Plant sulfolipid. III. Role in adaptation
The quality and/or relative content of plant sulfoquinovosyl diacylglycerol (SQDG) change in response to a stress action. Various types of stress action induce two types of response – more general to the oxidative stress and specific – to a concrete stress factor. Besides, two types of reaction take...
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Інститут молекулярної біології і генетики НАН України
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| Cite this: | Plant sulfolipid. III. Role in adaptation / O.I. Kosyk, A.A. Okanenko, N.Yu. Taran // Біополімери і клітина. — 2009. — Т. 25, № 2. — С. 85-94. — Бібліогр.: 78 назв. — англ. |
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| author | Kosyk, O.I. Okanenko, A.A. Taran, N.Yu. |
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| citation_txt | Plant sulfolipid. III. Role in adaptation / O.I. Kosyk, A.A. Okanenko, N.Yu. Taran // Біополімери і клітина. — 2009. — Т. 25, № 2. — С. 85-94. — Бібліогр.: 78 назв. — англ. |
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| description | The quality and/or relative content of plant sulfoquinovosyl diacylglycerol (SQDG) change in response to a stress action. Various types of stress action induce two types of response – more general to the oxidative stress and specific – to a concrete stress factor. Besides, two types of reaction take place in photosynthesizing and non-photosynthesizing tissues. SQDG molecules take part in the adaptation reaction being cytochrome oxidase, CF1, F1, ATPase regulators, protectors and stabilizing agents for D1/D2 dimers and LHC II. This compound in non-photosynthesising tissues could be connected with negative charge domination required for lipoprotein complex stabilisation. SQDG quantitative changes and acyl composition shifts take place at both abiotic and biotic factors impact.
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Plant sulfolipid. III. Role in adaptation
O. I. Kosyk, A. A. Okanenko, N. Yu. Taran
National Taras Shevchenko University of Kyiv
64, Volodymyrska Str, Kyiv, Ukraine, 01033
a_okanenko@yahoo.co.uk
The quality and/or relative content of plant sulfoquinovosyl diacylglycerol (SQDG) change in response to a
stress action. Various types of stress action induce two types of response – more general to the oxidative
stress and specific – to a concrete stress factor. Besides, two types of reaction take place in
photosynthesizing and non-photosynthesizing tissues. SQDG molecules take part in the adaptation reaction
being cytochrome oxidase, CF1, F1, ATPase regulators, protectors and stabilizing agents for D1/D2 dimers
and LHC II. This compound in non-photosynthesising tissues could be connected with negative charge
domination required for lipoprotein complex stabilisation. SQDG quantitative changes and acyl
composition shifts take place at both abiotic and biotic factors impact.
Keywords: sulfolipid, sulfoquinovosyl diacylglycerol, stress.
In re sponse to en vi ron men tal stresses, liv ing or gan isms
ac quired the ca pa bil ity of rec og niz ing such stresses and
adapt ing them selves to var i ous types of stress dur ing
evo lu tion. Stress af fects many phys i o log i cal pro cesses
in clud ing pro tein and lipid me tab o lism and en zyme ac -
tiv ity [1–5]. Since lipids are ma jor com po nents of
mem branes and are in flu enced con sid er ably by en vi -
ron men tal fac tors [6, 7] over all plant growth and deve-
lopment will be af fected by sig nif i cant changes in lipid
me tab o lism. The re sults of stud ies de voted to lipid in -
volve ment in ad ap ta tion pro cesses show that both
sulfoquinovosyl diacylglycerol (SQDG) quan ti ta tive
changes and fatty acid com po si tion shifts take place.
Oxidative stress. The questions of specificity of
plant adaptive reaction to the environment unfavorable
factor action are bound mainly with the oxidation
stress. Abiotic and biotic stresses can cause in plants
secondary oxidation stress. Plenty of oxygen and
highly power reactions of electron transfer associated
with the thylacoid membranes are the main source of
highly active oxygen intermediators in photosynthe-
sizing tissues of plants. Anion of superoxide of O2
.– or a
free oxygen radical is generated when electrons of PS II
are accepted by O2 instead of feredoxine and is
transformed with participation of SOD usually in H2O2
or sometimes in highly toxic free radical .OH via
Fe-catalyzed Haber-Weiss reaction [8]. Action of
abiotic and biotic stresses causes growth of main-
tenance of highly active oxygen derivates. Superoxides
are the charged molecules and can not pass via bio-
logical membranes.
Therefore subcellular compartmentation of
protective mechanisms is critical for elimination of
superoxide ions in the places of their origin in cells [9].
Oxygen radical action upon isolated membranes
caused an inccrease in their viscosity and permeability,
in temperature of lipid phase transition, lipid
phosphorus liberation and piling up free fatty acids
85
ISSN 0233-7657. Á³îïîë³ìåðè ³ êë³òèíà. 2009. Ò. 25. ¹ 2
Ó ²íñòè òóò ìî ëå êó ëÿð íî¿ á³îëî㳿 ³ ãå íå òè êè ÍÀÍ Óêðà¿ íè, 2009
[10]. In general the mechanism of attack of free radicals
is known to include the oxidation of acyl chains [11],
but it is not excluded, that free radicals can cause the
reactions of de-etherification of polar lipids and
liberation of fat acids which can be easily oxidized
[12]. Bearing in mind that the oxidative stress
accompanies many other stresses of plants, any
changes in the lipid composition caused under such
circumstances are of special significance.
