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Lectins of oil-seed flax plants exposed to

abiotic stress
ARTICLE in RUSSIAN JOURNAL OF PLANT PHYSIOLOGY JANUARY 2012
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ISSN 10214437, Russian Journal of Plant Physiology, 2013, Vol. 60, No. 1, pp. 7783. Pleiades Publishing, Ltd., 2013.
Original Russian Text A.N. Levchuk, E.N. Voitovich, V.A. Lyakh, 2013, published in Fiziologiya Rastenii, 2013, Vol. 60, No. 1, pp. 7581.

RESEARCH
PAPERS

Lectins of OilSeed Flax Plants Exposed to Abiotic Stress

A. N. Levchuk, E. N. Voitovich, and V. A. Lyakh
Zaporozhye National University, ul. Zhukovskogo 66, Zaporozhye, 69041 Ukraine;
email: anna.levchuck@yandex.ru
Received November 4, 2011

AbstractThe seedlings of six cultivars of oilseed flax (Linum humile Mill.) differing in the extent of adap
tation to abiotic stresses (hypo and hyperthermia, osmotic stress, and salinity) were used to assess hemag
glutination activity and carbohydrate specificity of total lectin preparations extracted from various cell com
partments. In the course of adaptation of plants resistant to hyperthermia, osmotic stress and salinity, we
observed a considerable rise in the coefficient of activity of membrane lectins, whereas the adaptation to
hypothermia elevated the coefficient of activity of cell wall lectins. As to total soluble lectins, the adaptation
of flax plants was associated with the changes in the range of their carbohydrate specificity. For instance, fol
lowing the adaptation to hyperthermia, they were found to bind glucose and glucosamine, to osmotic stress
mannose and xylose, to salinitygalactose, glucose, and glucosamine; after cold resistance was developed,
total soluble lectins were found to recognize lactose and fructose. It was concluded that lectins may partici
pate in specific adaptation of flax plants to various abiotic stress factors.
Keywords: Linum humile, adaptation, abiotic stress, hypo and hyperthermia, osmotic stress, salinity, range
of carbohydrate specificity, lectins.
DOI: 10.1134/S1021443712060106

such abiotic stress factors as hypo and hyperthermia,

osmotic stress, and sodium chloride salinity associated
with osmotic stress.

INTRODUCTION
Due to the lack of mobility, plant organisms cannot
avoid unfavorable effects of environmental factors;
therefore, they have to adapt to new environmental
conditions by changing their metabolism [1, 2].
Development of plant resistance to some extent
depends on lectins (glycoproteins capable of recogniz
ing and binding carbohydrates located on cell sur
faces) [3]. Lectins are quantitatively described by lec
tin activity (minimum protein concentration that
shows biological activity) and qualitatively by carbo
hydrate specificity (the ability to recognize particular
carbohydrates) [4, 5]. According to their location
within the cell, lectins are subdivided into three
groups: membrane (located in cytoplasmic membrane
and the membranes of organelles), soluble (located in
cytosol and vacuolar sap), and cell wall lectins [4, 6].
The role of lectins of different cellular fractions in
adaptation to abiotic stress was shown in several crops
[7]. For instance, lectins of cell walls and organelle
membranes participate in the development of resis
tance to low temperatures [8, 9] and soluble and mem
brane lectinsto salinity [1014].
The aim of this work was to look into the changes in
characteristics of oilseed flax lectins depending on
total level of nonspecific resistance at early develop
mental stages and on the extent of plant adaptation to

MATERIALS AND METHODS

The experiments were conducted with 7dold eti
olated seedlings of oilseed flax (Linum humile Mill.)
representing six genotypes (cvs. Tsian, Antares, Ais
berg, Avangard, Astral, and Zolotistyi), with different
extent of adaptation to particular abiotic stress factors
[15]. The seeds were preliminarily germinated for
5 days in Petri dishes on filter paper moistened with
distilled water at room temperature (22) in the dark.
In order to produce hypo and hyperthermia, 5d
old seedlings were kept for 24 h at 4 and 40, respec
tively. To simulate osmotic stress, the medium was
supplemented with sucrose up to 15% (1.13 MPa) or
sodium chloride to 3% (2.24 MPa). In the latter case,
osmotic stress was associated with toxic effect of salts
(salinity). In these solutions, the seedlings were incu
bated for 24 h.
Lectins were extracted from whole 7dold seed
lings after 24hlong exposure to stress and subsequent
24hlong adaptation.
Lectin extracts were described by quantitative (lec
tin activity) and qualitative (carbohydrate specificity)
indices.

