ELSEVIER Biochimica et Biophysica Acta 1242 (1996) 217-220
BB
Biochi~ic~a
et Biophysica A~ta
Meeting Report
Programmed cell death: from development to disease
Daniel J. Hoeppner a, Michael O. Hengartner a, David E. Fisher b
a Cold Spring Harbor Laboratory, Cold Spring Harbor, NY 11724, USA
b Dana Farber Cancer Institute, Harvard Medical School, 44 Binney St., Boston, MA 02115, USA
Accepted 1 December 1995
The first Cold Spring Harbor meeting on the topic of
Programmed Cell Death took place from September 20-24,
1995. The meeting provided a forum for the presentation
of apoptosis research in a wide diversity of systems rang-
ing from primitive development to human disease.
Methodologies being applied to the study of programmed
death were replete with in vivo manipulations (genetic
screens, knockout and transgenic animals), gene cloning
approaches (two-hybrid and novel death gene strategies),
and in vitro biochemical analyses. Several major themes
resounded. One was the constantly growing collection of
protein family members playing significant roles in the
regulation of apoptosis, either positively or negatively.
Another recurring observation was that while gene func-
tion and sequence are often conserved, inputs and outputs
of apoptosis are often system-specific. Finally, the need for
clearer mechanistic information, particularly with regard to
the biochemistry of apoptosis, was manifest, although a
gradual understanding of at least some of the central
players has begun to emerge.
1. Surface receptors as death signals
Apoptosis is a particularly attractive mechanism to ac-
count for positive and negative selection processes during
T cell development, when autoreactive clones are deleted
from the immune repertoire. S. Nagata (Osaka Bioscience
Institute) described the Fas knockout mouse which dis-
plays massive lymphadenopathy and splenomegaly, akin to
the phenotypes of Ipr and gld autoimmune mice (which
harbor mutations affecting Fas or Fas ligand, respectively).
These mice are defective in deletion of peripheral T cells,
but not thymic clonal deletion. In contrast, Tak Mak
(Amgen Institute, University of Toronto) described the
phenotype of mice nullizygous for CD30, a Hodgkin's
disease cell surface marker. These mice displayed 10-fold
thymic hypertrophy. Interestingly, these thymocytes were
profoundly resistant to anti-CD3 induced apoptosis, a mea-
0304-419X/96/$32.00 © 1996 Elsevier Science B.V. All rights reserved
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sure of thymic negative selection. Thus, apoptosis appears
to be regulated via CD30 for thymic selection and via Fas
for peripheral T cell deletion. Greater mechanistic detail
awaits understanding of T cell stimulatory events, such as
B7/CD28 costimulation (L. Boise, University of Chicago),
as well as further analysis of signaling events subsequent
to receptor:ligand binding.
2. Signal transduction in apoptosis
Fas and TNF-alpha receptors are both capable of trig-
gering apoptosis and share a cytoplasmic motif termed the
'death domain.' Protein-protein interactions of this potent
receptor family have been described by several investiga-
tors. Using two hybrid approaches, B. Stanger (Harvard
Medical School) described the identification of RIP, a
Jurkat cell-derived cDNA clone encoding a death domain
as well as an N-terminal putative protein kinase motif. A
more systematic biochemical analysis of Fas associated
proteins (A. Chinnaiyan, U of Michigan Medical School;
and P. Krammer, German Cancer Research Center, Heidel-
berg) resulted in the characterization of a Fas-associated
Death-Inducing Signaling Complex (DISC) which forms
upon oligomerization of the Fas receptor. This complex
consists of several (at least four, and probably many more)
CAPs, or Cytotoxicity-dependent APO-1 associated pro-
teins. Krammer and colleagues identified these factors by
co-immunoprecipitation of oligomerized Fas. Chinnaiyan
and colleagues identified FADD/MORT-1 in a two-hybrid
screen for proteins that interact with the cytoplasmic do-
main of Fas. CAP1 and CAP2 are phosphorylated forms of
FADD/MORT-1. Both groups demonstrate a requirement
of the death domain for DISC formation, and no DISC
formation with Fas carrying the Iprcg mutation.
The nature of the other CAPS has yet to be determined,
but a number of candidate proteins have been identified by
yeast two-hybrid screens. For example, K. Chu (Chiron
Corp.) presented the identification of a Fas-associated