Copyright © 2009 John Wiley & Sons, Ltd. Phytother. Res. 23, 1316–1325 (2009)
DOI: 10.1002/ptr
1316 K. B. CHRISTENSEN ET AL.
Copyright © 2009 John Wiley & Sons, Ltd.
PHYTOTHERAPY RESEARCH
Phytother. Res. 23, 1316–1325 (2009)
Published online 27 January 2009 in Wiley InterScience
(www.interscience.wiley.com) DOI: 10.1002/ptr.2782
Identification of Plant Extracts with Potential
Antidiabetic Properties: Effect on Human
Peroxisome Proliferator-activated Receptor
(PPAR), Adipocyte Differentiation and Insulin-
stimulated Glucose Uptake
Kathrine B. Christensen
1
*, Ariane Minet
2
, Henrik Svenstrup
2
, Kai Grevsen
3
,
Hongbin Zhang
2
, Eva Schrader
4
, Gerald Rimbach
4
, Silvia Wein
5
, Siegfried Wolffram
5
,
Karsten Kristiansen
2,6
and Lars P. Christensen
7
1
Department of Food Science, University of Aarhus, Kirstinebjergvej 10, 5792 Aarslev, Denmark
2
Department of Biochemistry and Molecular Biology, University of Southern Denmark, Campusvej 55, 5230 Odense M,
Denmark
3
Department of Horticulture, University of Aarhus, Kirstinebjergvej 10, 5792 Aarslev, Denmark
4
Department of Human Nutrition and Food Science, Christian-Albrechts University, Hermann-Rodewald-Str. 6, 24098 Kiel,
Germany
5
Department of Animal Nutrition and Physiology, Christian-Albrechts University, Hermann-Rodewald-Str. 9, 24098 Kiel,
Germany
6
Department of Biology, University of Copenhagen, Ole Maaløes Vej 5, 2200 Copenhagen N, Denmark
7
Institute of Chemical Engineering, Biotechnology and Environmental Technology, University of Southern Denmark, Niels
Bohrs Allé 1, 5230 Odense M, Denmark
Thiazolidinediones (TZDs) are insulin sensitizing drugs used to treat type 2 diabetes. The primary target
of the TZDs is the peroxisome proliferator-activated receptor (PPAR)
γ
γ
γ, a key regulator of adipogenesis and
glucose homeostasis. Currently prescribed TZDs are full PPAR
γ
γ
γ agonists, and their use is associated with
several side effects. Partial PPAR
γ
γ
γ agonists appear to be associated with fewer side effects but may still confer
the desired insulin sensitizing action. Extracts from common medicinal/food plants were tested in a screening
platform comprising a series of bioassays, including tests for PPAR
γ
γ
γ,
α
α
α and
δ
δ
δ transactivation, adipocyte
differentiation and insulin-stimulated glucose uptake, allowing identification of plants containing potentially
interesting PPAR agonists. Twenty-two plant extracts out of 133 were found to increase insulin-stimulated
glucose uptake and 18 extracts were found to activate PPAR
γ
γ
γ, 3 to activate PPAR
α
α
α and
γ
γ
γ, 6 to activate
PPAR
δ
δ
δ and
γ
γ
γ, and 9 to activate PPAR
γ
γ
γ,
α
α
α and
δ
δ
δ. Among the 24 different plant species tested in the platform,
50% were shown to contain compounds capable of activating PPAR
γ
γ
γ and stimulating insulin-dependent
glucose uptake with no or little effect on adipocyte differentiation warranting further studies and characteri-
zation. Copyright © 2009 John Wiley & Sons, Ltd.
Keywords: medicinal plants; food plants; type 2 diabetes; peroxisome proliferator-activated receptors (PPARs); adipocyte
differentiation; glucose uptake.
Received 21 August 2008
Revised 8 December 2008
Accepted 8 December 2008
* Correspondence to: Kathrine B. Christensen, University of Aarhus,
Faculty of Agricultural Sciences, Department of Food Science,
Kirstinebjergvej 10, 5792 Aarslev, Denmark.
E-mail: kathrine.bisgaard@agrsci.dk
Contract/grant sponsor: EU Interreg IIIA and the Danish Council for
Strategic Research; contract/grant number: 2101-01-0065.
INTRODUCTION
Obesity is the major factor predisposing individuals to
a complex of dyslipidaemic/metabolic disorders col-
lectively named the metabolic syndrome characterized
by a group of metabolic risk factors in one person.
They include: abdominal obesity, atherogenic dyslipi-
daemia, elevated blood pressure, insulin resistance or
glucose intolerance, a prothrombotic state and a pro-
inflammatory state. Eventually, this may lead to the
development of overt type 2 diabetes (T2D) and fatal
cardiovascular disorders (Mlinar et al., 2007). An
important early event preceding overt symptoms of the
metabolic syndrome is the unnoticed development of
insulin resistance. Insulin resistance is a state in which
the cells do not respond to normal levels of insulin and
one key factor conferring insulin resistance is excess
release of free fatty acids (FFAs) from visceral adipose
tissue (Gimeno and Klaman, 2001). FFAs impair insulin
signalling in insulin sensitive tissue such as the muscles.
Insulin resistance is also affected by the release of
certain hormones and cytokines from adipose tissue,
e.g. tumour necrosis factor α that increases insulin re-
sistance. By contrast, the adipocytokines adiponectin
and leptin are crucial for maintaining a normal level of
insulin sensitivity. Adiponectin increases the catabolism
of FFAs and thereby reduces insulin resistance (Mlinar
et al., 2007). Insulin resistance is initially counteracted
by an increased release of insulin from the β-cells
to maintain glucose homeostasis. Over time the β-cells
become unable to secure a sufficient insulin release
resulting in hyperglycaemia and eventually the patient
develops overt T2D (Mlinar et al., 2007). According to