American Journal of Plant Sciences, 2011, 2, 619-628
doi:10.4236/ajps.2011.24073 Published Online October 2011 (http://www.SciRP.org/journal/ajps)
Copyright © 2011 SciRes. AJPS
619
Trichome-Specific Expression of
Amorpha-4,11-Diene Synthase, a Key Enzyme of
Artemisinin Biosynthesis in Artemisia annua L., as
Reported by a Promoter-GUS Fusion
Hongzhen Wang, Linda Olofsson, Anneli Lundgren, Peter E. Brodelius
School of Natural Sciences, Linnaeus University, Kalmar, Sweden.
Email: peter.brodelius@lnu.se
Received August 16
th
, 2011; revised September 7
th
, 2011; accepted September 20
th
, 2011.
ABSTRACT
Artemisia annua L. produces small amounts of the sesquiterpenoid artemisinin, which is used for treatment of malaria.
A worldwide shortage of the drug has led to intense research to increase the yield of artemisinin in the plant. In order
to study the regulation of expression of a key enzyme of artemisinin biosynthesis, the promoter region of the key enzyme
amorpha-4,11-diene synthase (ADS) was cloned and fused with the β-glucuronidase (GUS) reporter gene. Transgenic
plants of A. annua expressing this fusion were generated and studied. Transgenic plants expressing the GUS gene were
used to establish the activity of the cloned promoter by a GUS activity staining procedure. GUS under the control of the
ADS promoter showed specific expression in glandular trichomes. The activity of the ADS promoter varies temporally
and in old tissues essentially no GUS staining could be observed. The expression pattern of GUS and ADS in aerial
parts of the transgenic plant was essentially the same indicating that the cis-elements controlling glandular trichome
specific expression are included in the cloned promoter. However, some cis-element(s) that control expression in root
and old leaf appears to be missing in the cloned promoter. Furthermore, qPCR was used to compare the activity of the
wild-type ADS promoter with that of the cloned ADS promoter. The latter promoter showed a considerably lower activ-
ity than the wild-type promoter as judged from the levels of GUS and ADS transcripts, respectively, which may be due
to the removal of an enhancing cis-element from the ADS promoter. The ADS gene is specifically expressed in stalk and
secretory cells of glandular trichomes of A. annua.
Keywords: Agrobacterium Tumefaciens, Amorpha-4,11-Diene Synthase, Artemisia annua, Artemisinin Biosynthesis,
β-Glucuronidase, Gene Regulation, Promoter Activity, Stable Transformation
1. Introduction
Artemisinin is an effective anti-malarial drug, which has
become an important component of artemisinin-based
combination therapies (ACTs) [1]. The content of ar-
temisinin in Artemisia annua is, however, very low and
ranges between 0.1% and 0.8% of dry weight [2]. Selec-
tion of high-producing varieties of A. annua has met lim-
ited success but hybrids producing increased amounts of
artemisinin have been produced [3]. However, there is
still a shortage in the supply of artemisinin [4]. Different
methods have been tried to improve the artemisinin pro-
duction such as treatment with abscisic acid [5], gibbere-
lic acid [6-8] and elicitors [9-11]. Various transgenic A.
annua plants have been produced to increase the yield of
artemisinin. These attempts include down-regulation of
squalene synthase [12,13] or up-regulation of farnesyl
diphosphate synthase (FDS) [14].
Cyclization of farnesyl diphosphate (FDP) to amor-
pha-4,11-diene by amorpha-4,11-diene synthase (ADS)
is the initial step of the artemisinin biosynthetic pathway
[15] and amorpha-4,11-diene is the committed precursor
[16]. Tissue specificity of ADS expression has been
shown by GUS expression in Arabidopsis thaliana using
a fusion of the ADS promoter and the reporter gene [17].
In the following step of the artemisinin biosynthesis,
amorpha-4,11-diene is hydroxylated to yield artemisinic
alcohol. This reaction is catalyzed by a cytochrome P450
dependent amorpha-4,11-diene 12-hydroxylase (CYP71AV1)