep in mind that the

Self so defined is not a thing (i.e. a sub-

stance), and therefore is not to be con-

strued as a referent. The Self, as defined

here, is the form of consciousness, itself

the emergent property of certain organ-

isms whose CIS is of a particular nature

and degree of complexity. Its referent is

the person, the ultimate ground of iden-

tity.

References Cited

1. L. Wittgenstein: Tractatus Logico-Philosophi-

cus.

2. L. Wittgenstein: Philosophical Investigations.

3. An object is private in the sense that its acces-

sibility is.

4. R. Descartes: Discourse On Method: Part IV,

33; Part V, 56-60.

Journal of the Washington Academy of Sciences,

Volume 77, Number 2, Pages 55-65, June 1987

The Influence of Natural

Resonances on Scattering and

Radiation Processes

D. Brill

Physics Department, U.S. Naval Academy, Annapolis,

MD 21402-5026

and

V. Ayres and G. Gaunaurd :

Research and Technology Department, Naval Surface Weapons

Center (R-43), Silver Spring, MD 20903-5000

ABSTRACT

Resonances present in the many physical processes associated with radiation and scat-

tering phenomena are being used in many applications ranging from radar target masking

to sonar target identification.

Introduction

Resonances have traditionally been re-

garded as an engineering problem. Many

an engineer has lived in fear of designing

a structure destined to go into an envi-

ronment where the ambient spectrum

contained a large amplitude at one of the

structure’s resonances. Two famous tra-

gedies which justify the concern are: the

collapse of Tacoma Narrows Bridge and

the loss while in flight of some Lockheed

Electras, an early turboprop airplane.

This has led to a historical revulsion to the

concept of resonance and probably leaves

many engineers feeling that nothing good

could ever come from such a troublesome

phenomenon. Not so!

Here, instead, we emphasize that the

spectrum of resonances inherent in a

physical structure carries information

about the structure’s size, shape, and ma-

terial composition. Looked at this way,

this phenomenon takes on the appearance

of a relatively unacknowledged reservoir

of information.

Singularities are the spice of life! With-

out them, many physical phenomena

56 D. BRILL, V. AYRES, AND G. GAUNAURD

would not exist. ““Analytic functions with-

out singularities must be constants,” states

a well-known theorem of analysis,’ and

eternally constant behavior is hardly an

interesting state of affairs. Here, we are

concerned with resonances, which are

singularities (or, mathematically, division

by zero) that serve to define the physical

characteristics.

Many physical processes of a wave-na-

ture contribute character to their sources

through their resonant content. For in-

stance, when we hear a familiar voice we

recognize the unique juxtaposition of

resonant qualities which serve to identify

the individual. We naturally store the ex-

perience of hearing a person’s voice for

later use in identification. The same sort

of recognition occurs using optical phe-

nomena where color provides a resonant

activity which serves to help distinguish

the source.

In scattering theory, singularities in the

scattering amplitudes of the waves re-

turned by objects are caused by their nat-

ural resonances. Each object has reso-

nances that are unique and which are

communicated, like a fingerprint, to its

echo. Once one deciphers the code about

shape and material composition that the

scatterer’s resonances impart to its echos,

it becomes possible to use them for char-

acterization purposes.

Resonances in physical processes have

a peculiar commonality and uniformity.

A recently developed theoretical tool, the

resonance scattering theory,’ (RST),

seems to possess a degree of universality

which makes it ideal for use as a general

investigative tool for the many physical

wave processes which exhibit resonance.

The details of the methodology have been

discussed in many technical?** and non-

technical? articles, but its essence may

be reviewed as follows. Whenever an in-

cident wave falls upon an object, the ob-

ject is invited to vibrate with all the fre-

quencies present in the incident wave. The

object will, however, naturally respond

by vibrating only with that peculiar subset

of frequencies which is also present in its

resonance spectrum. The RST consists of

a series of steps that ultimately isolate the

body’s resonances as they appear buried

within the active echo that the body re-

turns. The echo can be viewed as a char-

acteristic signature, in frequency or in

time, of the object that generated it. Thus,

RST is a potential target identification tool.

The first step of RST models the scatterer

as penetrable, with interior fields coupled

to outer fields through boundary condi-

tions at the object’s surface. The second

step requires an analysis of each one of

the modes or partial waves that form the

object’s summed cross section. The third

step separates the two basic contributions

contained within each mode. These are

in the form of smooth continous back-

grounds and discrete spiky resonances. The

backgrounds depend on the scatterer’s

shape, but assume it to be impenetrable.

The resonances are effects of the pen-

etrable composition of the body, and

they manifest themselves as discrete dips/

peaks superimposed on the smooth back-

grounds. They correspond to the pene-

tration of energy into the object which

only occurs through discrete spectral win-

dows centered at the object’s resonances.

For bodies with non-separable shapes, the

decomposition of the cross-section into

modal or partial wave portions is not pos-

sible, and the contribution from the back-

grounds and the resonances are sepa-

rated from each other in the complete or

summed cross-section instead of in each

of its partial waves.

It is essential to understand the differ-

ence that consideration of a body as pen-

etrable versus impenetrable makes. First,

one must remember that the Rayleigh re-

gion is where the wavelength is large com-

pared with the size of the target object,

and, hence, where the scattering process

is predominantly due to the phenomenon

of diffraction. At the other extreme of the

spectrum, the geometrical region, the

wavelength is much smaller than the ob-

ject’s size and here the scattering process

NATURAL RESONANCES a

[#294 (=) |

RAYLEIGH

REGION

3.00 4.00 8.00 1200 1600 20.00 24.00

|f a4 (=) |

RAYLEIGH

REGION

0.00 4.00 8.00 12.00 16.00 20.00 24.00

28.00 32.00 36.00 40.00 44.00 48.00

Fig. 1 (a) Form function of an impenetrable (rigid) sphere of radius ‘“‘a” in water is shown as a function

of ka (=27a/A = wa/c). Various scattering regions are shown on this spectral plot. (b) Form function of

an elastic (tungsten carbide) sphere in water. The resonance region widens.

is, for the most part, a result of refraction

and reflection. For an impenetrable body,

the resonance region separates the Ray-

leigh region from the geometrical regime.

(See Figure la.) For a penetrable body,

this is not the case. The resonance region,

above the Rayleigh region, propagates

out into the geometrical regime (see Fig-

ure 1b). The resonance region is there-

fore considerably enlarged. The impor-

tance of the Resonance Scattering Theory

for penetrable objects stems from recog-

nition of this fact.

Resonances appear everywhere in

physical processes which involve wave

phenomena. Typical examples are in areas

such as acoustics (sonar), ultrasonics/

elastodynamics, mechanics of solids,

electromagnetism (radar), geophysics,

optics, and nuclear physics. Complex res-

onance phenomena present in some of

these areas have been theoretically ex-

plained in the past. In other instances, the

explanation is still lacking. In all in-

stances, there has been little attempt at

unifying the treatment of resonances in

all of the above fields with a universal

approach.

In what follows, we will illustrate the

various areas with examples from each

one. This will form a collection of ex-

amples of quite dissimilar physical proc-

esses, that could all eventually be ex-

plained by this new methodology. In

Section I we shall point out the few cases

in which the methodology and the expla-

nation are already available; and in Sec-

tion II we call attention to the current

ongoing application of this method to new

situations.

58 D. BRILL, V. AYRES, AND G. GAUNAURD

I. Acoustic, Solid State, and

Electromagnetic Resonances—

A Common Bond

Let’s begin the development of our col-

lection with three examples from the dis-

ciplines of underwater and physical acous-

tics. When a sound wave emerging from

a distant source is incident on a sub-

merged cylindrical shell, the resulting

backscattering cross section is as shown

in Figure 2a, taken from Ref. 6, with res-

onance features clearly evident. In this

early study (1965) of small cylindrical

shells, the various types of resonances ob-

served in the backscattering cross sections

at various frequencies were categorized

according to their partial wave origin, an

RST-type idea. It was not until later, how-

ever, that a full RST analysis of a similar

problem,” a study of an air filled cylin-

drical aluminum shell in water, was per-

formed. A result from this study is shown

in Figure 2b, and what was previously

good guess work has now become clear.