A few data avail able ev i dence that ox i da tive pro -
cesses in duced by a high con cen tra tion of ozone run in
two phases. Dur ing the first phase (8 hours) a loss of
pig ments and lipids (mainly MGDG with some
DGDG) was observed. The loss in chloroplast lipids is
ac com pa nied by a small in crease in the malondial de -
hyde (MDA), TAG and DAG con tent. The sec ond
phase of ox i da tive in jury is char ac ter ized by a mas sive
de struc tion of pig ments and be gins with a dras tic fall in
MGDG and a smaller de crease in DGDG and PC con -
tents which are ac com pa nied by sig nif i cant in creases in
TAG, DAG and MDA [13]. How ever, the an ionic lipid
(SQDG and PI) con tent was sta ble for the pe riod of
ozone ex po sure (in spin ach leaves, at least). Sim i lar
lipid changes were also ob served in sev eral plant spe -
cies, and in broad bean leaves, a rel a tive in crease in
SQDG took place. Be cause both galactolipids were sig -
nif i cantly de stroyed dur ing ozone ex po sure, the SQDG
con tent ex pressed as mol% of the to tal glyco lipids in -
creased up to 45 (de pend ing on a spe cies) [14, 15]. In -
ter est ing re sults were ob tained with a sulfoglycolipidic
frac tion iso lated from the red microalga Porphyridium
cruentum. It was dem on strated that sulfolipid con -
tained large amounts of palmitic acid (26.1 %),
arachidonic acid (C20:4n-6, 36.8 %), eicosapentaenoic
(C20:5n-3, 16.6 %) ac ids and 16:1n-9 fatty acid
(10.5 %) and strongly in hib ited the pro duc tion of
superoxide an ion gen er ated by peritoneal leu ko cytes
[16]. Our ex per i ments with the in duced ox i da tive stress
showed that hy dro gen per ox ide af fect pig ment and
glycolipid com po si tion with in creas ing lipid pero-
xidation ac tiv ity in dose and time de pend ent man ner.
The treat ment with var i ous hy dro gen per ox ide con cen -
tra tion caused sig nif i cant SQDG con tent in crease in all
vari ants. The re sults of field ex per i ments showed a
dras tic fall of SQDG level in 24 hours (h) with a sub se -
quent sig nif i cant lipid ac cu mu la tion. This com pound
quan tity was sta ble af ter the sec ond treat ment when
light MGDG de crease was noted and DGDG level was
sta ble dur ing the whole ex per i ment [17]. Con cern ing
the mean ing of these changes the idea seems ac cept able
that this lipid may also pro vide a source of cysteine un -
der the con di tions of ox i da tive stress [18]. It func tions
as a com po nent in the sul fur cy cle in plants and is rap -
idly me tab o lized for protein production under the
conditions of sulfur depletion [19]. Thus the addi-
tionary quantity of thiol group needed at oxidative
agent action can be supplied.
Ir ra di a tion level. The quan tity of the light in nat u -
ral en vi ron ments can vary over sev eral or ders of mag -
ni tude and on a time scale that ranges from sec onds to
sea sons. Plant ac cli ma tion to light in ten sity de pends
upon the struc ture and func tion of the photosynthetic
ap pa ra tus. The photosystems and their subcomplexes
(LHCs, D1, D2) are an chored within the thylakoid mem -
brane through the lipids. This close con nec tion be -
tween lipids and photosystem subcomplexes in di cate
an in ter de pen dence be tween both, which sup ports the
con cept of pho to syn the sis reg u la tion by changes in the
thylakoid mem brane struc ture or the en tire chloroplast
[20]. Be sides, it was said ear lier that the cells of
Chlamydomonas SQDG-de fi cient mu tant tended to
suf fer from photoinhibition [21]. The data avail able in -
di cate that the red alga Tichocarpus crinitus ex posed to
low light (10 % of the in ci dent photosynthetically ac -
tive ra di a tion) con di tions ac cu mu lated the only SQDG
con tent char ac ter ized by palmitooleic (hexadecenoic,
16:1) acid 20-fold in crease with small con tent de vi a -
tion of the other FA res i dues [22]. But Ant arc tic sea ice
di a toms (the al gal com po si tion in the cul ture was 66 ±
± 11 % of Navicula gelida var. antarctica, 20 ± 7 % of
Fragilariopsis curta and 14 ± 9 % of Nitzschia medio-
constricta) did not show any dif fer ence in SQDG con -
tent at two dif fer ent pho ton flux den si ties – 15.0 ±
± 5.0 mmol pho tons m–2×s–1 and pho ton flux den sity of
2.0 ± 1.0 mmol pho tons m–2×s–1. The main dif fer ence was
palmitic res i due de crease ac com pa nied by stearic
(18:0) and eicosapentaenoic acid (20:5n-3) twice in -
crease [20]. The au thors con sid ered the in crease in pho -
ton trap ping and an in crease in elec tron trans port ve -
locity at PS II un der 2 mmol pho tons m–2×s–1 be tween
bound QA and QB as a con se quence of in creas ing FA
desaturation of typ i cal chloroplast lipids (MGDG,
86
KOSYK O. I., OKANENKO A. A., TARAN N. Yu.
SQDG, PG), par tic u lar by in creas ing 20:5 n-3 of
MGDG and SQDG. It sup ports the QA and QB in ter ac -
tions and thus the ve loc ity of elec tron flow.
