Метод імітації водного стресу на Arabidopsis thaliana L.
A method for the imitation of drought stress in Arabidopsis thaliana L. combining the advantages of rapid and prolonged treatments was developed. The proposed method allows decreasing water content in the growth medium gradually and homogenously, growing plants under sterile conditions, and observin...
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| Дата: | 2006 |
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M.M. Gryshko National Botanical Garden of the NAS of Ukraine
2006
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Plant Introduction| _version_ | 1860124417446641664 |
|---|---|
| author | Bobrownyzky, J. |
| author_facet | Bobrownyzky, J. |
| author_sort | Bobrownyzky, J. |
| baseUrl_str | https://www.plantintroduction.org/index.php/pi/oai |
| collection | OJS |
| datestamp_date | 2019-12-28T11:07:04Z |
| description | A method for the imitation of drought stress in Arabidopsis thaliana L. combining the advantages of rapid and prolonged treatments was developed. The proposed method allows decreasing water content in the growth medium gradually and homogenously, growing plants under sterile conditions, and observing a root system during the growth. In the course of experiment, gradual growth retardation of stressed plants was observed. The rates of cell division and cell elongation of the tap root were measured and it was defined that cell division and cell elongation are both responsible for growth retardation. Morphological changes in the roots of stressed plants were described. |
| doi_str_mv | 10.5281/zenodo.2567226 |
| first_indexed | 2025-07-17T12:46:55Z |
| format | Article |
| fulltext |
98 ISSN 1605�6574. Інтродукція рослин, 2006, № 1
Plant growth is greatly affected by environ%
mental abiotic stresses such as drought, high
salinity and low temperature. These stresses
induce various biochemical and physiological
responses in plants to acquire stress tole%
rance. Abiotic stresses are severe limiting
factors of plant growth and crop production
[31, 22, 23]. Among these abiotic stresses,
drought or water deficit is the most severe
limiting factor of plant growth and crop pro%
duction [22, 11]. It causes 24 million tons of
yield loss in maize annually by inhibiting
plant growth and photosynthesis [25]. Typi%
cally, however, roots are affected less then
shoots. In fact, even under mild water deficit,
shoots may stop growing completely while
roots continue to grow. Continued root
growth allows plant to plumb soil for water
and can be especially important for seedling
establishment [31].
Although the responses of plants to water
deficit have been studied in many species, we
thought to use Arabidopsis thaliana L. to take
advantage of the potent molecular and genetic
tools available for this species [31, 35]. Its many
advantages include a small genome, short life
cycle, small stature, prolific seed production,
and ease of transformation. In addition, a
wealth of genomic resources exists, such as a
completely sequenced genome, a near satura%
tion insertion mutant collection, a genome
array that contains the entire transcriptome,
and more then 50,000 molecular markers [35].
In addition, roots of A. thaliana seedlings have
a well%defined anatomy [31].
In the last years a number of methods have
been worked out, which allow imitating con%
ditions of natural drought stress. These treat%
ments could be divided into two big groups:
prolonged [30] and rapid treatments [9, 26, 34].
In the case of prolonged treatments, stress
action lasts from several days to several
weeks. Plants are grown on sand or soil, and
desiccation occurs through ceasing of wate%
ring. Stress conditions are more similar to tho%
se of natural water stress. Another advantage
is the possibility to observe long%term mor%
phological changes. At the same time difficul%
ties are possible by reliable determination of
the humidity of sand or soil and hence it may
be problematic to estimate the stage of stress.
Root system can't be observed in vivo, and it
could be complicated to isolate whole plants
including roots for analysis. As a result of
using of non%sterile growing substrate, ste%
rility is not maintained.
Taking into account disadvantages of pro%
longed treatments, another group of methods
has been established and widely used. We
called them rapid treatments because stress
УДК 58.032.3
J. BOBROWNYZKY
M.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine,
Ukraine, 01601 Kyiv, 2 Tereshchenkiwska St.