The resonances which are only dependent

upon the cylinder’s composition have been

isolated here from the geometry or shape

dependent background. From plots such

as these we can reexamine the cross sec-

tion diagram and identify every one of its

features (Fig. 2c). Many theoretical pre-

dictions of RST have been confirmed ex-

perimentally.”

An even earlier (1951) but similar

study,® also foucused on acoustic reso-

nances. Here the bistatic (i.e., receiver

and source at different locations) scat-

tering cross section of a solid elastic cyl-

inder in water is shown versus the angle

6 between the incident and scattered beams

for three different values of frequency. It

was observed that when the frequency of

the incident sound wave matches one of

the resonant frequencies of the object, a

significant decrease occurs in the back-

scattered signal (0 = 180°).

Yet another RST analysis’ in 1979 has

furnished a clear picture of what is ac-

tually happening in this problem. The null

is caused by absoption of energy at a fre-

quency which corresponds to a natural

resonance of the object. These resonance

frequency values are given by the roots

of characteristic equations which occur

as a result of applying the RST. If we

choose one of these resonances which

cause a quasi-null and plot the form func-

tion or scattered wave amplitude versus

angle 8 we should expect to see a null in

the backscattering direction, @ = 180°,

and we do. Also, the near null at @ =

180° seen in Figure 3, will quickly be filled

in and disappear when it is redone for any

frequency value slightly away from the

resonance value (even 1% away).

Further study of acoustic resonances

appear in the analysis of a different type

of problem: that of wave propagation

through bubbly liquids. When a sound

wave travels through a cloud of gas bub-

bles in water, the wave exhibits an effec-

tive sound speed and effective attenuation

much larger than if the bubbles were not

present. This phenomenon has been

measured” and then predicted by RST"

with good agreement between the two. It

is shown that this behavior is completely

controlled by the acoustic resonances of

the individual bubbles, which taken to-

gether produce a cumulative effect. Anal-

ogous studies have been carried out for

viscoelastic solids containing perfora-

tions.''°

We continue the list with two examples

of ultrasonic resonances in solids.

A fluid filled cavity within a solid can

be excited by an incident elastic wave. Of

particular interest to those who work with

underwater sound absorbers is the case of

a spherical, air filled cavity in a rubber

matrix. Such a cavity was set into oscil-

lation by an incident compressional wave,

and the cavity’s pressure amplitude was

plotted versus frequency in Ref. (12). The

simple theory presented in Ref. (12) to

describe a vibration problem was later

generalized within the context of scatter-

ing, and in particular, of resonance scat-

tering,’ with the following results. The

cavity exhibits a big resonant peak which

is caused by its monopole mode of oscil-

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60 D. BRILL, V. AYRES, AND G. GAUNAURD

(5,2) RESONANCE, ka = 19.5429

Fig. 3 Bistatic form function versus 6 at a single

selected resonance frequency (viz, xs.) = 19.5429).

A Faran type® of minimum or dip at 6 = 180° is

clearly observed. This dip disappears for values of

x = k,a slightly away from the value of the (5,2)

resonance at 19.5429. (From Ref. 9).

lation (n=0), and which is quite similar

to that seen in bubbles” and in nuclear

physics.'* For a single cavity, the spectral

location of the peak changes with the cav-

ity contents and size, and with the matrix

composition.

Metal matrix composites have also been

investigated for resonant behavior. A

water filled cylindrical cavity contained

within an aluminum matrix, has been ex-

cited by an incident elastic compressional

MODES

RELATIVE AMPLITUDE

pulse. Its backscattered spectrum is plot-

ted versus frequency in Figure 4, which is

taken from Ref. (15). Oscillations in this

cross-sectional plot are due to the reso-

nances of the fluid in the cavity.

It should be noted that the literature in

this area is mostly concerned with mea-

surements and, thus far, the inherent

spectrum of resonances has only been

analyzed for the purpose of sizing the

cylinder’s diameter. In fact, a variety of

(largely experimental) studies of dif-

ferent types of elastic waves scattered by

inclusions within elastic media, such as

those in Refs. (12), (15), and (16), have

been completed in recent years, and still

await general theoretical treatments

which focus on the observed resonant be-

haviors.

Our next two examples come from a

seemingly very different area of physics—

electromagnetism. Resonance effects in

the radar cross-sections of conducting and

dielectric bodies are as abundant and fa-

miliar as they are for the acoustic case of

sonar cross-sections. The literature in this

area is quite extensive. For instance,

FREQUENCY (MHZ)

Fig. 4 Ultrasonic resonances. A compressional wave

incident upon a water-filled cylinder in an aluminum

medium is backscattered (8 = 7) into another com-

pressional wave. The form-function (f°?) for this

process is displayed versus frequency: (

), ex-

periment; (----), spectrum of the incident pulse. (From

Ref. 15) Various resonance modes are identified in

the displayed power spectrum.

NATURAL RESONANCES 61

10 log 19 (o/wa*)

25.13

0.0 12.57

kya

Fig. 5 Radar resonances. Radar cross-section (dB)

of a perfectly conducting sphere coated with a mask-

ing layer of dielectric material, in the (non-dimen-

sional) frequency band 0 = k,a = 25.13 (=8m).

Resonance oscillations are identified for various TE

and TM modes. (From Refs. 18 and 19).

backscattering from a perfectly conduct-

ing cylinder (of radius aj) coated with a

layer of dielectric material was studied in

Ref. 17, in 1957. The normalized cross-

section versus non-dimensional frequency

can be displayed for various thickness and

composition values in the dielectric coat-

ing layer. The resonance peaks shift with

thickness and composition changes in the

dielectric coating. A general theory was

developed which agrees with the experi-

mental observations, however, no fun-

damental analysis of the resonances was

performed here. In a similar study in 1964,

a conducting spherical core covered with

a layer of dielectric material was analyzed

by Rheinstein.'’* Its radar cross-section

(RCS) was obtained by classical means

without a fundamental study to sort out

the resonances present. A resonance scat-

tering analysis” of this particular work was

carried on later, in 1980 (see Fig. 5). It

showed how RST could be used in this

instance to: (a) identify the object, even

though it is coated, since the composition

of the dielectric layer can be extracted

from the wiggles in the RCS, in whichever

frequency band one has access to and, (b)

best mask the object or make its presence

less apparent in spite of the fact that it is

being illuminated since a well-chosen

coating reduces the cross-sectional peaks

by damping the material resonances. This

analysis has served to model the echoes

from a_ telecommunications _ satellite

coated with a dielectric layer. It could also

apply to any other large coated conduct-

ing object of spherical shape. Extensions

of this manner of treatment to other geo-

metrical shapes, coated or bare, are also

possible.

II. Some Noted Resonance Behaviors

In all of the diverse physical processes

described above, natural resonances oc-

cur. Consequently, there is acommon bond

between them since all could be analyzed

using RST. This would yield important

information about their shape and ma-

terial properties. Other examples come to

mind, all of which display notable reso-

nance features, and for which a funda-

mental, unifying analysis has yet to be

performed. We will briefly discuss some

of these “‘future possibilities” here.

Wave propagation through the Earth—

in an earthquake, for example—imme-

diately brings out the resonances of the

Earth itself. In Geophysics, the Earth is

usually modelled as a large layered, elas-

tic sphere. Even without the layering, the

analysis is quite complicated and it leads

to the so-called radial and spheroidal vi-

brations.”” More recent works’! have

analyzed and displayed theses vibrational

spectra from novel viewpoints. The graphs

in Ref. 21 show some of the earth’s radial

and spheroidal eigenfrequencies in the

MHz region, caused by various types of

surface tremors. Resonance features are

clearly evident in the graphs presented in

these papers. Some of these features were

examined with RST in Ref. 22. This work

continues.