Thermoresistance. Many stud ies on the lipid
changes caused by tem per a ture have been made with
cyanobacteria be cause these prokaryotic or gan isms are
ho mo ge neous in cul ture and re spond very quickly to
the tem per a ture shifts. The ex per i ments per formed
showed that a tem per a ture de crease caused an in crease
in the SQDG con tent or did not af fected it. For ex am -
ple, in Anacystis nidulans an in crease in MGDG and
SQDG and a de crease in DGDG were ob served. Al -
most all of the palmitate at the sn-1 po si tion of SQDG
was con verted to palmitoleate at low growth tem per a -
tures [23–25]. In the cyanobacterium Spirulina
platensis cul ti vated at 35, 30 and 27 °C the low tempe-
ratures caused the SQDG con tent in crease ac com pa -
nied by the desaturation of palmitate at the sn-2 po si -
tion of SQDG [26]. On the other hand, for Anabaena
variabilis grow ing at low tem per a tures, the level of
SQDG was sta ble with only a small con ver sion of
palmitate to palmitoleate [23–25]. Low tem per a tures
also in duced sim i lar ef fect in the higher plant lipid
com po si tions. Our ear lier in ves ti ga tions showed that,
dur ing au tumn hard en ing, SQDG ac cu mu la tion took
place in one-year-old ap ple shoot bark and wood (!)
and was es pe cially strik ing for a hardy ap ple spe cie
(Malus boccata Borh) [27]. Oquist [28] found a 2-fold
in crease in the SQDG con tent in pine thylakoid prep a -
ra tions dur ing the au tumn which re mained high dur ing
the win ter and then low ered in spring. SQDG from pine
in the win ter was en riched in linoleate whereas
palmitate was dom i nant in the sum mer.
The re sults avail able in lit er a ture in di cate that
superoptimal tem per a tures cause an in crease of SQDG
con tent in most plants stud ied. For ex am ple, Atriplex
lentiformis plants grown in coastal and desert re gions
ac cu mu lated SQDG (by 260 % and 64 % re spec tively)
at high growth tem per a tures [1]. When Nerium ole an -
der was grown at 45 °C rather than 20 °C the main
changes in mo lec u lar spe cies were an in crease in the
pro por tion of dipalmitoyl-SQDG from 12 up to 20 %
and a de crease in that of linolenoyl-palmitoyl SQDG
from 40 to 30 % at the higher growth tem per a ture. At
the same time the phase tran si tion tem per a ture in -
creased from 19 ± 3 °C to 24 ± 3 °C. Tak ing into ac -
count the fact that dipalmitoyl-SQDG un der goes a
tran si tion at 42 °C, it was sug gested that this mo lec u lar
spe cies could be a ma jor lipid com po nent in volved in a
phase tran si tions in the thylakoid po lar lipids [29]. Our
data showed that growth at high tem per a ture in duced
sulfolipid ac cu mu la tion but only in the wheat leaves
and chloroplasts of drought re sis tant va ri et ies. In con -
trast, in the only sen si tive spe cies (Myronivska 808)
there was a de crease in the SQDG con tent. The heat ing
at tem per a tures 40, 45 and 50 °C caused the SQDG
con tent changes in chloroplasts de scribed by curve
with one apex at 45 °C with fol low ing drop at 50 °C in
all re sis tant plants [30].
The ex plo ra tion of phys i cal prop er ties of the
SQDG showed that only in sponge cu cum ber did phase
sep a ra tion be gin at 15 °C for SQDG. Nev er the less,
SQDG from the chill ing sen si tive spe cies con tained up
to three times more stearic acid than re sis tant plants
[31]. Fur ther more, Kenrick and Bishop [32] con cluded
that if the pri mary event in chill ing sen si tiv ity of higher
plants is a phase tran si tion in bulk chloroplast mem -
brane lipids, then not only PG, but also SQDG might be
in volved. In par tic u lar, the main dif fer ence be tween
Carica pa paya (trop i cal or i gin) and C. pubescens
(adapted to tem per ate cli mates) was in their SQDG spe -
cies (57 % and 36 % sat u rated, re spec tively), and not in
PG. Their re sults showed that linolenic acid res i due
was abun dant in SQDG from the chill ing-tol er ant spe -
cies whereas palmitate was a ma jor com po nent in the
tem per a ture-sen si tive plants.
Wa ter def i cit. Wa ter def i cit af fected the SQDG
amounts in wheat de pend ing upon wheat va ri ety
drought re sis tance. Our data showed that wa ter def i cit
in duced some SQDG ac cu mu la tion in drought re sis tant
wheat plants and the dras tic de crease of it con tent in
sen si tive plants [33]. Sim i lar re sults were pre sented by
Quartacci et al. [2]. So, thylakoids of tol er ant wheat va -
ri ety were char ac ter ized by SQDG ac cu mu la tion
whereas sen si tive va ri et ies loosed it. Palmitic res i due
con tent in creased while palmitooleic and linolenic
ones de creased in tol er ant plant SQDG whereas
palmitic res i due level di min ished and linolenic en -
larged in soft one. Thus, the SQDG ac cu mu la tion in
tol er ant plants with a par al lel rise in sat u ra tion was ob -
served while changes in sen si tive plants were ex actly
the op po site. Field ex per i ments with ar ti fi cial ir ri ga -
87
PLANT SULFOLIPID. III. ROLE IN ADAPTATION
tion (con ferred as «con trol») dur ing drought showed an
in crease in SQDG at two stages of de vel op ment (the
stage of stooling and the stage of milk ripe ness) for
wheat ex posed to drought (by 73.7 and 51.1 % re spec -
tively) [34]. Sim i lar data were ob tained by Pancratova
and Karimova [35] when they ex posed rye to drought.