Abbreviations: LAlectin activity.

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LEVCHUK et al.

Lectins were extracted from various cell fractions

(membranes, soluble cytoplasm, and cell walls)
according to standard protocols [9, 16] with some
modifications: cellular fractions were separated by
centrifugation, lectins were isolated with 0.02 M
potassiumphosphate buffer at pH 6.8 with the addi
tion of 0.35 M sucrose, 0.1 M ascorbic acid, and 0.01
M Trilon B.
A seedling sample (0.5 g) was homogenized with 8
ml of 20 mM potassiumphosphate buffer (pH 6.8). In
order to separate cell walls, the suspension was
squeezed through two layers of linen cloth. The cell
wall pellet remaining on the cloth after squeezing was
homogenized with 5 mL of 20 mM potassiumphos
phate buffer (pH 4.0) supplemented with Triton X100
detergent to 0.05%, left for extraction for 12 h at 4,
and then centrifuged for 15 min at 10000 g.
Supernatant (the fraction enriched in cell wall lec
tins) was used for subsequent analysis. In order to iso
late organelles, the suspension remaining after separa
tion of cell walls was centrifuged at 10000 g for 15 min;
the pellet contained nuclei, mitochondria, and plas
tids, and supernatantpredominantly cytosol and
vacuolar sap. In order to isolate membranes from the
organelle pellet, it was homogenized with 0.5 mL of
initial potassiumphosphate buffer (pH 6.8) with the
addition of Triton X100 to 0.05% and left for extrac
tion for 20 min. The suspension was centrifuged simi
larly; the resulting supernatant contained membrane
lectins. Solubilized lectins of all the fractions were
concentrated with 70% ammonium sulphate, and the
pellets were dissolved in 0.4 mL of 0.9% sodium chlo
ride. This lectin solution was additionally purified by
precipitating with acetone to 60% saturation (two
parts of acetone per part of lectin solution). Lectins are
resistant to such high acetone concentration, and
when the pellet is later dissolved in physiological solu
tion, lectins restore their initial structure. Other pro
teins are irreversibly denatured by such acetone con
centration [4].
Lectin activity was determined by hemagglutina
tion with 2% suspension of rabbit erythrocytes [4].
Protein concentration was determined spectrophoto
metrically according to WarburgChristian [17]. Lec
tin activity (LA) was expressed as 1/LA coefficient in
(g/mL)1. Carbohydrate specificity was determined
by the suppression of hemagglutination with individ
ual carbohydrates [4]. Investigations were repeated
five times, and the results were processed using stan
dard statistical methods [18].
RESULTS
Earlier we demonstrated that seedlings of various
genotypes of oilseed flax differed by the extent of their
adaptation to particular abiotic stress factors. For
instance, it was shown that cvs. Antares, Tsian, and