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NATURAL RESONANCES 63

Another more localized geophysical

problem exhibits resonance features with

a growing RST connection. When a sound

wave is incident on a layered ocean bot-

tom, the reflection coefficient exhibits

certain losses that depend on the angle

that the incident pulse makes with the sea-

bed.* The angle dependence of this bot-

tom-loss exhibits a marked resonant be-

havior. Some of these features have been

analyzed with RST in Ref. 24. This work

studies lossy multilayered configurations,

analogous to, but not quite the same as,

stratified ocean bottoms. Thus, the RST-

analysis still has to be extended to yield

a more accurate model for ocean bottom

reflections.

Resonance scattering of light is useful,

among other applications, for the high

resolution sizing of liquid droplets in aer-

osols or in clouds.” Existing techniques

can reach size resolutions of one part in

10°. Investigations have been made by

shining laser light at 90° on individual

droplets and polystyrene spheres in the

6.5 to 11.5 wm diameter range. Levitated

single droplets, evaporated and non-

evaporated, have been studied, as well as

spherical and non-spherical single or mul-

tiple droplets. The unprecedented size

resolution resulting from this method finds

application in the measurement of diffu-

sion coefficients and in studies of areosol

growth and evaporation.

Figure 6, taken from Ref. 25, displays

spectral resonances useful for particle siz-

ing, and compares prediction with obser-

vation which appear to agree. Although

these resonances have been analyzed, the

emphasis has been on the determination

of the sphere’s size. The problem remains

to extract information on size distribu-

tions, concentrations and overall material

properties from the structure of reso-

nances in the scattered returns. This seems

to be a task perfectly suited for RST.

Nuclear resonance reactions have re-

ceived considerable attention since the

early work of Breit and Wigner.*° Of par-

ticular interest, is the photonuclear giant

resonance in Si**. Plots of the yield from

the reactions Al’’ (p,y) Si*® and Si?§ (y,n)

Si’’ are shown in Figure 7 and taken from

Ref. 14. The cross-sectional plots contain

the rapid oscillations typical of resonant

behavior. Additional evidence of reso-

nant nuclear behavior was analyzed in

C” + C” scattering studies carried out in

1980 (Ret127).

All these articles display cross-sec-

tional plots (versus energy) as predicted

by theory* and as observed in measure-

ments.’’ The general connection between

the resonant behavior in these plots and

those of analogous disciplines, however,

remains unexploited and largely unex-

plored.

It is interesting to note that the original

development of the RST was in fact an

extension of Breit and Wigner’s work in

quantum theory to a classical scattering

situation. Perhaps we can bring these ideas

around full circle and find new insights

into the problems of nuclear physics from

6 Ai”\p.y 0) Si”

= 90° YIELD

si y,n)Si”

b 5

16 18 20 22 24

Ey,MeV

Fig. 7 Nuclear resonances. Top: The reaction dif-

ferential cross section for ground state energy pho-

tons emitted at 90° to the incident protons direction

is plotted versus proton energy. The cross section

units are microbarns (10° *° cm?) per unit solid scat-

tering angle (steradians). Bottom: Total reaction

cross section in microbarns (10~”’ cm?) as a func-

tion of the incident photon’s energy.

64 D. BRILL, V. AYRES, AND G. GAUNAURD

a technique that began there, by studying

a properly chosen acoustic or electro-

magnetic analogue.

Iii. Future Paths

With so many possible applications being

investigated, research into resonance

scattering is growing ever more exciting.

Advances in one field are quickly trans-

ferable to another, since the governing

equations and the general methodology

have so many similarities. One can see

that advances in underwater acoustics can

be carried over to areas such as nuclear

physics—the field first investigated with

a resonance expansion. Extensions of the

theory appear constantly with the partic-

ular and peculiar details of each appli-

cation requiring individual and often con-

siderable effort.

Scattering processes in the resonance

region are certainly the hardest to study

since in this region one cannot take ad-

vantage of any low or high frequency

asymptotic approximation. Exact solu-

tions developed for this region can be ex-

tended to the far limits of the respective

spectra to yield novel ways of looking at

resonance processes everywhere. The ul-

timate goal is a broadly useful analytical

method which is both informative and es-

thetically pleasing. This should result in a

macroscopic spectroscopy applicable to

objects of all sizes interacting with all types

of radiation in all conceivable spectral

bands. Also, there is now hope that the

inverse scattering process will be ame-

nable to interpretation and solution.

Techniques for target identification and

active signature analysis will prosper. We

look forward to sharing this excitement

with the many professionals in diverse

fields who serve to benefit from this com-

mon goal.

References Cited

1. G. Carrier, M. Krock and C. Pearson, Func-

tions of a Complex Variable, McGraw-Hill Book

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isliar

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the resonances of viscoelastic and electromag-

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Acoustics XV, pp. 191-294, Ch 3, W. P. Mason

and R. N. Thurston, Editors, (1981).

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can identify targets,’ Microwave Systems

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excited compressional ring modes of model

submarines,” Navy Electronics Lab (currently

NOSC), Report NEL-TR-1273, 17 March

1965, (Unclas.), San Diego, CA.

G. Gaunaurd and D. Brill, ‘“Acoustic spectro-

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Amer. 75, 1680-1693, (1984).

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experimental verification,’ J. Acoust. Soc.

Amer. 80, 382-390, (1986).

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spheres,” J. Acoust. Soc. Amer. 23, 405-418,

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in bubbly mixtures in standing wave tubes,” J.

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theory of bubbly liquids,” J. Acoust. Soc.

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and cavity-size distribution effects on the dy-

namic effective properties of perforated elas-

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(1983).

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nances in the scattering of waves from gas-

filled spherical cavities and bubbles,” J.

Acoust. Soc. Amer. 65, 573-594, (1979).

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inant state for the El giant resonance,” Phys.

Rev. Lett. 13 (#21), 628-631, (1964).

Y. H. Pao and W. Sachse, “Interpretation of

time records and power spectra of scattered

ultrasonic pulses in solids,” J. Acoust. Soc.

Amer. 56, 1478-1486, (1974).

NATURAL RESONANCES 65

16. E. R. Cohen and B. R. Tittmann, “Analysis

of ultrasonic wave scattering for characteri-

zation of defects in solids,’ Rockwell Inter-

national, Science Center, Tech. Report SCTR-

fale. Dec. (1975).

17. C. C. H. Tang, “Backscattering from dielec-

tric-coated infinite cylindrical obstacles,” J.

Appl. Phys. 28, 628-633, (1957).

18. J. Rheinstein, “Scattering of electromagnetic

waves from dielectric coated conducting

spheres,” IEEE Trans. A P-12, 334-340, (1969).

19. G. Gaunaurd and H. Uberall, “Electromag-

netic spectral determination of the material

composition of penetrable radar targets,” Na-

ture 287, 708-709, (1980). (Also U.S. Patent

No. 4,415,898 was granted Nov. 15, 1983.)

20. A. E. H. Love, A Treatise on the Mathematical

Theory of Elasticity, (Dover, New York, 1944)

4th Ed., Ch. 12, p. 284.

21. F. Gilbert and A. M. Dziewonski, “‘An appli-

cation of normal mode theory to the retrieval

of structural parameters and source mechanics

from seismic spectra,” Philos. Trans. Roy. Soc.

(London) A, 278, 24-269, (1975).

22. G. Gaunaurd and H. Uberall, ““RST-analysis

of monostatic and bistatic echoes from an elas-

Biographical Sketches

Donald Brill is a professor of Physics

at the U.S. Naval Academy. Since his

graduation from Catholic University

(Ph.D., 1971) his reasearch activity has

centered around acoustic scattering. He

is a Sigma Xi member.

Virginia Ayres has been a research phy-

tic sphere,” J. Acoust. Soc. Amer. 73, 1-12,

(1983).

23. E. H. Morris, ‘““‘Bottom reflection loss model

with velocity gradient,” J. Acoust. Soc. Amer.

48, 1198-1202, (1970).

24. P. D. Jackins and G. Gaunaurd, “‘Resonance

reflection of acoustic waves by a perforated

bilaminar coating model,” J. Acoust. Soc.

Amer. 73, 1456-1463, (1983).