Com bine ac tion of heat and wa ter def i cit in duced the
SQDG ac cu mu la tion in the al most all re sis tant plant
chloroplasts, whereas the SQDG con tent de creased in
sen si tive va ri ety more dras ti cally than in the cases of
sin gle fac tor ac tion [36]. Thus the shifts ob served at the
heat are sim i lar to those un der the wa ter def i cit ac tion
and their com bine ef fect in duced the SQDG ac cu mu la -
tion some times much more sig nif i cant than when sin -
gle fac tor ac tion. The im por tance of SQDG was also in -
di cated from ex per i ments on rehydration of air-dried
cells of the des ic ca tion-tol er ant fil a men tous cyanobac -
te rium Nostoc com mune. The radiolabeled pool sizes of
PG and SQDG reached steady-state within sev eral min -
utes, whereas the two abun dant mem brane glyco lipids,
MGDG and DGDG, achieved uni form la bel ing only
within 2 h [37]. This rapid re sponse may be con nected
with the abil ity of this cyanobacterium to tol er ate de-
siccation.
Sa lin ity stress. Higher plant Calystegia soldanella
R. Br. (Convolvulaceae) – a halophyte plant that can
grow in some ar eas along the Black Sea where the soil
may con tain up to 700–900 mg salts in 100 g soil. The
main glycolipid was SQDG (31.7 %) with al most equal
con tent of MGDG (11.1 %) and DGDG (14.3 %). Con -
trary to the other halophyte plants from the same regi-
on, the con tent of phospholipids is rel a tively low [38].
Ad ap ta tion of the plant cells to high sa lin ity in -
volves os motic ad just ment and the compartmentation
of toxic ions, whereas an in creas ing body of ev i dence
sug gests that high sa lin ity also in duces ox i da tive stress
[39–43]. Sev eral lab o ra to ries have re ported that salini-
ty im pairs photosynthetic ac tiv ity in a num ber of
photosynthetic or gan isms [44, 45] and af fect lipid com -
po si tion.
Early re sults showed that the SQDG con tent was
sta ble (rel a tive to chlo ro phyll) dur ing ad ap ta tion of
bar ley seed lings to high con cen tra tions of so dium chlo -
ride while the MGDG con tent of thylakoid mem branes
de creased con sid er ably. The lat ter de crease meant that
the rel a tive per cent age of SQDG com pared to to tal
thylakoid lipids in creased by 30 % [46]. In wheat roots,
in creased salt lev els (cal cium sul fate) re sulted in higher
(two-fold in crease) lev els of SQDG, while the re verse
oc curred in lipids from oat roots [47]. It may be in ter -
est ing to note in this con nec tion (and in view of the pos -
si ble in volve ment of SQDG in ATPase ac tiv ity) that
Ca2+ ac ti va tion (via annexin) of the root ATPase is im -
por tant in wheat [48] whereas in oat roots the ATPase
ac tiv ity is mainly in creased by Mg2+. How ever, treat -
ment of sugar beet with so dium sul phate did not con -
firm the sug ges tion that SQDG was in volved in salt-ad -
ap ta tion [49]. On the other hand, SQDG ranged from 5
in spin ach to 20 mol e cules in the halotolerant alga
Dunaliella sa lina per CF0-CF1 ATPase com plex and
this lipid could not be readily ex changed [50]. Growth
of Synechococcus 6311 in the pres ence of 0.5 M NaCl
is ac com pa nied by sig nif i cant changes in both thy-
lakoid and cy to plas mic mem brane lipid com po si tion.
MGDG con tent de creased while DGDG quan tity
raised. The to tal con tent of an ionic lipids (PG and
SQDG) was al ways higher in the iso lated mem branes
and the whole cells from high salt-grown cul tures. The
ob served changes in mem brane fatty ac ids and lipids
com po si tion cor re late with the al ter ations in elec tron
and ion trans port ac tiv i ties, and it is con cluded that the
re ar range ment of the mem brane lipid en vi ron ment is an
es sen tial part of the pro cess by which cells con trol
mem brane func tion and sta bil ity [51]. And it should be
kept in mind that in ex treme halophiles (Halobac-
terium cutirubrum, Haloferax volcanii T., Planococcus
H8) po lar lipid ex tracts con tained near 14 % sul fated
glyco lipids and that lipids ex tracted from the cry-
stallizer ponds of the sal terns of Margherita di Savoia
(It aly) and Eilat (Is rael) and from cul tures of rep re sen -
ta tive spe cies of the Halobacteriaceae showed that a
sulfated diglycosyl diether was the major glycolipid
detected in the biomass of both salterns [52].