Aisberg were able to adapt themselves to low tempera

ture; cvs. Aisberg, Astral, and Zolotistyi were resistant
to high temperature; cvs. Tsian and Zolotistyito
salinity. At the same time, a pronounced adaptation to
osmotic stress was characteristic of all the investigated
cultivars except for cv. Avangard [15].
We found that in response to abiotic stress, the
coefficient of lectin activity of the seedlings (1/LA)
considerably changed; at the same time, these changes
depended on the type of stress action, location of lec
tins in the cell, and the level of genotype resistance to
a certain stress factor.
In the course of low temperature acclimation of the
seedlings of coldresistant genotypes (cvs. Antares,
Tsian, and Aisberg), the coefficient of cell wall LA
notably increased: from 300 times in cv. Antares to
1000 times in cvs. Aisberg and Tsian (Fig. 1). At the
same time, in the seedlings of nonresistant genotypes
(Avangard, Astral, and Zolotistyi) LA coefficient con
siderably decreased. For instance, in the flax seedlings
of cvs. Avangard and Zolotistyi, 1/LA decreased five
fold, and in cv. Astralby 30 times as compared with
the control level (Fig. 1). In case of exposure to other
stress factors (hyperthermia, osmotic stress, and
osmotic stress associated with salinity), changes in LA
coefficient did not depend on the level of resistance. At
high temperature, LA coefficient in the seedlings of
cvs. Zolotistyi, Tsian, and Avangard increased, and in
the seedlings of other genotypes decreased. At the
same time, cvs. Astral, Aisberg, and Zolotistyi turned
out resistant to this type of stress (Fig. 2). Under
osmotic stress and salinity, coefficient of cell wall LA
increased in the seedlings of all investigated genotypes
irrespective of their level of resistance (Figs. 3 and 4,
respectively).
In the course of adaptation to all abiotic stress fac
tors under study, except hypothermia, the coefficient
of membrane LA considerably increased. This rela
tionship was characteristic only of resistant genotypes.
For instance, at high temperature the coefficient of
membrane LA in resistant cvs. Aisberg, Astral, and
Zolotistyi increased by 2 to 15 times depending on
genotype, whereas in nonresistant cultivars, it
decreased by 2100 times (Fig. 2).
The similar pattern was observed under osmotic
stress produced by sucrose added to 15% and salinity
associated with osmotic stress produced by NaCl
added 3% (Figs. 3 and 4, respectively). Under osmotic
stress, the coefficient of membrane LA increased from
3 to 15 times in all the genotypes except cv. Avangard
where LA coefficient decreased almost twofold as
compared with the control plants (Fig. 3). At the same
time, it was the only cultivar that did not show adapta
tion to this stress [11].
When salinity was associated with osmotic stress,
the elevation of activity of membrane lectins was
observed only in two genotypes (cvs. Tsian and Zolo

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LECTINS OF OILSEED FLAX PLANTS EXPOSED TO ABIOTIC STRESS

79

Increase in LA coefficient,
times

10000
1000

100
2

10
1

0.1

0.01

Antares Avangard

Tsian

Aisberg

Astral

Zolotistyi

Increase in LA coefficient,
times

Fig. 1. Changes in activity of total lectins of different cellular location in the seedlings of oilseed flax exposed to low temperature.
(1) Membrane lectins; (2) soluble lectins; (3) cell wall lectins.

100

10
1

0.1

0.01

Antares Avangard

Tsian

Aisberg

Astral

Zolotistyi

Fig. 2. Changes in activity of total lectins of different cellular location in the seedlings of oilseed flax exposed to high temperature.
(1) Membrane lectins; (2) soluble lectins; (3) cell wall lectins.

tistyi) where LA coefficient rose 3 and 12fold,

respectively, as compared with the control (Fig. 4).
These particular genotypes were earlier recognized as
the cultivars with a high adaptive potential regarding
salinity [15].
The coefficient of activity of soluble lectins
changed under stress; however, these changes did not
depend on the resistance of seedlings of the examined
genotypes (Figs. 14).
Thus, the obtained results show that there is some
correlation between resistance and LA coefficient. For
instance, in the resistant genotypes exposed to low
temperatures, the coefficient of cell wall lectin activity
considerably rose, and at high temperatures, osmotic
stress, and salinity, the coefficient of activity of mem
brane lectins increased.
In the course of adaptation of oilseed flax, the
qualitative characteristics of lectins also changed but
these changes were only observed in the fraction of sol
uble lectins. Same as lectin activity, carbohydrate
specificity of total lectin preparations were related to
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the extent of resistance (changes were observed only in