25. T. R. Lettieri, W. D. Jenkins, ““A review of

ultra high resolution and sizing of single drop-

lets by Resonance Light Scattering,’ Amer.

Soc. for Testing and Materials, Special Tech.

Publication 848, pp. 98-108, 1984 (J. M. Tish-

koff, R. D. Ingebo and J. B. Kennedy, Edi-

tors).

26. G. Breit and E. Wigner, “Capture of slow neu-

trons,” Phys. Rev. 49, 519-531, (1936).

27. K. A. Erb, et al., “Resonant and average be-

havior of the C’+C” total reaction cross sec-

tion: 5.6)= E =10:0 MeV.) Phys: Rev, GC;

22, (#2), 507-514, (1980).

28. L. L. Hill, ““Application of the particle-hole

model to electron excitation and muon capture

in light nuclei,” NOL Tech. Report -67 -88,

131 pp., 24 May 1967 (Unclas.).

sicist at NSWC since her graduation from

Purdue University (Ph.D., 1985). She is

a member of Sigma Si.

Guillermo Gaunaurd has been a re-

search physicist at NSWC since his grad-

uation from Catholic University (Ph.D.,

1971). He leads a team engaged in all areas

of classical radiation and scattering the-

ory. He is a Sigma Xi member.

Journal of the Washington Academy of Sciences,

Volume 77, Number 2, Pages 66-75, June 1987

Fatty Acid Synthase: A Protein

Having Many Catalytically

Active Domains

Soma Kumar

Department of Chemistry, Georgetown University, Washington, DC

20057

ABSTRACT

Fatty acid synthesis requires the participation of a protein, the acyl carrier protein

(ACP) and six or seven different enzymes. In prokaryotes these proteins are separable

and monofunctional. Such an organization of the proteins is called Type II. In animal

tissues and yeast, the same enzyme activities are located on only one or two peptide

chains. These polyfunctional enzyme system is referred to as Type I. Plants have retained

Type II enzyme systems which are confined to plastid-related organelles. This lends sup-

port to the bacterial origin of these organelles.

Type I fatty acid synthase is believed to have originated by the fusion of the ancestral

genes encoding Type II enzymes and ACP. A comparison of the distribution of the various

enzymatic loci in yeast and vertebrate synthase suggests that the diversion during the

course of their evolution occurred by independent fusion.

Living organisms are able to perform

incredible feats of chemical activities re-

quired for their survival because these ac-

tivities are mediated by enzymes which

are extremely efficient catalysts. The ef-

ficiency of these catalysts is manifest in

their increasing the rates of chemical re-

actions several thousand- or even million-

fold, and in exhibiting extreme specifici-

ties with regard to their substrates and the

reactions they catalyze. Hence, virtually

no product other than the desired ones

are obtained during the course of the re-

action. The metabolic pathways that con-

stitute the catabolic and anabolic reac-

tions involve a reaction sequence in which

a starting precursor molecule undergoes

a sequence of reactions in which the prod-

uct of the first reaction becomes the sub-

strate of the second reaction and the prod-

uct of the second reaction the substrate

of the third reaction and so on until the

desired final product is obtained. At each

step a single chemical event occurs. This

might be the addition of a particular group

or the removal of one, cleavage or the

formation of one specific bond or a simple

rearrangment of atoms or groups of atoms

within the molecule. Underlying the com-

plex network of chemical reactions, con-

stituting the various reaction sequences,

shown in the metabolic charts adorning a

biochemist’s office, there is a correspond-

ing network of enzymes. The general rule

FATTY ACID SYNTHASE 67

|a>

Ey ES Ez Final

: ns

— mm Product

Fig. 1A. Metabolic reaction sequences in which the concerned enzymes (E,, E>, E;) are separate entities.

©)

OV. AYO

Fig. 1B. Schematic representation of (A) the soluble, separable, unaggregated enzymes catalyzing the

reactions shown in Figure 1A; and (C) a multienzyme complex, of which pyruvate dehydrogenase (Figure

2) is an example, in which there is an aggregation of the separable enzymes.

NAD*

FADH,

Lip-S.]

Bape

Zane

[Lip-(SH).,

eooH aa

ay O=C-S-CoA

[TPP]

CoA-SH

Fig. 2. Mechanism of action of pyruvate dehydrogenase. Three enzymes are involved. They are pyruvate

dehydrogenase (E,), dihydrolipoyl transacetylase (E,) and dehydrolipoyl dehydrogenase (E;). The complex,

an aggregate of these enzymes, each containing several peptide chains, has a particle weight of several

million daltons.

68 SOMA KUMAR

is, ONe enzyme catalyzes only one reac-

tion. The oxidation of glucose to lactic

acid (glycolysis) occurs in ten steps and

requires ten different enzymes.

Soluble, Separable Enzyme System

In this system (Figure 1A) the product

of the first reaction, S,, can be presumed

to move about in the intracellular space

available until it encounters E,, the en-

zyme catalyzing the second reaction re-

sulting in S; which has to find E; (Figure

1B). The efficiency of the overall reaction

sequence under this system would depend

upon the catalytic efficiency (k,,,) of the

various enzymes and the steady state con-

centrations of all the intermediates. The

requirement for coenzymes and other co-

factors in some of the reactions is another

complication. Enzymes catalyzing the re-

actions of glycolytic and citric acid cycle

are examples of such a system.

Multienzyme Complexes

The efficiency of a reaction sequence

would be increased considerably if the en-

zymes catalyzing consecutive reactions are

aggregated, or associated, so that the

product of a particular reaction finds the

enzyme catalyzing the subsequent reac-

tion right next to the first enzyme. The

intermediates in such a system would be

transferred from one enzyme to the next,

“bucket brigade”’ fashion. In effect, this

increases the local concentration of each

of the intermediates immensely and

thereby increases greatly the efficiency of

the overall process. Such an arrangement

is found in several enzyme systems. One

of the most thoroughly studied is pyruvate

dehydrogenase which oxidizes pyruvate

according to the following equation:

This reaction occurs in five discrete steps

(Figure 2) and requires three different

enzymes and thiamine pyrophosphate

(TPP), enzyme-bound lipoic acid (Lip S,)

and enzyme-bound flavin-adenine dinu-

cleotide (FAD) as cofactors. E,, the de-

hydrogenase decarboxylates pyruvate, it

releases CO, but keeps the acyl product

formed covalently linked to TPP which

itself is bound to E;. The subsequent step

involves the transfer of the acyl group to

E,-bound lipoate which requires the re-

duction of lipoate and oxidation of the

acyl group. E,, then transfers the acetyl

group formed to CoA of the medium but

this leaves lipoate in the reduced state.

FAD containing E; oxidizes lipoate to en-

able it to accept another acyl group from

E, while NAD* of the medium is used by

E; to oxidize the reduced form of FAD

in order for it to function as an oxidant

of reduced lipoate. The point to notice is

that the different intermediates are trans-

ferred from the one enzyme to another

and are not released into the medium.

The three enzymes concerned while ex-

isting as a complex can be dissociated from

each other and can function independ-

ently of each other provided the appro-

priate substrates are made available.!

A variation of the association of pro-

teins catalyzing consecutive reactions is

the respiratory enzyme complex that

carries out electron transport with the

concomitant formation of ATP. In this

system, the aggregates of the different pro-

teins are held together by their being

embedded in the membranes of the mi-

tochondria. Being aggregates these en-

zymes are also separable but because of

their natural association with membranes

special handling is required.

CH: €— COOH = GoA sh NAD

<— Pyruvic acid

! <—— Acetyl-CoA

FATTY ACID SYNTHASE 69

An entirely different kind of complex

is that of aminoacyl-tRNA synthetase en-

zyme systems found in eukaryotic species.

Several synthetases, specific for as many

as seven to ten amino acids, occur as ag-

gregates.”1 These aggregates carry out

parallel reactions, rather than sequential

reactions, and the enzymes concerned ap-

pear to be functionally independent. The

biochemical significance of this type of

organization of enzymes is at present not

clear.

Enzymes with multicatalytic sites.

