ATLAS
of
PROTEIN
SEQUENCE
and
STRUCTURE
1965 zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Margaret
0.
Dayhoff
Richard
V.
Eck
Marie
A.
Chang
Minnie
R.
Sochard
NATIONAL
BIOMEDICAL
RESEARCH
FOUNDATION
8600
16~~
STREET zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Silw
Spring,
Ma
yhnd
ATLAS
OF
PROTEIN SEQUENCE AND STRUCTURE zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
1965
Library
of
Congress Card Catalogue Number
65-29342 zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
@
Copyright
1965
by
The National Biomedical Research Foundation
Extra copies may be purchased from:
The National Biomedical Research Foundation
8600 16th Street, Silver Spring, Maryland
DEDICATION zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
To zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
all
the investigators who have developed the techniques
necessary for the grand accomplishments represented by this tabu-
lation, and to
all
those who have spent zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
so
much tedious effort in
their application.
We
would gratefully appreciate receiving suggestions, correc-
tions, new data (even
if
fragmentary or provisional), and
refer-
ences to any data omitted from this volume.
M. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
0.
D.
R.
V.
E.
M.
A. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
C.
M.
R.
S.
.001 zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
PREFACE
This
Atlas
voluminously illustrates the triumph of experimental technique over
the secretiveness of nature. Perhaps nowhere has the power of the scientific method
been more brilliantly demonstrated than in the development of procedures for the
study of the chemistry of life.
As
recently
as
twenty years ago,
it
was customary for
biologists
to
have
a
hopeless attitude about biochemistry. Some details might be
elicited, perhaps, but living things were thought to be
so
very complex and intricate
that
there surely was no hope of fully "understanding" them in
all
their chemical de-
tail.
Who,
if
he really comprehended the difficulty of the problem, would dare
to
think
'of man's ever knowing the detailed structure of
a
protein, for example, much
less
be
able
to synthesize
it?
Who
would ever understand the mechanism of
an
enzyme
as
clearly
as
a
chemist understands the details of an inorganic reaction? How could we
ever hope to know the atomic details of
a
protein crystal zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
?
Today some of these ambitions have already been attained, and the others no
longer seem out of reach.
We
now rationally hope to be able to discover and under-
stand the finest chemical details of living processes.
These accomplishments and
hopes have been made possible by the combined
effect
of several new approaches.
Techniques which permit the separation of chemically similar compounds have
been developed for microgram samples.
Among these
are
ion-exchange columns,
paper chromatography, electrophoresis, and counter-current distribution. Radio-
active
tracer
techniques and other micro-quantitative analytical procedures, often de-
pendent on electronics and automation, have aided the 'analyses. X-ray crystallography,
starting with
the
art
of protein crystal production and ending with the processing of
great numbers of experimental observations in the high-speed computer, has permitted
a
glimpse of three-dimensional structure.
Confidence in our understanding of experimental procedures and relationships
among proteins
has
grown
so
great that sequences of amino acids
are
inferred from
those found in homologous proteins. This technique requires only
a
small proportion
of
the analytical work needed to sequence
a
protein with no known relatives.
The ef-
fectiveness of laboratory effort
is
thus magnified.
Some
of
the insights which have been developed cannot be attributed to any partic-
ular
worker
or
school. Perhaps the greatest
of
these insights
is
that nature always
uses "building blocks."
A
living cell
is
extremely complex and almost unimaginably
intricate in detail.
But
it
consists of
a
limited, understandable number of types of
processes, reduplicated with variations.
To understand the cell,
we
must have
a
few
examples of each
type
of process, from which we can see the overall principles. For
understanding,
we
need not work out the details of
all
the variations on these princi-
ples, although we may eventually choose to
do
so
for medical
or
other practical
rea-
sons. Similarly, the analysis of such large, complex chemical molecules
as
proteins
has been made possible by the recognition of their essential modularity, their building-
block nature. Proteins
are
precise chemical structures built from
regular
subunits,