Re sults ob tained by Zhang et al. [53] showed that
trans gen ic Bras sica napus plants overexpressing
AtNHX1, a vacuolar Na+/H+ antiport from Arabidopsis
thaliana, were able to grow, flower, and pro duce seeds
in the pres ence of 200 mM so dium chlo ride. The data
sug gest that the ma jor struc tural lipids of the
extraplastidic com part ments (PC and PE) and of the
chloroplasts (DGDG and MGDG) were un af fected by
the overexpression of AtNHX1 and by the growth of the
88
KOSYK O. I., OKANENKO A. A., TARAN N. Yu.
trans gen ic plants at high sa lin ity. Some dif fer ences,
how ever, were seen in the mi nor chloroplastic lipids,
SQDG and PG in 200 mM NaCl. Al though the 16/18C
ra tios were the same, there was lesser unsaturation of
the 18C fatty ac ids in both SQDG and PG from trans -
gen ic plants grown in 200 mM NaCl. SQDG ex pressed
as mol% of four ma jor chloroplast lipids (MGDG,
DGDG, SQDG and PG) in the case of the
overexpression of AtNHX1, a vacuolar Na+/H + antiport
from A. thaliana, overpassed wild type (at 10 mM
NaCl) by 31 % at 10 mM NaCl and by 58 % at 200 mM
NaCl. This phe nom e non al lows us to sup pose SQDG to
take part in the Na+/H + antiport func tion ing (per haps as
ATPase sta bi liz ing agent).
The study of dif fer ent NaCl con cen tra tion ef fect
upon halophytes and a glycophyte showed that in the
halophyte As ter tripolium the SQDG con tents in -
creased in dose-de pend ing man ner [54]. The differen-
ces in SQDG con tents be tween the con trol, 258 and
517 mM NaCl in the wa ter ing so lu tion were al ready
sub stan ti ated af ter one day and change unsignificantly
for 10 days of ex per i ment. The in crease in SQDG in
As ter treated with high salt cor re lated pos i tively with
the in crease in chlo ro phyll con tents. The larg est diffe-
rences were ob served on the 7th day of the treat ment. In
roots of the same As ter plants the sulfolipid con tents
are about 5 ́ lower than in As ter leaves and in creased in
roots of salt treated plants es pe cially dis tinctly on the
5th day.
The sulfolipid con tent of the other the halophyte
Sesuvium portulacastrum was also higher in the leaves
of plants treated with NaCl when com pared to the
non-treated plants. SQDG con tained pre dom i nantly
palmitic acid and g-linolenic acid, and lesser amounts
of linoleic acid and com mon tend was g-linolenic acid
de crease. In Sesuvium af ter 7 days of salt treat ment the
18:2 spe cies clearly in creased in com par i son to the
con trol in a sim i lar way as in As ter. Main SQDG mo -
lec u lar spe cies in the halophytes were palmi-
toyl-linolenoyl (16:0/18:3) and dilinolenoyl
(18:3/18:3) forms. ATPase ac tiv ity study showed that
F-ATPase lo cal ized in chloroplast and mi to chon dria
in creased it cor re lated to NaCl con cen tra tions in the
growth me dium in As ter and, in less ex tent, Sesuvium.
This phe nom e non was ac com pa nied by sig nif i cant in -
crease of SQDG con tent in As ter and Sesuvium. To
con firm the re sults sim i lar in ves ti ga tion was car ried
out in Thellungiella halophila, a salt tol er ant rel a tive of
Arabidopsis and in Arabidopsis it self. In Thellungiella
SQDG con tent in creased due to the ef fect of NaCl
echo ing the re sult of As ter and Sesuvium, but no signi-
ficant changes could be ob served in Arabidopsis
[54–56]. These re sults con firmed the sug ges tion that
sulfolipids sta bi lize and/or activate F-ATPases.
Thus, tak ing into ac count that plenty of met a bolic
pro cesses and adaptative re ac tions flow en gag ing
ATPases, one could sup pose the size of SQDG in volve -
ment in these pro cesses. Au thors con cerned this SQDG
in crease sup posed that this lipid is in volved in sta bi li -
za tion of ATPase com plexes and PS II – As ter
F-ATPase activity increased with increasing NaCl
concentration in the medium of growth [55] and might
also play a role in signalling processes.
Our ex per i ments [57] re vealed that SQDG ac cu mu -
la tion in euhalophyte Salicornia europaea L. and
crinohalophyte Halimione pedunculata L. was pos si -
bly con nected with its abil ity to ac cu mu late high con -
cen tra tions of salts (NaCl) in its tis sues. In con trast,
SQDG lev els were sta ble in Ar te mi sia arenaria DC
dur ing changes in os motic pres sure caused by
photoassimilate ac cu mu la tion and de creased in
Atriplex pedunculata dur ing the elim i na tion of salts
from this or gan ism. Hence, any in volve ment of SQDG
in salt ad ap ta tion seems to be de pend ent upon the var i -
ous mech a nisms of salt tolerance used by different
organisms.
Thus, SQDG role in this case should be con sid ered
in fol low ing di rec tions: as com pound closely con -
nected with photophosphorylation cou pling fac tor and
ATPase sta bi liz ing en ergy sup ply for antioxidative ac -
tiv ity and Na+/H+ antiport func tion ing; as com pound
neu tral iz ing cat ion ex cess and capturing extra water.
Heavy metal ac tion. The quan tity of heavy met als
in soil is of ten in creased greatly in in dus trial ar eas and,
when they are taken up by plants, they can in duce both
toxic and adap tive re sponses. There are a few stud ies
con firms that lipid com po si tion can be af fected [58].