There are enzymes which catalyze more

than one chemical reaction and which,

therefore, must possess more than one

catalytic site. A good example is the DNA

polymerase that carries out the template-

directed DNA synthesis in the 5’ — 3’

direction but which also possesses exo-

nuclease activity for clearing the path and

for “proofreading and editing” mis-

matched bases to ensure fidelity of DNA

replication. The exonuclease activity re-

sides on the same peptide chain and in

the native, folded state must be near the

polymerization site.” The nuclease activ-

ity can be excised enzymatically.

Dual Organization of enzymes

catalyzing de novo fatty acid synthesis.

Fatty acid synthesis occurs according to

the following equation:

O Fee O

the synthetic pathway failed because, as

it was later realized, the acyl intermedi-

ates were covalently bound to a relatively

large-molecular protein.* The enzyme from

both yeast and pigeon liver, the two sources

of the enzyme in these studies, was very

specific for acetyl-CoA and malonyl-CoA

and exhibited little reaction with sus-

pected intermediates, like acetoacetyl-

CoA, butanoyl-CoA or hexanoyl-CoA.

These investigations also failed to resolve

the enzyme into smaller components with

partial activities.

Two strategies were adopted for the

elucidation of the reaction sequence. Ly-

nen and his coworkers, after having

obtained evidence that the acyl inter-

mediates were bound to the enzyme

via thiol ester bonds, employed acyl thiol

esters of N-acetylcysteamine as model sub-

strates.’ Their structures are shown in

Figure 3. 5

If relatively high concentrations of the

model substrates and the enzymes are

used it was possible to “‘cheat”’ the en-

zyme into binding the substrate at the ac-

tive site long enough for a specific reac-

tion to occur at a measurable rate. For

instance yeast synthase would reduce ace-

toacetyl-NAC (Figure 3A) and crotonyl-

NAC (Figure 3C) to 3-hydroxybutanoy]l-

NAc and butanoyl-NAc, respectively, in

the presence of NADPH and would de-

hydrate B-hydroxybutanoyl-NAc (Figure

1B) to crotonyl-NAc. With such sub-

strates only individual partial reaction

CH;C—S—CoA + 7 CH,—C—S—CoA + 14 NADPH + 14 H* —>

acetyl CoA malonyl-CoA

ie. Chl. CH, jCH «CH, CH. *€H> CH.

O77 fee a ee a ESO FS

GEL (He -acCHy ‘CH; “CHL CHSC, COOH

Palmitic Acid

Early attempts to isolate the intermedi-

ates in the process in order to elucidate

because the product would diffuse away

and its concentration would be too low

70 SOMA KUMAR

CH 3 CH 2 eR 2 an CH 3 CH 2 S-NAC

ae ss ont, pele cH, SS One

J O v" aA OH -

NAC

Acetoacetyl-N-acetyl cysteamine (NAC) 8 -hydroxybutanoy1-NAC

(A) (B)

ay wa

trans-crotonoyl-NAC

(C)

Fig. 3. Structures of model compounds of N-acetylcysteamine.

would occur and not the overall reaction phopantetheine is also the tail portion of

for the subsequent reaction. coenzyme A, the carrier of many of the

An alternate strategy employed by Va- reactive acyl groups in biological systems.

gelos and his group of workers was to The structure of the prosthetic group and

attempt to fractionate the enzyme system its relationships to the rest of the molecule

from a prokaryote, a simpler organism of coenzyme A is shown in Figure 4.

such as E.coli, into component enzymes. Following the discovery of ACP, all of

In this they were successful. The fatty acid the enzymes concerned were purified and

synthesis by this system was found to re- the reaction sequence was rapidly estab-

quire a heat and acid stable protein of lished. This is shown in Figure 5A. This

relatively low molecular weight (M, ~ sequence is similar to that deduced for

10,000) which bound the acyl intermedi- yeast enzyme using model substrates by

ates. Characterization of this protein, Lynen and coworkers.’ After the first

which was named acyl carrier protein round of six reactions requiring six en-

(ACP), revealed the presence of a pros- zymes the chain lengthening continues with

thetic group, 4’-phosphopantetheine, the addition of two-carbon units succes-

which was bound to ACP via anesterlink- sively, employing the enzymes in reac-

age to a specific serine residue.* 4’-Phos- _ tions 3-6 until palmitoyl-ACP is formed.

HO H

CH NH CH NH ae CH oa

VW ee

CHS i CH, f a ee

cH, ‘CH,

a ee)

Adenine

4'-Phosphopantetheine

Fig. 4. Structure of coenzyme A showing the 4'-phosphopantetheine prosthetic group of acyl carrier

protein.

FATTY ACID SYNTHASE 71

5.A.

CH S acetyl CH S

Ne Wahcon + ACP-SH Ne J ~rcr + CoA-SH

i transferase 5

10) 10)

Acety1-ACP

2 £ t salar sh

coo Va ae ae malonyl co, y ACP

+ ACP- aE + CoA-SH

Sa Ss ets ee Nea \s/

ransferase 2

Malony1-ACP

CH, Ss CORE i ACP condensing c @ iS

Sey + ea, SS Bet ay Bau AN + ACP-SH

Cc ACP CH Ss

2 enzyme f ji ACP

A | + €05

Acetoacety1-ACP

CH CH S B-ketoacyl CH CH S

3 2 + 3 2 +

+ NADPH + H + NADP

Ne No neo reductase No Ne nee

(@) H \ou O

8 -Hydroxybutanoy1-ACP

CH H B-hydroxyacyl CH CH S

3 ve 2 Wi x 3

No »: Sie dehydratase SA Soups: eee ’

u/ OH Oo fe)

trans-crotony1-ACP

CH, WX S A enoyl CH3 Tax S i

; + NADPH + H eS + NADP

pe NaN reductase Nat ~~ Zo

(@)

5 ACP

Butanoy1-ACP

Fig. 5A. Reaction sequence for the de novo synthesis of fatty acids. Successive reactions in the first

round of reactions is shown. Butanoyl-ACP formed condenses with a mol of malonyl-ACP (Reaction 3)

and reactions 4—6 are repeated. This process is repeated until the 16-carbon palmitic acid is formed.

5.B.

CH, S CH, Ss

ve +o 5 -Ssq —_—_-— + ACP-SH

aioe cond —_—_—_——_—_ a NewiaNe

Tl II cond

O 1@)

O O O

if | HI

(e CH, S Cc ~<a ACP

oe TOR + LEG BRON ee AL NIA eS +aNcOr

CH, S-Eoond CO, (e ACP CH, CH,

-- Eoond 2H

Fig. 5B. Mechanism of the condensing reaction. This reaction occurs in two steps. Acetyl (or a saturated

acyl) group is transferred by the enzyme to one of its own specific cysteinyl SH group. This is followed by

condensation with an ACP-bound malonyl group so that the B-ketoacyl group formed is now bound to

ACP via its pantetheinyl SH.

72 SOMA KUMAR

Such a dissociable enzyme system was

not found in yeast, avian and mammalian

tissues. Yeast enzyme had a molecular

weight of 2.1 x 10°. Fatty acid synthase

from various animal tissues were similar

in size with a molecular weight of about

500,000. The reaction sequence by which

the acyl chain is elongated is also similar

which indicates the requirement for the

same catalytic activities. These enzymes

also contained 4'-phosphopantetheine as

prosthetic group and its role in the bind-

ing of incoming malonyl goup and of the

reactive acyl intermediates were similar

to E. coli and yeast enzymes.

Classification of Fatty Acid Synthase.

Two distinct types of organization of

the enzymes constituting the fatty acid

synthesizing system is, thus, found in na-

ture. They have been called Type I and

Type II by Bloch.*® Type I consists of the

organization of the various enzymes into

a tight complex, not amenable to disso-

ciation into individual enzymes. This type

is found in animal and insect tissues, yeast

and certain bacteria. Fatty acid synthase,

Type II, characterized by the component

enzymes being separable, is found in pro-

karyotes, e.g., E. coli, and most bacteria.

