MEE T I N G REP 0 R
Endothelin: a knockout in London
Bruno Battistini, Renia Botting and Timothy D. Warner
At a recent conference* on endothelin,
all the major areas of endothelin
physiology and pharmacology, and
the roles of the endothelins in health
and disease, were discussed. In ad-
dition, attention was focused on the
application of the technique of gene
knockout on endothelins.
Gene targeting
Knockout of the endothelin 1 gene
in mice (homozygous ET-l-null
mice) causes severe craniofacial and
thoracic blood vessel malformations
(Y. Kurihara, University of Tokyo),
suggesting an important role for
endothelin 1 (ET-1) in the normal
development of the pharyngeal
arches, heart and large blood vessels 1.
This appears to be dependent upon
the action of ET-1 on endothelin ET A
receptors, as the frequency and sever-
ity of these malformations are in-
creased by treatment with either the
ET A receptor antagonist BQ123 or
antibodies against the ET A receptor,
and are mimicked by knockout of
the etA.
Homozygous ET-l-null and ET A-
null mice die soon after birth but
ET-l-heterozygotic mice are viable.
Studies on these animals show them
to have lower tissue and plasma lev-
els of ET-1 than wild-type mice
(T. Kuwaki, University of Tokyo), but
unexpectedly elevated blood press-
ures. This increase in blood pressure
should not be taken to indicate that
ET-1 has a hypotensive role in the
adult animal as it may be explained
by changes in the chemoreceptor
reflex and increases in the sym-
pathetic drive, as observed in the
heterozygote mice (Kuwaki). Indeed,
overexpression of ET-1 in the rat
elevates blood pressure (S. T616maque,
Howard Hughes Medical Institute,
Dallas).
Interestingly, mice lacking endo-
thelin 3 (homozygous ET-3-null
mice) unlike ET-l-null mice, are
viable at birth, although they die at
three to four weeks of age as a result
of toxic megacolon (A. Baynash,
Howard Hughes Medical Institute,
Dallas). These mice have spotted
coats (Fig. 1) as a result of regional
lack of epidermal melanocytes,
which is an identical phenotype to
that found in the lethal spotted-rat
model of Hirschsprung disease.
Similar changes were seen in homo-
zygous ETB-null mice (K. Hosoda,
Kyoto University) indicating a role
for endothelin 3 (ET-3) and ET B
receptors in human Hirschsprung
disease. Together these results show,
unexpectedly, that ET-1 and ET-3,
acting on ET A and ET B receptors,
respectively, are essential for the
development of neural crest derived
structures.
Endothelin converting
enzymes: cloned and expressed
A recent major advance in the field
of endothelin research has been the
purification and cloning of endothe-
lin converting enzyme (ECE), which
transforms big endothelin into its
active form2-7. It is apparent that
there is not a single endothelin con-
verting enzyme but a group of
related proteins, named ECE-la,
ECE-lb and ECE-2, which have dif-
ferent preferences for big ET-1,
big ET-2 and big ET-3 as substrates
(Table 1).
In general terms these ECEs are
type II integral membrane-bound
proteases with considerable struc-
tural homology to endopeptidase
N.E.P. 24.11 and Kelt blood group
protein, and consist of a single trans-
membrane domain, conserved Zn2÷-
binding motifs, N-glycosylation sites,
a short N-terminal cytoplasmic tail
and a large extracellular portion that
contains the catalytic domain. The
ECE-la enzyme is expressed and
localized in the vascular endothelial
cells of all organs and in airway, gas-
trointestinal, and urinary epithelial
cells (F. Bayan, Montreal General
Hospital). Gel filtration suggests that
ECE-la has a molecular weight of
T
250-280 kDa (Refs 6, 8), whereas
SDS-PAGE under reducing condi-
tions reveals a single band at
120-130 kDa (Ref. 6) suggesting that
ECE-la may exist as a heterodimer.
ECE-lb is present in human renal
adenocarcinoma cells, which pro-
duce ET-2 (Ref. 7) (K. Yorimitsu,
Chiba University School of Medi-
cine), and also in human umbilical
vein endothelial cells, bovine adrenal
cortex and bovine aortic endothelial
cells (O. Valdenaire, Hoffmann-
La Roche, Basel).
Development of ECE inhibitors
Because the production of the
mature endothelins requires the
action of ECE(s), these are attractive
targets at which to aim pharmacolo~-
cal interventions.
Phosphoramidon inhibits both the
conversion of exogenously adminis-
tered big ET-1 and the de novo pro-
duction of ET-1. Phosphoramidon
analogues have, therefore, been stu d-
ied in the hope of producing selective
ECE inhibitors. For example, removal
of the rhamnose moiety of phospho-
ramidon followed by the addition of
2-(2-naphthyl)ethyl to phosphonyl-
L-Leu-L-Trp increases the potency
against ECE sixfold, but decreased it
twofold against N.E.P. 24.11 (A. Jertg,
Ciba-Geigy, Summit). However, per-
haps of more interest is the develop-
ment of CGS26303 and CGS263c)3,
which are dual N.E.P. 24.11 and ECE
inhibitors (S. De Lombaert, Ciba-
Geigy, Summit), and inhibit the
in vivo hypertensive effects of ex-
ogenous administration of big ET-1
and reduce the blood pressure of
spontaneously hypertensive rats
(De Lombaert).
Endothelin receptors
In mammals two endothelin recep-
tors have been cloned and expressed:
the ET A receptor, rank order of
potency of ET-1 = ET-2 > ET-3, and
the ET B receptor, rank order of
potency of ET-1 = ET-2 = ET-3.
Despite this, evidence from the use of
selective and nonselective endothelin
receptor antagonists has been re-
ported to suggest the existence of
subtypes of the ET Breceptor9. There
B. Bonistini,
PostdoctoralFellow,
R. Boning.
Information Scientist,
and
T. D. Wemor,
Lecturer.
William Harvey Research
Institute,
St Bartholomew's
Hospital Medical College,
Charterhouse Square,
London.UK ECIM 6BQ
*4th International
Conference on
Endothelin,
23-26 April 1995,
London, UK.
©1995, Elsevier Science Ltd TiPS-July 1995 (Vol. 16) 2 1 7