1944 Studies on the Chemical Nature of the

Substance Inducing Transformation of

Pneumococcal Types. Induction of
Transformation by a Desoxyribonucleic Acid
Fraction Isolated from Pneumococcus Type III
O. T. AVERY, C. M. MACLEOD, AND M. MCCARTY

T
he authors of this paper were the first to report that the biochemical
material responsible for the transfer of genetic information was
deoxyribonucleic acid (DNA). The observations presented in this man-
uscript laid the foundation for the evolution of our understanding of the
molecular function of DNA and for the development of Watson and Crick’s
Nobel Prize-winning model of that structure. In this study, Avery and col-
leagues pursued an observation reported by F. Griffith (J. Hyg. 27:113–159,
1928) that, until that time, had not been fully appreciated. Griffith found that
when a culture of avirulent, live, rough Streptococcus pneumoniae type II was
mixed with virulent, heat-killed smooth-type III pneumococci and injected
subcutaneously into mice, the animals frequently died. Smooth-type III S.
pneumoniae were isolated from the heart blood of dying animals. Avery et al.
made the critical connection that transformation of S. pneumoniae from an
avirulent phenotype to a virulent one was a consequence of transfer of DNA
from dead smooth organisms to live rough ones. The authors correctly con-
cluded that the “. . . chemically induced alterations in cellular structure and
function are predictable, type-specific, and transmissible.”
ALISON O’BRIEN

Reproduced from The Journal of Experimental Medicine 79:137–156. Copyright © 1944,

by permission of The Rockefeller University Press.

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