It is interesting that plants, although eu-

karyotic in nature have Type II fatty acid

synthase.’ The exclusive location of the

enzymes in proplastids, plastids and chlo-

roplasts has been taken as evidence that

supports the prokaryotic origin of these

organelles and the symbiotic relationship

between them and the host cells. It would

appear that Type I synthase is cytosolic

and Type II is organellar in location. The

chloroplast containing green alga, Eu-

glena gracilis, is endowed with both types

of the enzyme system. Type I is present

in the cytoplasm and Type II in the chlo-

roplasts. It is interesting to note that Type

II enzyme system is induced when these

organisms are grown in light and is absent

when grown in the dark.® It appears highly

likely that Type II synthase in plants has

all the constituent enzymes aggregated in

a loosely-associated complex in the con-

fines of the organelles and that it is the

disruption of the organelles in the course

of the preparation of the enzymes, that

disaggregates the complex.’

Type I Fatty Acid Synthase.

The demonstration of the various par-

tial reactions by the enzyme complex iso-

lated from yeast and animal tissues and

the failure to dissociate the complex into

component enzymes led Lynen to pro-

pose a model, by analogy with pyruvate

dehydrogenase, in which ACP, with a 20

A long 4'-phosphopantetheine arm, was

pictured as constituting a hub around

which the various enzymes were arranged

in the order necessary to carry out the

reaction sequence.’ This is the model cur-

rently shown in all the textbooks. The

prosthetic group with the reactive acyl

group is pictured as swinging from the

catalytic site of one enzyme to the next

successively to complete a round of re-

actions. The saturated unsubstituted acyl

group formed after one round of reactions

is transferred to a specific cysteinyl-SH of

the condensing enzyme. The free SH group

of pantetheine then accepts a malonyl

group. Condensation occurs between the

acyl group at the cysteinyl site and the

malonyl group (Figure 5B), which is fol-

lowed by another round of reactions. This

process is repeated until chain lengthen-

ing ceases by a mechanism still not under-

stood.

Subunit compostion of Type I Synthase.

Efforts to dissociate the enzyme com-

plex from animal tissues was finally met

with partial success. The 500,000 molec-

ular weight species of the enzyme com-

plex yielded a species with 250,000 mo-

lecular weight in buffers of low ionic

strength.'?! These could be reassociated

with the restoration of full activities. So

convinced were the workers of the mul-

tienzyme nature of the complex that the

250 kDA species were considered sub-

FATTY ACID SYNTHASE 73

complexes, more loosely associated with

each other than the components of each

of the subcomplex. Gradually evidence

began to accumulate, however, that the

two 250 kDa species formed upon disso-

ciation of the synthase were identical and

consisted of a single peptide chain. This,

naturally led to the concept that the var-

ious catalytic activities, representing each

of the partial reactions, were distributed

on the same peptide chain.

Yeast fatty acid synthase, of molecular

weight, 2.1 x 10°, had an entirely differ-

ent kind of organization. It consists of two

different peptide chains and the various

partial activities were distributed on these

two chains. To complicate matters even

further there are six copies of each chain.”

The subunit composition of yeast and an-

imal synthase are represented as a,8, and

a, respectively.

Comparative Aspects of Fatty Acid Synthase,

Type I.

While in all essential features the fatty

acid synthase from various animal tissues

are very similar there are some striking

differences. 1) Acetyl-CoA was recog-

nized early as the specific primer of the

overall reaction. Higher homologues of

acetyl group yields poor reaction with

yeast and avian liver enzyme. The enzyme

from mammalian tissues was later found

to utilize butanoyl-CoA more efficiently

than acetyl-CoA.” 2) Starting with acetyl-

CoA, palmitic acid was the main product

with yeast and avian enzymes. The en-

zyme from mammalian sources, on the

other hand, produces a high proportion

of butanoic acid. This derailment of the

C, acid intermediate is attributed to the

lack of specificity of the acyl transfer re-

actions.'’ Butanoyl group leaves the pan-

tetheinyl SH group by being transferred

to CoASH. Butanoyl-CoA produced in

this manner, most probably, accounts for

the butanoic acid content of the butterfat

of ruminants. 3) A lack of specificity with

regard to the acyl group transfer by bo-

vine mammary synthase also accounts for

the high rates of reduction of acetoacetyl-

CoA and crotonyl-CoA observed with this

enzyme. This enabled us to characterize

each of the partial reactions using esters

of CoA at natural concentrations.!!4 Em-

ployment of model substrates was useful

to demonstrate the occurrence of a re-

action but not to characterize it. 4) Goose

uropygial fatty acid synthase is able to

form multibranched chain fatty acids from

acetyl-CoA as primer if methylmalonyl-

CoA is used for chain elongation.” 5) Yeast

enzyme transfers palmitoyl group upon

termination of chain elongation to coen-

zyme A.” Animal synthase, on the other

hand, hydrolyzes palmitoyl group from the

prosthetic group to form free palmitic acid.

This requires the presence of an addi-

tional enzyme, a thioesterase in the com-

plex.'° 6) There is a separate transferase

for acetyl and malonyl groups in the yeast

enzyme but the same transferase serves

to transfer both groups in the vertebrate

enzyme.”

Multicatalytic function of fatty acid synthase I.

The resistance of the 250 kDa subunit

of fatty acid synthase from animal tissues

to dissociate into smaller peptides, de-

spite treatment with chemicals that dis-

rupt all chemical interactions except the

covalent bonds, led to the recognition that

all the catalytic activities reside on the

same peptide chain. Such a concept and

the identical nature of the two peptide

chains finally gained acceptance. This im-

plied that the five or six different chemical

activities, necessary for the addition of

each 2-carbon unit, reside on different do-

mains of the relatively large peptide

chain.”

The creation of a functional map began

by the cleaving off of the thioesterase by

treatment with trypsin.’° A 29 kDa active

thioesterase was obtained which proved

to be the carboxyterminal end region of

both the chains. Using the approach of

limited proteolysis and employing differ-

ent enzymes Wakil and coworkers,” in

74 SOMA KUMAR

Jae

Ch Cys

Fig. 6. Functional map of fatty acid synthase of animal tissues. Each subunit is juxtaposed in a head-to-

tail manner. The domains representing the different activities are: acyl carrier protein shown with its

prosthetic group (ACP); condensing enzyme (CE, reaction 3); acetyl- and malonyl-transferases (AT re-

actions 1 and 2); B-ketoreductase, dehydratase and enoyl reductase (KR, DH and ER, reactions 4, 5 and

6 respectively). For the reaction numbers refer to Figure 4. The different domains are not drawn to scale.

Adapted from the results of Wakil et al’? and McCarthy and Hardie.”

Houston, Texas, and Hardie and col-

leagues’’ in Dundee, Scotland, have con-

structed a functional map of the synthase

identifying the various partial enzyme ac-

tivities. Both groups have arrived at sim-

ilar conclusions. The map is presented

schematically in Figure 6. The two sub-

units are held by non-covent forces in a

head-to-tail arrangement such that ACP

domain of one subunit is juxtaposed

against the condensing domain of the ad-

jacent subunit. This would facilitate the

condensing reaction (Figure 5B) occur-

ring at both ends of the enzyme molecule.

The formation of B-ketoacyl group is fol-

lowed by reactions 4, 5 and 6 sequentially

(Figure 5A) to complete one round of re-

actions. The details of the reactions and

the arrangement of the different domains

involved are yet to be determined.

Occurrence of fatty acid synthesis in living

organisms.

Fatty acid synthesis is an essential fea-

ture of all dividing cells. The de novo syn-

thesis provides palmitic acid from which

the various fatty acids of the membranous

lipids are derived. The fatty acid synthase

is absent, or is present in extremely low

levels, in fully transformed cells except

certain specialized cells. All vertebrates

are able to store excess food intake in the

form of fat (triacylglycerols) to be used

at times of need. Fatty acids for the for-

mation of this concentrated form of

‘stored’ energy occurs primarily in liver,

whence it is exported to the various site

for deposition. Besides liver, lactating

mammary gland is extremely active in the

synthesis of fatty acids and its conversion

to milkfat in order to enrich milk with a

high energy dietary product.