adapted genotypes) and on the type of stress effect (see
tables 14).
Table 1 shows that in the seedlings of all the geno
types resistant to hypothermia, the range of carbohy
drate specificity of total lectin preparations expanded
in the course of adaptation, and we observed the
capacity for recognizing fructose and lactose and in
the seedlings of cv. Tsianalso glucose, mannose, and
xylose.
In the genotypes resistant to hyperthermia (Table 2)
the range of carbohydrate specificity also changed,
and these total lectin preparations recognized and
bound glucose and glucosamine.
In the course of adaptation to osmotic stress, total
soluble lectins of resistant cultivars manifested the rec
ognition and binding mannose and xylose (Table 3); in
saltresistant genotypes exposed to this type of stress in
the course of adaptation, total soluble lectins
responded to galactose, glucose, and glucosamine
(Table 4).
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LEVCHUK et al.

Increase in LA coefficient,
times

10000
1000
100
10

0.1

Antares Avangard

Tsian

Aisberg

Astral

Zolotistyi

Fig. 3. Changes in activity of total lectins of different cellular location in the seedlings of oilseed flax exposed to osmotic stress.
(1) Membrane lectins; (2) soluble lectins; (3) cell wall lectins.

Changes in the coefficient

of lectin activity as compared
with control material, times

10000
1000

100
10
1

3
1

0.1
0.01

Antares Avangard

Tsian

Aisberg

Astral

Zolotistyi

Fig. 4. Changes in activity of total lectins of different cellular location in the seedlings of oilseed flax exposed to salinity associ
ated with osmotic stress.
(1) Membrane lectins; (2) soluble lectins; (3) cell wall lectins.

DISCUSSION
Comparison of the levels of lectin activity in the
seedlings with the extent of adaptation of different flax
genotypes shows the following relationships.
(1) The coefficient of membrane lectin activity when
plants are affected by high temperature, osmotic stress
and salinity, considerably increased (by 215 times) in
the seedlings of adapted flax cultivars and decreased by
2100 times in undated seedlings. (2) Low tempera
ture considerably increased LA coefficient of cell walls
(by 3001000 times) in the seedlings of adapted flax
cultivars (Antares, Tsian, and Aisberg) and reduced it
by 530 times in nonadapted seedlings. (3) The coef
ficient of soluble lectin activity changed under the
effect of stresses independently on plant adaptation.
Thus, the adaptation of oilseed flax seedlings to
abiotic stress depends on lectins located in different
parts of the cell. In response to stress, the range of car
bohydrate specificity of total soluble lectins changed,
whereas the coefficient of activity of membrane lectins

and the lectins of cell walls rose. At the same time, cell
wall lectins responded to hypothermia and membrane
lectinsto other abiotic stresses: hyperthermia,
osmotic stress, and salinity. All the detected changes
were observed only in the seedlings of resistant geno
types.
Our data on the participation of cell wall lectins of
flax seedlings in cold acclimation are corroborated by
other researchers. For instance, one of the possible
causes for cold resistance of winter wheat plants
depends on the ability of cortical microtubules to pre
serve their native state [19]. An important role in this
phenomenon belongs to extensin that shows LA and
therefore ensures interaction between cell wall and
microtubules. In this relation, it is assumed that an
increase in the activity of lectins of the cell wall caused
by disturbance of its integrity is a compensatory mech
anism ensuring stabilization of depolymerized cytosk
eletal structures governed on genetic level [16, 19].

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81

Table 1. Carbohydrate specificity of total soluble lectins in coldresistant genotypes of oilseed flax
Carbohydrate

Type
of treatment

gal

No treatment
Hypothermia

+
+

No treatment
Hypothermia

glu

man

xyl

No treatment
Hypothermia

ara
Antares
+
+
Tsian
+
+
Aisberg
+
+

malt

+
+

suc

lact

fru

++

++

++

++

++

++

glA

+
+

Note: Carbohydrate specificity was tested using 0.6 M solutions of carbohydrates. (malt) maltose; (lact) lactose; (glA) glucosamine. Plus
designates suppression of hemagglutination by the specified carbohydrate. Empty cells show that the suppression was not
observed. Double plus designates broadening the range of carbohydrate specificity under the effect of stress as compared to the
control.