A METHOD FOR THE IMITATION
OF DROUGHT STRESS IN ARABIDOPSIS THALIANA L.
A method for the imitation of drought stress in Arabidopsis thaliana L. combining the advantages of rapid and prolonged
treatments was developed. The proposed method allows decreasing water content in the growth medium gradually and
homogenously, growing plants under sterile conditions, and observing a root system during the growth. In the course of
experiment, gradual growth retardation of stressed plants was observed. The rates of cell division and cell elongation of the
tap root were measured and it was defined that cell division and cell elongation are both responsible for growth retardation.
Morphological changes in the roots of stressed plants were described.
© J. BOBROWNYZKY, 2006
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action doesn't exceed several hours. Desicca%
tion occurs through removing of plants from
growth medium and putting them on filter
paper under controlled temperature and hu%
midity. These methods are brief and reliable;
whole plants can easily be separated from
medium and taken for biochemical as well as
for microscopic analysis. At the same time,
they are very simplified imitation of natural
water stress; duration of stress doesn't allow
observing long%term morphological changes.
Despite the advantages of A. thaliana, to
our knowledge only few attempts were under%
taken to imitate conditions of prolonged water
stress on plants growing on sterile agar medi%
um [26]. Unfortunately, our attempts to repeat
these experiments resulted in unequal drying
of medium and contaminations. Nutrient%agar
media are often used in studies of A. thaliana,
and hence a large amount of physiological data
has been obtained for these conditions [31].
Therefore, the aim of this study was to deve%
lop a method to grow A. thaliana seedlings at a
lowering water potential on an agar%solidified
medium; and, using this system, we charac%
terized several growth responses of A. thali�
ana seedlings to water deficit.
Materials & Methods
Plant Material
Arabidopsis thaliana L. line used in this study
was Columbia wild type. The CycB1; 1: CDB:
:GUS marker line was kindly provided by
John Celenza.
Plant Growth Conditions
A. thaliana seeds were sterilized in 5% NaClO
for 3 minutes, washed 3 times in sterile water,
and distributed in 1% low melt agarose. After%
wards the seeds were sown onto the standard
plastic agar plates containing 1x Murashige
and Skoog (MS) salt mixture supplemented
with 4.5% sugar, pH 5.7, and left to imbibe 2
days at 4° in the dark. The plants were then
cultivated in a growth chamber in a near ver%
tical position at 22 °C, and 16%h light (80 µmol
m%2s%1) /8h dark cycle till the age of 6 days
which corresponds to the stage of fully deve%
loped cotyledons. Then, the plants were trans%
ferred onto plates with low MS concentration
(0.25x) and without sugar. Sucrose, often com%
pared to a hormone, was added to the medium
because it promotes plant growth [1, 31]. In the
experiments with the medium lacking sugar,
germination and growth were irregular to
such an extent that it was difficult to select
enough uniform seedlings for transplanting
[31]. At the same time, to minimise the conse%
quences of osmotic stress, we used a second
medium without sugar and with the low con%
centration of mineral nutrients (0,25x MS).
Additionally, the plates intended for the
modulation of water stress conditions were
modified. Firstly, the amount of medium was
fixed to 80 ml. Then, the medium was poured
under determined angle so that the thickness
of agar layer on the top of the plate is less as
on its bottom, and hence the drying gradient
is created. Finally, the standard plate plastic
cover was replaced with cellophane foil (Roth,
Germany), which is transparent for water
vapour. Control plates were standard i.e. they
contained 0.25x MS 40 ml each and were cov%
ered with usual plastic cover. Plants were
viable under these conditions till the age of
5—6 days after transfer.
Measurement of Substrate Moisture
The small pieces of agar substrate were taken
from the surface of angle plates, which were
placed into growth chamber, at the level of 2,
6, and 10 cm from the top of the plate. Then,
the agar pieces were dried for 4 h at 70 °C,
and the dry weight was calculated.