The level of hepatic fatty acid synthase

can be easily manipulated by dietary re-

gime. Starvation for just two days reduces

the synthase activity to barely detectable

levels. It has been established that the

enzyme is in fact degraded and not just

inactivated. Refeeding a starved animal

with a high carbohydrate diet results in a

surge in the synthesis of the enzyme,

reaching 3 to 4 times the normal level

within forty-eight hours.'* This feature has

been exploited to obtain large quantities

of the enzyme for purposes of purifica-

tion. Lactating mammary enzyme, on the

other hand, is less sensitive to dietary ma-

nipulation.

FATTY ACID SYNTHASE 75

Genetic Implications of the Multicatalytic

Function of Type I Synthase.

The presence of type II fatty acid syn-

thase in an organism necessitates the en-

coding of the various enzymes in different

genes, the expression of which will have

to be closely synchronized to ensure the

proper ratio of the enzymes. Type I sys-

tem has the advantage over Type II sys-

tem in that all the activities are encoded

in only one or two genes. Control of the

expression of the gene(s) is, presumably,

facilitated.

A comparison of the aminoacid se-

quence of ACP of various bacterial, plant

and animal origin shows striking homol-

ogy indicating a common ancestral gene

from which the ACP genes of modern

species evolved.'? Although data are still

more limited regarding the other enzymes

or domains they do suggest similar ho-

mologies.'? These observations constitute

the basis for the current hypothesis that

the respective enzymes of the system have

also common ancestors and which were

monofunctional (Type II). During the ev-

olution of fungi and animals, fusion of

genes occurred to form polyfunctional

peptides. The encoding of the different

proteins in two genes in the fungi and one

gene in the animal organisms suggest that

a divergence occurred during their evo-

lution by two independent processes of

gene fusion rather than by one consecu-

tive process.*’ The order of the arrange-

ment of the loci of the different domain

in the two subunits of yeast synthase sup-

ports this concept.

References Cited

1. L. J. Reed (1974) Multienzyme Complexes.

Accts. Chem. Res. 7, 40—46.

2. A. Kornberg (1974) DNA Synthesis, pp 67-121.

W. H. Freeman & Co., Publishers.

3. F. Lynen (1961) Biosynthesis of saturated fatty

acids. Fed eration Proc. 20, 941-951.

4. Vagelos, P. R., P. W. Majerus, A. W. Alberts,

A. R. Larrabee and G. P. Ailhaud (1966) Struc-

ture and function of the acyl carrier protein.

Federation Proc. 25, 1485-1494.

5. Volpe, J. J. and P. R. Vagelos (1973) Saturated

fatty acid biosynthesis and its regulation. Annu.

Rev. Biochem. 42, 21-60.

6. Bloch, K. and D. Vance (1977) Control mech-

10.

12.

15:

14.

15.

16.

1 fe

18.

19:

20.

anisms in the synthesis of saturated fatty acids.

Annu. Rev. Biochem. 46, 263-298.

. J. B. Ohlrogge (1982) Fatty acid synthetase:

plants and bacteria have similar organization.

Trends Biochem. Sci. 7, 387-388.

. A. J. Fulco (1983) Fatty acid metabolism in

bacteria. Prog. Lipid Res. 22, 133-160.

. Lynen, F., D. Oesterhelt, E. Schweizer and K.

Willecke (1968) The viosynthesis of fatty acids

in Cellular Compartmentalization and control of

fatty acid metabolism, pp 1-24 (F. C. Gran, ed)

Academic Press.

Kumar, S., R. A. Muesing and J. W. Porter

(1972) Conformational changes, inactivation and

dissociation of pigeon liver fatty acid synthetase

complex. J. Biol. Chem. 247, 4749-4762.

Dodds, P. F., M. G. F. Guzman, S. C. Chalberg,

G. J. Anderson and S. Kumar (1981) Aceto-

acetyl-CoA reductase activity of lactating bo-

vine mammary fatty acid synthase. J. Biol. Chem.

256, 6282-6290.

Wakil, S. J. and J. K. Stoops (1983) Structure

and mechanism of fatty acid synthetase. in The

Enzymes, Third Edition, Vol. XVI, pp 3-61 (P.

D. Boyer, ed) Academic Press.

Abdinejad, A., A. M. Fisher and §. Kumar

(1981) Arch. Biochem. Biophys. 208, 135-145.

Ghayourmanesh, S. and S. Kumar (1981) Syn-

thesis of acetoacetyl-CoA by bovine mammary

fatty acid synthase. FEBS Lett. 132, 231-234.

Buckner, J. S. and P. E. Kolattukudy (1975)

Lipid biosynthesis in the sebaceous glands: syn-

thesis of multibranched fatty acids from meth-

ylmalonyl-coenzyme A in cell free preparations

from the uropygial gland of goose. Biochemistry

14, 1774-1782.

Smith, S., E. Agradi, L. Libertini and K. N.

Dileepan (1976) Specific release of the thioes-

terase component of fatty acid synthetase mul-

tienzyme complex by limited trypsinization. Proc

Natl. Acad. Sci. 73, 1184-1188.

McCarthy, A. D. and D. G. Hardie (1983) The

multifunctional polypeptide chains of rabbit-

mammary fatty-acid synthase. Stoichiometry of

active sites and active-site mapping using limited

proteolysis. Eur. J. Biochem. 130, 185-193.

A. G. Goodridge (1986) Regulation of the gene

for fatty acid synthase. Federation Proc. 45, 2399-

2405.

McCarthy, A. D. and D. G. Hardie (1984) Fatty

acid synthase—an example of protein evolution

by gene fusion. Trends Biochem. Sci. 9, 60-63.

Hardie, D. G., A. D. McCarthy and M. Brad-

dock (1986) Mammalian fatty acid synthase: a

chimeric protein which has evolved by gene fu-

sion. Biochem. Soc. Transactions 14, 568-570.

Yang, D. C. H., J. V. Garcia, Y. D. Johnson

and S. Wahab (1985) Multienzyme Complexes

of Mammalian Aminoacyl-tRNA Synthetases.

Current Topics in Cellular Regulation 26, 325-—

335:

Journal of the Washington Academy of Sciences,

Volume 77, Number 2, Pages 76-80, June 1987

Dental Alteration in

Prehistoric Ecuador

A New Example

from Jama-coaque

Douglas H. Ubelaker

‘Department of Anthropology, Smithsonian Institution,

Washington, D.C.

ABSTRACT

A recently discovered prehistoric skull of Jama-coaque culture from Manabi, Ecuador

shows a pattern of incised lines on seven of the anterior maxillary teeth. This example of

dental incision is unique in South America and has been documented previously in the

New World only in Mexico.

The practice of dental alteration in

the prehistoric New World includes tech-

niques of drilling, filing, chipping, pro-

ducing designs ranging from filed points

and incised patterns to drilled holes con-

taining stone insets. Romero! has docu-

mented over 127 types, mostly originat-

ing in Classic and Post-Classic Mexico.

In South America, dental mutilation by

chipping occurs only in historic times, and

appears to have been introduced by

Blacks. Prehistoric dental mutilation in

South America outside of Ecuador con-

sists only of filing, with published exam-

ples of teeth filed to points from Argen-

tina, Bolivia, and Chile.’ In prehistoric

Ecuador, mutilation by filing also oc-

curred, but in addition, examples of den-

tal inlay have been reported that closely

resemble those reported from Mexico. As

early as 1909, Saville called attention to

the early Spanish accounts of Zarate and

Gomara which mention that when they

explored the area of Atacames on the north

coast of Ecuador, they observed Indians

with “‘their faces sown with gold nails.’”

Saville’ also reported skulls found in the

Atacames area with small disks of gold

set into circular drilled holes in the an-

terior teeth and one skull with wide plates

of gold inset into the labial surfaces of the

maxillary teeth.

In 1912, Joyce’ illustrated a skull from

the Atacames collection of the British

Museum which shows drilled circular per-

forations in seven anterior maxillary teeth,

with some still containing gold insets.

