Copyright © Physiologia Plantarum 2000 PHYSIOLOGIA PLANTARUM 108: 87 – 93. 2000
Printed in Ireland —all rights resered ISSN 0031-9317
Growth in excess copper induces changes in the lipid composition and
fluidity of PSII-enriched membranes in wheat
Mike F. Quartacci
a
, Calogero Pinzino
b
, Cristina L. M. Sgherri
a
, Francesca Dalla Vecchia
c
and Flavia Navari-Izzo
a,
*
a
Dipartimento di Chimica e Biotecnologie Agrarie, Uniersita ` di Pisa, Via del Borghetto, 80, I -56124 Pisa, Italy
b
Istituto di Chimica Quantistica ed Energetica Molecolare, CNR, Via Risorgimento, 35, I -56126 Pisa, Italy
c
Dipartimento di Biologia, Uniersita ` di Padoa, Via Trieste, 75, I -35121 Padoa, Italy
*Corresponding author, e -mail: fnaari@agr.unipi.it
Received 6 April 1999; revised 6 August 1999
decreased, whereas the degradation of the polar lipids caused Changes in the lipid composition and fluidity of PSII-enriched
thylakoids were studied in seedlings of wheat (Triticum du - an accumulation of free fatty acids (FFA). The total amount
of unsaturated lipids associated with the PSII-enriched mem- rum Desf. cv. Adamello) grown in nutrient solution supple-
mented with CuSO
4
to achieve a final concentration of 10 branes of wheat was not affected by excess copper supplies,
even though changes in the individual fatty acids occurred. and 50 M Cu. Metal content increased in the chloroplasts of
the 50 M Cu-grown plants. PSII isolated from wheat sup- The effect of copper on the fluidity of PSII membranes was
evaluated by electron paramagnetic resonance (EPR) mea- plied with 10 M Cu did not show any alteration in the lipid
composition or in the lipid and protein levels of the mem- surements, using spin-probed fatty acids as probes. The PSII
membranes, spin probed by means of 5- and 16-doxylstearic branes, nor was any change in the ultrastructure of the
membranes detected. The 50 M Cu-grown plants showed acids, showed that only the fluidity of the surface region of
thylakoid swelling, particularly in the stroma and terminal the bilayer close to the polar head group was reduced follow-
ing the 50 M Cu supply. In contrast, the fluidity of the inner grana thylakoids. Furthermore, an alteration in the lipid
composition of PSII preparations was observed together with membrane region of the bilayer did not show any change. The
a decrease in the lipid content, which resulted in a reduction implications of changes in the lipid composition and lipid-
protein interactions on the fluidity of specific transversal in the lipid to protein ratio. The monogalactosyldiacylglycerol
membrane regions are discussed. (MGDG) to digalactosyldiacylglycerol (DGDG) molar ratio
under excess copper supply the production of harmful oxy-
gen species, such as hydroxyl radicals arising from superox-
ide, has been demonstrated (Navari-Izzo et al. 1998, 1999).
In addition to a direct toxic action, the accumulation of
high levels of reactive oxygen species may also be responsi-
ble for an indirect action of copper on the photosynthetic
apparatus of plants (Stohs and Bagchi 1995, Navari-Izzo et
al. 1999).
The thylakoid intrinsic pigment-protein complexes of
higher plants are distributed within the lipid bilayer so that
any change in the lipid composition and fluidity may alter
the conformation, the orientation and the function of
Introduction
Copper is known to be an essential micronutrient for higher
plants, however, when present in excess, copper causes
severe damage to plant organelles through inhibitory effects
to photosynthetic electron transport and degradation of the
chloroplast inner structure and pigment content (Ouzou-
nidou 1996, Ciscato et al. 1997). Studies on the inhibitory
effects of copper on photosynthetic activity have shown that
PSII is more sensitive to copper than PSI (Jegerscho ¨ld et al.
1995, Horva ´th et al. 1998). Copper-mediated damage to the
photosynthetic apparatus may be the result of a direct toxic
action, both on the oxidising and on the reducing sides of
PSII (Baro ´ n et al. 1995). In PSII isolated from wheat grown
Abbreiations – DGDG, digalactosyldiacylglycerol; EPR, electron paramagnetic resonance; FFA, free fatty acids; MGDG, monogalactosyl-
diacylglycerol; PC, phosphatidylcholine; PG, phosphatidylglycerol; SASL, doxylstearic acid spin probe; SQDG, sulfoquinovosyldiacylglyc-
erol.
Physiol. Plant. 108, 2000 87