Table 2. Carbohydrate specificity of total soluble lectins of heatresistant genotypes of oilseed flax
Type
of treatment

Carbohydrate
gal

glu

man

xyl

ara

malt

suc

lact

fru

glA

Aisberg
No treatment
Hyperthermia

++

++

Astral
No treatment
Hyperthermia

++

++

Zolotistyi
+

No treatment
Hyperthermia

++

++

Note: Conditions and designations see in Table 1.

There are publications concerning the participa

tion of membrane lectins in formation of cold resis
tance of wheat [16]; however, in the case of flax plants
we did not observe such participation, probably due to
particular crop physiology.
Lectins are known to participate in the develop
ment of resistance to different types of stress. However,
lectins are believed to be components of nonspecific
resistance [6, 10]. In our case, lectins of different cel
lular fractions differently responded to particular types
of stress. For instance, low temperatures induced a
considerable rise in the coefficient of cell wall lectin
activity, while osmotic stress, salinity or high tempera
ture activated membrane lectins.
One can assume that the development of resistance
to various abiotic stress factors depends on lectins
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located in different parts of the cell because the range

of carbohydrate specificity of total lectins of cytosol
and vacuolar sap changed under all types of stress. In
the course of acclimation to hyperthermia, total solu
ble lectins of flax seedlings manifested capacity for
binding glucose and glucosamine, in the course of
adaptation to osmotic stress, mannose and xylose, to
salinity, galactose, glucose, and glucosamine, and
upon the formation of cold resistance, lactose and
fructose. In membrane lectins, coefficient of activity
rose upon the effect of hyperthermia, osmotic stress
and salinity associated with osmotic stress; in cell wall
lectins, upon the effect hypothermia. Therefore, one
can believe that the lectins of oilseed flax directly par
ticipate in the development of resistance to abiotic
stress by modifying their quantitative and qualitative
characteristics. Moreover, one can suggest simulta
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LEVCHUK et al.

Table 3. Carbohydrate specificity of total soluble lectins in osmotically resistant genotypes of oilproducing flax
Carbohydrate
Type of treatment
gal

glu

man

xyl

ara

malt

suc

+
+

lact

fru

+
+

+
+

+
+

+
+

glA

Antares
No treatment
Osmotic stress

+
+

No treatment
Osmotic stress

++

+
+

++
Tsian

+
+

++

++
Aisberg

No treatment
Osmotic stress

++

++
Astral

No treatment
Osmotic stress

++

++
Zolotistyi

No treatment
Osmotic stress

++

+
+

+
+

+
+
+
+

++

Note: Conditions and designations see in Table 1.

Table 4. Carbohydrate specificity of total soluble lectins of saltresistant genotypes of oilproducing flax
Type
of treatment

Carbohydrate
gal

glu

man

xyl

ara

malt

suc

lact

fru

glA

Tsian
No treatment
Salinity

++

++

++

Zolotistyi
No treatment
Salinity

+
++

++

++

Note: Conditions and designations see in Table 1.

neous but nonequivalent participation of various lec

tins in the process of adaptation of flax seedlings. For
instance, in total soluble lectins, qualitative character
istic (the range of carbohydrate specificity) changed,
whereas in the lectins of cell walls and membranes,
quantitative characteristic (LA coefficient) altered.
Thus, lectins may be the components of specific
adaptation of flax plants to abiotic stress factors of dif
ferent nature. Depending on the type of stress, adapta
tion of resistant genotypes was associated with eleva
tion of the coefficient of activity of membrane lectins
or the lectins of cell walls. Total lectins of soluble part
of cytoplasm were found to recognize and bind new
carbohydrates, which is a prerequisite for adaptation

because under such conditions the composition of

sugars on cellular surfaces changed.
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