GUS�Staining
The mitotic activity of A.thaliana roots was
investigated by using the cycB1;1:CDB:GUS
marker line. The plants were removed from
agar every day after transfer, submerged in
staining solution (0.1 M NaPO4 pH 7.0, 0.01 M
EDTA, 0.01% (v/v) Triton X%100, 0.5 mM
K3[Fe(CN)6], 0.5 mM K4[Fe(CN)6] and 0.05%
(w/v) X%Gluc substrate solved in dimethyl%
formamide). Samples were mounted on mic%
ro%scopic slides and incubated at 37 °C for 2
hours. The number of dividing cells in the
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meristematic zone of the main root was
counted under the light microscope.
Imaging of Roots and Cell Length Measurement
Images of roots of growing plants were cap%
tured with a Leica DC500 digital camera. The
control and stressed plants were harvested at
the age of 5 days after transfer, fixed in a
mixture of methanol and acetic acid (3:1, v/v).
The samples were mounted on the micro%
scopic slides in mounting solution (76% (w/v)
chloralhydrate, 19% (v/v) glycerine 87%). For
the measurement the fully elongated epi%
dermal cells were taken, which do not form
root hairs (atrichoblasts) along the entire
main root, starting at 2mm from the distal end
of the main root and finishing at the point
where transfer occured. This point could be
easily seen by the abscence of the root hairs,
which were lost at the moment of transfer.
The cells were measured using micrometric
ocular (10×). One randomly selected cell
within an interval of 0,5 mm was measured.
The measurements were taken using a Zeiss
Axiowert 35 microscope.
Results and Discussion
System for observation of growth of A. thaliana
plants upon slow desiccation stress
Arabidopsis thaliana plants grown till the age
of 6 days on the full MS medium supple%
mented with 4.5 % sugar under standard con%
ditions were subjected to slow desiccation
stress by transferring on the plates specially
constructed for this purpose. We measured
the dry weight of the substrate at 2, 6 and 10
cm from the top of the angle plate (Fig. 1).
This resulted in a progressive depletion of
water in the substrate. From the onset of the
drought conditions (0 d), the dry weight of
substrate increased from the initial value of
1,1% to 100% (no water) within 5, 6 and 7 days
at the level of 2, 6 and 10 cm from the top of
the plate, respectively. The character of
curve testifies the loss of approximately
equal portions of water day after day.
As a result of water loss, the stressed plants
were visually much more retarded in compa%
rison to control. To gain further insight into the
response of primary root elongation to water
deficit, the kinetics of the response during the
5 d exposure was studied. As shown on Fig. 2, in
the control plants, soon after transfer, we ob%
served the subsequent activation of growth,
return to the log%phase, and reaching of the
plateau. In contrast, the stressed plants have
undergone after the period of slight activation
on 2 d and 3 d the phase of slow subsequent
retardation of growth followed by death on 6 d
as a consequence of total water loss. Under
Fig. 1. Change of substrate humidity of an angle plate
Fig. 2. Change of length of main root of plants grown
under stress conditions and in control. Symbols indi�
cate: K — control plants; S — plants subjected to slow
desiccation stress. Bars indicate standard deviations
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stress conditions, the total elongation of the
main root at the end of the 5 d treatment was
approx. 6 times less as in the control.
Most studies have found that water deficit
inhibits primary root elongation rate [10, 13,
16, 27, 31]. Stimulation has been shown for
Pinus pinaster [18, 29], Glycine max (L.) Merr.
[5], and Arabidopsis thaliana [31]. In our expe%
riment, the decrease of primary root growth
rate is more pronounced at the later stages of
stress, which is in accordance with an obser%
vation that at moderate stress the root elon%
gation increases whereas at severe water
stress it decreases [31]. As described above,
the desiccation stress results in a progressive
retardation of root growth. It is well known
that adverse conditions inhibit root growth
and that cell division and cell cycle regulation
are involved in this response [3, 12, 20, 21, 33].
To investigate whether this phenomenon is a
result of more rare division of meristematic
cells or it reflects the lower level of cell elon%
gation, we undertook two experiments.