Cad mium ex po sure in duced a de crease in MGDG and
SQDG con tents in B. napus leaves which was ac com -
pa nied by a sharp in crease in ex tra-chloroplastic lipids
(PC and PI) [59]. Our re sults [60] showed that lead,
sup plied at var i ous con cen tra tions to hy dro ponic cul -
89
PLANT SULFOLIPID. III. ROLE IN ADAPTATION
tures, caused a de crease in SQDG in wheat seed ling
leaves and roots. Thus, most heavy met als tested cause
a de crease of SQDG con tent. Con cern ing the phe nom e -
non ob served we can as sume, that this lipid be ing
strong an ion can cre ate com plex com pound with lead
atom (we ob served it in vi tro at least). It is pos si ble
com pet i tive us age of sul fur for sul fur con tain ing
cys-rich pep tides (phytochelatins) and pro tein syn the -
sis (judg ing on in creas ing pro tein con tent) ac cord ing to
the sug ges tion that the cells uti lize sul fur pref er en tially
for the syn the sis of es sen tial me tab o lites, such as pro -
teins, rather than for SQDG syn the sis [61]. But in for -
ma tion pre sented al lows also to sug gest that me tab o -
lism changes in duced by lead de pend upon plant spe -
cies and con di tions of metal ac tion. For ex am ple, while
acute ex po sure of the moss, Rhytidiadelphus squar-
rosus, to low lev els (1–10 mM) of lead ni trate did not
change the radiolabelling of SQDG sig nif i cantly, po-
pulations gath ered from lead-pol luted soils showed
more la bel ing of chloroplast lipids than moss from un -
pol luted ar eas. It was con sid ered the in creased me tab o -
lism of chloroplast lipids in moss from lead-pol luted
re gions may rep re sent an adap tive re sponse. Thus, the
mem brane lipids, dam aged as a re sult of heavy metal
pol lu tion, could be re placed and any det ri men tal ef fect
on pho to syn the sis be ing min i mized [62].
Me chan i cal in jury. Plants re act to wound ing, ei -
ther me chan i cal or caused by her bi vore feed ing, by ac-
tivating the tran scrip tion of a set of genes, the func tion
of which is mainly de voted to wound heal ing and the
pre ven tion of any sub se quent patho gen at tack. On the
other hand it is known that wound ing in duce ROS and
H2O2 form ing [63] fol low ing by lipid com po si tion cha-
nge re ac tions [15]. Our data showed that wound ing
(cut ting the leaf apex) in duce in crease (» on 20 %) both
leaf galactolipids in 4 h with re turn ing to con trol in
24 h. Much more in ter est ing were the SQDG changes
both through 4 and in 24 h. In leaves this com pound ac -
cu mu la tion up to 150 % of con trol was ob served in 4 h
and changed by dras tic fall (twice com par ing to con -
trol) in 24 h. But most in ter est ing was the fact (tak ing
into ac count that wound ing was ren dered as cut ting off
1 cm of leaf apex) sig nif i cant SQDG con tent increase
(twice) in roots in both 4 and 24 h. We con sider this
phe nom e non to be an ev i dence of SQDG tak ing part in
plant sys tem re sponse (un pub lished re sults).
In fec tion. A few ex per i ments sug gest that in fec -
tion can af fect mem brane lipids. In our ex per i ments, a
rel a tive ac cu mu la tion of SQDG in the to tal glyco lipids
in wheat plants in fected by Puccinia graminis [34] and
kid ney bean plants in fected by po tato x potexvirus
(PXV) or to bacco mo saic (TMV) vi rus was ob served.
The lat ter phe nom e non was con firmed by a sig nif i cant
in crease of the SQDG/DGDG ra tio (used be cause
DGDG was the most sta ble com po nent in the ex per i -
ments) [64]. These changes could be use ful for the
plant if the ac tion of SQDG in in hib it ing vi ral de vel op -
ment (and against DNA-poly mer ase and re verse trans-
criptase ac tiv ity [65, 66]), also ap ply to in vivo situati-
ons. On the other hand, it could be con nected with ox i -
da tive stress ac com pa nied the vi rus in fec tion [67, 68].
But in bar ley stripe mo saic vi rus strains and poa
semilatent hordeivirus losses of MGDG, PG, SQDG
and chlo ro phyll as well as rel a tive in creases in
phosphatidylcholine, phosphatidylethanolamine and
phosphatidylinositol con tents were ob served [69].
Thus, in for ma tion pre sented ar gues that qual ity and/or
rel a tive con tent of SQDG changes in re sponse to a
stressor ac tion.
Con clu sions. Thus, an a lyz ing in for ma tion pre -
sented one could con clude that SQDG func tions known
to date are mul ti ple. The qual ity and/or rel a tive con tent
of SQDG changes in re sponse to a stress ac tion. Con -
cern ing the in ter pre ta tion of the phe nom ena ob served
at var i ous type stress ac tion we con sider to take place
two type of re sponse – more gen eral to the ox i da tive
stress and spe cific – to the con crete stress fac tor ac tion.
Be sides, we should keep in mind the ne ces sity of con -
sid er ation of two types of re ac tion tak ing place in pho -
to syn the siz ing and non-pho to syn the siz ing tis sues. In
pho to syn the siz ing tis sues it seems pu ta tive to as sume
avail abil ity of all struc tural and func tional SQDG mol -
e cules pe cu liar i ties known for to day to sup ply their
tak ing part in ad ap ta tion re ac tion as cytochrome
oxidase, CF1, F1, ATPase reg u la tors, pro tec tors and sta -
bi liz ing agents for D1/D2 dimers and LHC II [50, 70].