Estrada’ indicates that skulls with

“‘dientes con oro” were recovered from

Elisita and La Tolita, but provides no de-

tailed descriptions.

Evans and Meggers°® provided illustra-

DENTAL ALTERATION IN PREHISTORIC ECUADOR 77

tions and descriptions of examples of den-

tal decoration from the sites of Elisita (G-

M-5, Guayas Province), Atacames, and

La Piedra on the Rio Esmeraldas in Es-

meraldas Province. They note that the

form of some of the dental inlays is similar

to those reported from Mexico, especially

from the state of Oaxaca. An important

difference is that the Mexican inlays were

mostly of pyrites, jadeite, or turquoise,

while those from Ecuador are all of gold.

They argue that the custom suggests con-

tact between Ecuador and Mexico some-

time during the Integration period, not

earlier than A.D. 500-600. This compar-

ison was made earlier by Saville’ and by

Stewart” who traced the form of dental

mutilation to the Mayan area.

Carlos Zevallos Menendez’ provides a

summary of dental mutilation in Ecuador.

Within this article he reports additional

examples of isolated teeth with gold insets

recovered during his excavations in the

Chanduy Valley. He also describes a man-

dible excavated by Jorge Marcos at the

site Congrejitos that displays small gold

insets in the incisors, canines and pre-

molars. A photograph of the mandible in

site is provided by Marcos.*®

All of the examples mentioned above

were found in skulls originating from

young adults who had died at a young age,

before much of the crown surface was lost

through attrition. An exception was re-

ported by Ubelaker’ from the Guayas

Province site of La Compania on the Rio

Babahoyo. Five anterior maxillary teeth

were found in a mound urn burial dating

from perhaps the 16th century. The teeth

originate from an older adult, in which

attrition has destroyed most of the crown

surface. Each tooth displayed a single cir-

cular perforation at the junction of the

crown and root, probably for the purpose

of displaying inlays. The burial also con-

tained extensive exotic artifacts suggest-

ing the individual enjoyed considerable

status.

In 1985, an articulated skull and man-

dible were brought to the attention of Olaf

Holm, Director of the Museo Antropol-

ogico, Banco Central del Ecuador. The

material had been found in the culture

area of Jama-coaque near Manabi and is

estimated to date between 200 B.C. and

800 A.D. The material was delivered to

the Museo with the skull and mandible

articulated and still embedded in the soil

matrix. Several of the anterior teeth ap-

parently had fallen out and had been rear-

ticulated with adhesive to the approxi-

mate original locations, but in incorrect

anatomical order (Figure 1).

On July 10, 1986, at the request of the

Museo, the author carefully removed the

skeletal remains from the soil matrix,

cleaned all skeletal parts and teeth, and

subsequently reconstructed them as much

as possible. Analysis revealed a nearly

complete skull and mandible of a young

adult (Figure 2). Most cranial vault su-

tures were open and dental attrition was

minimal, suggesting an age at death of

between 28 and 34 years. The occipital

shows marked flattening, strongly sug-

gesting intentional cranial deformation.

All teeth were present except the maxil-

lary left premolars, and the mandibular

Fig. 1. Jama-coaque skull and mandible with in-

cised teeth prior to cleaning and reconstruction.

78 DOUGLAS H. UBELAKER

Fig. 2. Jama-coaque skull and mandible after cleaning and reconstruction.

right canine and first premolar. None of

the teeth present had carious lesions and

deposits of calculus were absent on some

teeth, but heavy on others.

Male sex is suggested by the presence

of moderately large supra orbital ridges,

deep grooves behind the mastoid proc-

esses and a mandibular gonial angle ap-

proaching 90°.

The only evidence of disease consists

of porosity on the superior interior sur-

face of both orbits. This “‘cribra orbitale”

suggests the individual may have suffered

from anemia.

DENTAL ALTERATION IN PREHISTORIC ECUADOR 79

Fig. 3. Incised maxillary teeth from Jama-coaque skull.

Seven of the anterior teeth display a

series of incised lines on their buccal

crown surfaces. The maxillary canines and

incisors all show a series of intersecting

incised lines of a cross hatch design (Fig-

ure 3). The right central mandibular in-

cisor shows a single diagonal incised line

on its buccal crown surface. All of the

designs would have been visible through

the mouth and clearly were made during

life for aesthetic purposes. The anterior

mandibular teeth show more advanced at-

trition than those of the maxilla. Thus, it

is possible that the lower teeth may once

have had more incisions, but lost them

through attrition.

This type of dental modification is pre-

viously unknown in Ecuador and to the

best of the author’s knowledge, in South

America as well. The design is very sim-

ilar to a single example reported from

Mexico by Romero.’”!! The Mexican skull

is a male from ‘“‘entierro 17 de Xaloztoc,

Edo. de Mexico, Preclasico Superior.”

Romero’s”’ brief description is as follows:

‘Casi todo la extension de la cara anterior

se encuentra ocupada por una serie de

lineas entrecruzadas. Existe en incisivos

laterales y caninos superiores, en com-

binacion con la forma anterior en los cen-

trales -y . (Bicure 4):

Fig. 4. Incised maxillary teeth from Preclassic skull

from Xaloztoc, Mexico (from Romero, 1958: 45).

80 DOUGLAS H. UBELAKER

This newly discovered dental alteration

suggests that methods of dental decora-

tion and alteration in prehistoric Ecuador

were even more varied than previously

thought. The incised teeth also add yet

another cultural trait shared by prehis-

toric peoples of Ecuador and Meso Amer-

ica.

References Cited

1. Romero, Javier. 1974. La Mutilacion Dentaria,

pp. 230-250 in Antropologia fisica, Epoca Pre-

hispanica. Hidalgo: La Casa Chata.

2. Stewart, T.D. 1950. Deformity, Trephining, and

Mutilation in South American Indian Skeletal

Remains. Pp. 43-52 in Handbook of South

American Indians, Vol. 6, Physical Anthropol-

ogy, Linguistics and Cultural Geography of

South American Indians, Julian H. Steward,

Editor, Washington: G.P.O.

3. Saville, Marshall. 1909. Archaeological Re-

searches on the Coast of Esmeraldas, Ecuador,

Verhandlungen des XVI. _ Internationalen

Amerikanisten-Kongresses, Wien, 1908, pp.

331-345.

4. Joyce, T.A. 1912. A Short Guide to the Amer-

ican Antiquities in the British Museum, Uni-

versity Press, Oxford,

10.

. Estrada, Emilio. 1957. Ultimos Civilizaciones

Pre-Historicas de la Cuenca del Rio Guayas.

Publication del Museo Victor Emilio Estrada.

No. 2. Guayaquil.

. Evans, Clifford and Betty J. Meggers. 1966.

Meso America and Ecuador. Chapter 12, pp.

243-264 in Handbook of Middle American In-

dians, Volume 4, Gordon F. Ekholm and Gor-

don R. Willey, editors. Austin: University of

Texas Press.

. Zevallos Menendez, Carlos. 1982. La Mutila-

cion Dentaria en el Antiguo Ecuador. Chapter

2 of Seccion II pp. 233-257. Primer Simposio

de Correlaciones Antropologicas Andino-Meso

America. Escuela Superior Politecnica del Li-

toral. Guayaquil.

. Marcos, Jorge G. 1982. Arqueologia de la Pen-

insula de Santa Elena (11). Espejo. No. 6, pp.

123-128. Quito.

. Ubelaker, Douglas H. 1977. Drilled Human

Teeth from the Coast of Ecuador. Journal of

the Washington Academy of Sciences, Vol. 67,

No. 2, pp. 83-85.

Romero, Javier. 1958. Mutilaciones Dentarias

Prehispanicas de Mexico y America en General.

Serie Investigationes, Inst. Nac. Antr. Hist. Dir.

Invest. Antr. No. 3. Mexico.

Romero, Javier. 1970. Dental Mutilation, Tre-

phination, and Cranial Deformation. Chapter

4, pp. 50-67 in Handbook of Middle American

Indians, Vol. 9, Physical Anthropology, T.D.

Stewart, Editor. Austin: University of Texas

Press.

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