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Antioxidant composition of selected high alp... (1996)
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Plant, Cell and Environment (1996) 19,138-146

Antioxidant composition of selected high alpine
from different altitudes

species

B. WILDI & C. LUTZ

GSF-Forschungszentrum für Umwelt und Gesundheit, Expositionskammern, Ingolstädter Landstr. 1, D-85764 Oberschleißheim,
Germany

ABSTRACT

Selected high alpine plant species were collected from dif¬
ferent elevations in the Obergurgl/Ötztal subnival and
nival regions in Austria to investigate the content of
antioxidants in plants growing under the particular in vivo
conditions experienced in this area (e.g. chilling stress,
short Vegetation period and high irradiation). The con-
tents of antioxidants (ascorbic acid, tocopherol and glu-
tathione) and photosynthetic pigments were measured
throughout the day. The Contents of most compounds were
found to follow a diurnal rhythm, with the maximum
occurring at midday and the minimum during the night. It
was not clear whether these fluctuations were tempera-
ture-dependent or light-dependent.

Analyses of the antioxidant spectrum in the same plant
species at different altitudes (and thus under different envi-
ronmental conditions: as altitude increases, for example,
day temperature decreases and light intensity increases)
revealed that the total amount of antioxidants increases as
altitude increases. This enhancement was mainly due to
ascorbic acid Contents. Each plant species displayed a spe-
cific reaction to the increase in stress that accompanies an
increase in altitude, resulting in a broad adaptation spec¬
trum for these plants. The present study suggests that the
combined effect of Iower temperature and higher light
intensity induces higher antioxidant Contents.

Key-words: alpine plants; altitude effects; antioxidants;
pigments; stress avoidance.

INTRODUCTION

Alpine environments are characterized by short, often cold
and unpredictable growing seasons. At increasing altitudes
(here above the timber line), plants are exposed to decreas-
ing mean temperatures and increasing light intensities
(Larcher 1983, 1994a). Photosynthesis in high alpine plants
seems to be adapted to these conditions, as has been found
in the field by Moser (1973), Larcher & Wagner (1976),
Bergweiler (1987), Bergweiler & Lutz (manuscript in
preparation) and Körner & Diemer (1987, 1994). In a study

Correspondence: Cornelius Lutz, GSF-Forschungszentrum für
Umwelt und Gesundheit, Expositionskammern, Ingolstädter
Landstr. 1, D-85764 Oberschleißheim, Germany.

138

on Carex curvula, Körner (1982) described the light envi-
ronment in a canopy as being more important than tempera¬
ture for CO 2 -dependent photosynthetic yield. There nor-
mally seems to be no impairment of photosynthesis by high
light and low temperature under field conditions, which
indicates a very well regulated system of carbohydrate
turnover together with a functional system of antioxidants.
Intense irradiation may generate oxygen radicals, such as
singlet oxygen, in chloroplasts, which activate a defence
system (Knox & Dodge 1985). In lowland plants, only
species adapted to winter show similar protection of photo¬
synthesis (Somersalo & Krause 1990). Such protection is
probably mediated by an increase in photosynthetic elec-
tron transport (Huner 1985). This hypothesis is supported
by our studies on different high alpine species, including
green Soldanella alpina leaves taken from the snow, where
the rate of electron transport in photosystem II is up to 30%
higher than in lowland species (Bergweiler 1987).

Numerous studies describe regeneration of metabolism
and photosynthetic activities after climatic extremes, which
seems to occur much faster in alpine Vegetation than in cold-
exposed and photoinhibited species such as Hordeum
vulgäre or Spinacia oleracea. In addition to an effective elec¬
tron transport system, the unavoidable formation of radicals,
mainly in photosynthesis, requires a scavenging system.

To add a new dimension to the excellent ecophysiologi-
cal work already carried out on high alpine plant photo¬
synthesis, it is of interest to pose the following question:
what role does the antioxidative system play in plants from
higher altitudes in protecting leaves from climatic stress?

Ascorbic acid, a-tocopherol, glutathione, ß-carotene and
additional photosynthetic pigments were analysed. It has
been demonstrated in many experiments that the first three
compounds protect against oxidative damage by inhibiting or
quenching free radicals and reactive oxygen species (Ong &
Packer 1992). The antioxidant compositions of typical high
mountain plants are compared in relation to growth sites and
local light and temperature conditions. The antioxidant com¬
positions of high alpine plant species have not, to our know-
ledge, been examined before. This is also the first field study
on diurnal changes of antioxidants in alpine plants.

MATERIALS AND METHODS
Plants and study Sites

We investigated nine different plant species (shown in

© 1996 Blackwell Science Ltd