Tak ing into ac count the SQDG lo cal iza tion on the na -
tive heterodimer D1/D2 sur face [71], one could as sume
that it might hold mono mers to gether as dimer [72].
There fore it is not ex cluded that SQDG cer tain mo lec u -
lar spe cie ac cu mu la tion can pre vent RC PS II deg ra da -
tion. Func tion of the com pound in nonphotosyn-
90
KOSYK O. I., OKANENKO A. A., TARAN N. Yu.
thesizing tis sues could be con nected with neg a tive
charge dom i na tion re quire ment for uni va lent cat ion
(Na+ and K+) be ing nec es sary for li po pro tein com plex
sta bili sa tion. Be sides, the com pound can re al ize
ATPase and PL A2 ac tiv ity reg u la tion [73] both in
photosynthetic and non-photosynthetic tis sues. We as -
sume also the pos si bil ity of the ad di tional wa ter
amount cap tur ing be cause of SQDG ab sorp tion in the
space be tween the SQDG lay ers in mem brane which
in ten sity in creases grad u ally when tem per a ture is ris -
ing [74]. It could also in hibit the nonbilayer struc ture
form ing by means of mak ing bilayer MGDG struc ture
or ga ni za tion and takes part in the MGDG syn the sis via
reg u la tion UDP-galactoso: diacylglycerol galactosyl
transferase ac tiv ity thus cor rect ing MGDG/DGDG ra -
tio in mem brane [75, 76]. A cer tain role SQDG plays in
the pro cesses of pro tein syn the sized trans port. It is im -
por tant be cause of the en forced syn the sis of a wide set
of spe cific pro teins dur ing a stress ac tion. The tran sit
pep tide in serts most ef fi ciently in monolayers of PG,
SQDG and MGDG sug gest ing that these lipid classes
are mainly re spon si ble for in ser tion into the tar get lipid
of mem brane [72]. But among all func tions the main
seems to be energetical one. In pho to syn the siz ing tis -
sues SQDG mol e cules sta bi lize F-ATPase, pro tect and
sta bi lize D1/D2 dimers and LHC II [50, 70]. SQDG
and the Rieske pro tein in ter ac tion in the cyt b6f struc -
tures is also very im por tant [77]. Photoinhibition aris -
ing at a stressor ac tion in duces deg ra da tion and cleav -
age of D1 pro tein of RC PS II [78]. How ever, SQDG
lo cal ized on the sur face of the na tive D1/D2 he-
terodimer might hold mono mers to gether as a dimer
[71, 72].
Thus, this lipid seems to be in volved in the stress
ad ap ta tion re ac tions and can be a unit of the ad ap ta tion
mech a nism chain.
Î. ². Êî ñèê, Î. A. Îêà íåí êî, Í. Þ. Òà ðàí
Ðîñ ëèí íèé ñóëü ôîë³ï³ä. III. Ðîëü â àäàï òàö³¿
Ðåçþìå
ßê³ñíèé i â³äíîñ íèé âì³ñò ñóëü ôîõ³íî âî çèëä³àöèëãë³öå ðî ëó
(ÑÕÄÃ) y ðîñ ëè íàõ çì³íþºòüñÿ â³äïîâ³äíî äî 䳿 ñòðå ñî ðà. гçí³
÷èí íè êè ³íäó êó þòü äâà âèäè â³äïîâ³ä³: çà ãàëü íó – íà îêèñ íþ -
âàëü íèé ñòðåñ ³ ñïå öèô³÷ó – íà êîí êðåò íèé ñòðå ñî âèé ôàê òîð.
Îêð³ì òîãî, äâà âèäè â³äãó êó ñïîñ òåð³ãà þòü ñÿ ó ôî òî ñèí òå çó -
âàëü íèõ ³ íå ôî òî ñèí òå çó âà ëè íèõ òêà íè íàõ. Ìî ëå êó ëè ÑÕÄÃ
áå ðóòü ó÷àñòü ó ðå àêö³¿ àäàï òàö³¿ ÿê ðå ãó ëÿ òî ðè öè òîõ ðî ìîê -
ñè äà çè, CF1, F1 ³ ATPaç òà ÿê àãåí òè, ùî ñòàá³ë³çó þòü äè ìå ðè
D1/D2 ³ LHC II. Íà ÿâí³ñòü ö³º¿ ñïî ëó êè ó íå ôî òî ñèí òå çó âàëü íèõ
òêà íè íàõ ìîæå áóòè ïî â’ÿ çà íà ç âè ìî ãîþ äîì³íó âàí íÿ íå ãà -
òèâ íî ãî çà ðÿ äó îä íî âà ëåí òíî ãî êàò³îíà (Na+ àáî K+) äëÿ ñòà-
á³ë³çàö³¿ ë³ïî ïðî òå¿ íî âî ãî êîì ïëåê ñó. ʳëüê³ñí³ çì³íè âì³ñòó
ÑÕÄà òà aöèëü íî ãî ñêëà äó â³äáó âà þòü ñÿ ïðè 䳿 ÿê àá³îò è÷ íèõ,
òàê ³ á³îò è÷ íèõ ñòðå ñîð³â.
Êëþ ÷îâ³ ñëî âà: ñóëü ôîë³ï³ä, ñóëü ôîõ³íî âî çèëä³àöèëãë³öå ðîë,
ñòðåñ.
