Genetic Modification of the Heart
The use of knockout mouse technology to achieve tissue selective
expression of angiotensin converting enzyme
Hong D. Xiao, Sebastien Fuchs, Kristen Frenzel, Lu Teng, Ping Li, Xiao Z. Shen,
Jonathan Adams, Hui Zhao, George T. Keshelava, Kenneth E. Bernstein *, Justin M. Cole
Department of Pathology and Laboratory Medicine, Rm. 7107A WMB, Emory University, Atlanta, GA 30322, USA
Received 6 January 2004; accepted 26 February 2004
Abstract
The resin angiotensin system (RAS) plays an essential role in blood pressure regulation and electrolyte homeostasis. The effecter peptide
of the RAS, angiotensin II, is produced by angiotensin converting enzyme (ACE) in multiple tissues. Genetic deletion ofACE in mice resulted
a phenotype of low blood pressure, anemia and kidney defects. However, it is not clear whether the lack of the systemic or the local production
of angiotensin II caused these defects. To understand the role of local angiotensin II production, we developed a method to achieve tissue
specific ACE expression through homologous recombination. In this review, we discuss mouse models in which endothelial ACE was
eliminated and replaced by hepatic ACE. These studies suggest that both circulating angiotensin II and local angiotensin II production play a
role in angiotensin II generation; the elimination of local angiotensin II generation up-regulates systemic production and maintains
physiologic homeostasis.
© 2004 Elsevier Ltd. All rights reserved.
Keywords: Knockout mouse technology; ACE
1. Introduction
The first indication that something in the kidney partici-
pated in the control of blood pressure was studies by Tiger-
stedt in 1898 [1]. Tigerstedt had discovered renin, but his
experiments were difficult to reproduce and, more impor-
tantly, the clinical appreciation of blood pressure was so
limited that these studies were not expanded. Barely 15 years
later, the clinical measurement of human blood pressure had
become routine. When Goldblatt et al. [2] identified the
kidney as a central regulator of blood pressure in 1934, the
modern era of blood pressure investigation began. And like
many areas of science, increased understanding paralleled
the development of new scientific tools. For instance, the
identification of angiotensin converting enzyme (ACE) in the
production of angiotensin II, published in 1956, derived from
the development of new techniques for separating small
peptides, such as angiotensin I and angiotensin II [3,4]. Also,
the development in 1977 of captopril, the first non-peptidic
inhibitor of ACE led to a much greater understanding of the
physiology of the renin–angiotensin system (RAS) [5]. Fi-
nally, the onset of the molecular era in the 1980s led to the
cloning of all the individual components of the RAS, includ-
ing the first cloning of angiotensin receptors [6,7].
We now live in the genetic age where DNA is easily
manipulated. Now, targeted homologous recombination in
mouse embryonic stem (ES) cells is a unique tool that allows
us to manipulate the mouse genome such that virtually any
genetic change imagined can be created in a mouse [8]. In
turn, this allows us to ask new questions about blood pressure
control. Consider ACE, a protein expressed on the surface of
vascular endothelial cells where it makes angiotensin II in
immediate proximity to vascular smooth muscle, a critical
target organ for this vasoconstrictor. Other tissues make
ACE, including activated macrophages, ciliated gut epithe-
lium, and areas of the brain. In the midst of this complexity,
how can we investigate the specific physiologic role of ACE
expression by a single tissue such as endothelium. ACE
inhibitors could be used, but these pharmaceuticals are sys-
temic in their effects. ACE could be genetically eliminated
through knockout mutation of the ACE gene but this too
would eliminate all protein production and not help us to
understand the specific role of ACE made by endothelium.
* Corresponding author. Tel.: +1-404-727-3134; fax: +1-404-727-8540.
E-mail address: kbernst@emory.edu (K.E. Bernstein).
Journal of Molecular and Cellular Cardiology 36 (2004) 781–789
www.elsevier.com/locate/yjmcc
© 2004 Elsevier Ltd. All rights reserved.
doi:10.1016/j.yjmcc.2004.02.013