Î. È. Êî ñûê, À. À. Îêà íåí êî, Í. Þ. Òà ðàí
Ðàñ òè òåëü íûé ñóëü ôî ëè ïèä. III. Ðîëü â àäàï òà öèè
Ðå çþ ìå
Êà ÷åñ òâåí íîå è îò íî ñè òåëü íîå ñî äåð æà íèå ðàñ òè òåëü íî ãî
ñóëü ôî õè íî âî çèë äè à öèë ãëè öå ðî ëà (ÑÕÄÃ) èç ìå íÿ åò ñÿ â îò âåò
íà äå éñòâèå ñòðåñ cî ðîâ. Ðàç íûå ñòðåñ ñî ðû âû çû âà þò äâà
âèäà îò âå òà: áî ëåå îá ùèé – íà îêèñ ëè òåëü íûé ñòðåññ è ñïå öè -
ôè ÷åñ êèé – íà äå éñòâèå êîí êðåò íî ãî ôàê òî ðà. Êðî ìå òîãî,
äâà âèäà ðå àê öèè íà áëþ äà þò ñÿ â ôî òî ñèí òå çè ðó þ ùèõ è íå ôî -
òî ñèí òå çè ðó þ ùèõ òêà íÿõ. Ìî ëå êó ëû ÑÕÄà ó÷àñ òâó þò â ðå àê -
öèè àäàï òà öèè êàê ðå ãó ëÿ òî ðû öè òîõ ðî ìîê ñè äà çû, CF1, F1,
ATPaç è êàê ñòà áè ëè çè ðó þ ùèå àãåí òû äè ìå ðîâ D1/D2 è LHC II.
Ðîëü ýòî ãî ñî å äè íå íèÿ â íå ôî òî ñèí òå çè ðó þ ùèõ òêà íÿõ ìî -
æåò áûòü ñâÿ çà íà ñ íå îá õî äè ìîñ òüþ äî ìè íè ðî âà íèÿ íå ãà òèâ -
íî ãî çà ðÿ äà îä íî âà ëåí òíî ãî êà òè î íà (Na+ èëè K+) äëÿ ñòàáè-
ëè çà öèè ëè ïîï ðî òå è íî âî ãî êîì ïëåê ñà. Êî ëè ÷åñ òâåí íûå èç ìå -
íå íèÿ ñî äåð æà íèÿ ÑÕÄÃ è aöèëü íî ãî ñî ñòà âà ïðî èñ õî äÿò ïðè
äå éñòâèè êàê àáè î òè ÷åñ êèõ, òàê è áè î òè ÷åñ êèõ ñòðåñ ñî ðîâ.
Êëþ ÷å âûå ñëî âà: ñóëü ôî ëè ïèä, ñóëü ôî õè íî âî çèë äè à öèë ãëè -
öå ðîë, ñòðåññ.
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ÓÄÊ 755.121
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94
KOSYK O. I., OKANENKO A. A., TARAN N. Yu.
|
| id | nasplib_isofts_kiev_ua-123456789-5652 |
| institution | Digital Library of Periodicals of National Academy of Sciences of Ukraine |
| issn | 0233-7657 |
| language | English |
| last_indexed | 2025-11-30T16:49:18Z |
| publishDate | 2009 |
| publisher | Інститут молекулярної біології і генетики НАН України |
| record_format | dspace |
| spelling | Kosyk, O.I. Okanenko, A.A. Taran, N.Yu. 2010-02-01T14:20:23Z 2010-02-01T14:20:23Z 2009 Plant sulfolipid. III. Role in adaptation / O.I. Kosyk, A.A. Okanenko, N.Yu. Taran // Біополімери і клітина. — 2009. — Т. 25, № 2. — С. 85-94. — Бібліогр.: 78 назв. — англ. 0233-7657 https://nasplib.isofts.kiev.ua/handle/123456789/5652 755.121 The quality and/or relative content of plant sulfoquinovosyl diacylglycerol (SQDG) change in response to a stress action. Various types of stress action induce two types of response – more general to the oxidative stress and specific – to a concrete stress factor. Besides, two types of reaction take place in photosynthesizing and non-photosynthesizing tissues. SQDG molecules take part in the adaptation reaction being cytochrome oxidase, CF1, F1, ATPase regulators, protectors and stabilizing agents for D1/D2 dimers and LHC II. This compound in non-photosynthesising tissues could be connected with negative charge domination required for lipoprotein complex stabilisation. SQDG quantitative changes and acyl composition shifts take place at both abiotic and biotic factors impact. en Інститут молекулярної біології і генетики НАН України Огляди Plant sulfolipid. III. Role in adaptation Рослинний сульфоліпід. III. Роль в адаптації Растительный сульфолипид. III. Роль в адаптации Article published earlier |
| spellingShingle | Plant sulfolipid. III. Role in adaptation Kosyk, O.I. Okanenko, A.A. Taran, N.Yu. Огляди |
| title | Plant sulfolipid. III. Role in adaptation |
| title_alt | Рослинний сульфоліпід. III. Роль в адаптації Растительный сульфолипид. III. Роль в адаптации |
| title_full | Plant sulfolipid. III. Role in adaptation |
| title_fullStr | Plant sulfolipid. III. Role in adaptation |
| title_full_unstemmed | Plant sulfolipid. III. Role in adaptation |
| title_short | Plant sulfolipid. III. Role in adaptation |
| title_sort | plant sulfolipid. iii. role in adaptation |
| topic | Огляди |
| topic_facet | Огляди |
| url | https://nasplib.isofts.kiev.ua/handle/123456789/5652 |
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