In course of the first one (Fig. 3) A. thaliana
plants (ecotype Columbia) were fixed on the 5
d and observed in light microscope. We mea%
sured the length of mature epidermal cells,
which do not produce root hairs, so called atri%
choblasts. Cells were measured only in the
part of the main root, which had grown after
the transfer. This "post%transfer" distal part
of the root could easily be distinguished from
the "pre%transfer" one by the presence of root
hairs in it. At first we expected the change of
a cell length corresponding to the phase of
growth, e.g. longer cells at the beginning of
stress action and shorter at its end. Neverthe%
less, no considerable change of cell length
within two groups could be observed, though
elongation rates changed considerably in both
groups. Cell length in the control increased
quite slightly compared with stressed plants.
At the same time, this difference was not sig%
nificant, and it could not be responsible for
much more strong elongation of a main root of
control plants. Therefore, a length of epider%
mal cells seems to be parameter, which should
be regulated to fall within a preferred range.
This allows us to presume that the changes
in division of root meristematic cells are
responsible for changes in root growth rate
and hence for the reaction of plants to the
conditions of the slow desiccation stress. In
order to check this hypothesis, we investi%
gated the mitotic activity using the plants of a
transgenic marker line Cyc B1;1: CDB: GUS,
which were expressing the labile translational
fusion protein between the mitotic cyclin B1
and the β%glucuronidase. The promoter and
the region coding the first 150 amino acids of
A. thaliana cyclin B1 were combined with the
Fig. 4. Number of dividing cells in the root meristem
of plants grown under stress conditions and in con%
trol. Symbols indicate: K— control plants; S — plants
subjected to slow desiccation stress. Bars indicate
standard deviations
Fig. 3. Length of mature root epidermal cells in con%
trol and under slow desiccation stress. Symbols indi�
cate: K — control plants; S— plants subjected to slow
desiccation stress. Bars indicate standard deviations
102 ISSN 1605�6574. Інтродукція рослин, 2006, № 1
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β%glucuronidase (GUS) gene. The expressed
chimeric protein contains the destruction box
(CDB), which resides between amino acid 30
and 38 of cyclin B1. Owing to a mitotic degra%
dation signal in the protein, reporter gene
activity marks only actively dividing cells [4,
15, 32]. The plants were daily taken from agar,
stained, and the number of stained cells was
measured (Fig. 4). Parallel we carried out the
measurement of root elongation of transgenic
plants, which was similar to the root elonga%
tion of Columbia wild type plants (not shown).
In the control group, the division of meriste%
matic cells of main root changed over time in
parallel with an elongation rate. The same
goes for the group of stressed plants with one
noteworthy exception: division rates on 3 d,
4 d and 5 d were much higher as it could be
expected from their elongation rates. This
may be due to a fact that root elongation is
always a result of previous cell divisions.
To our knowledge, in A. thaliana, the chan%
ge in root elongation rate is paralleled by the
change in cell production rate. Our data pro%
vide another support for this hypothesis. In
species where water deficit similarly inhi%
bited primary root elongation rate, cell pro%
duction rate was similarly inhibited [6, 8].
This suggests that the root's elongation rate
at given levels of water deficit is determined
principally by the supply of cells to the zone
of rapid elongation [2, 31].
The most sensitive cells of a root might be
root hairs. They display a tendency to a big
morphological plasticity. Under slow desicca%
tion stress, the roots exhibited hair decay
(Fig. 5B), which was stronger under more
severe stress conditions. Typical for the root
hairs of stressed plants are also such abnor%
malities, as root hairs with vesicular exten%
sion on their distal end and root hairs with
bifurcations (Fig. 5C, D). To our knowledge,
this is the first report of the enhanced pro%
duction of root hair branching under condi%
tions of water deficit. Bifurcated root hairs is
one of the numerous root hair deformations,
which is promoted by several treatments,
such as influence of Nod factors (bacteria%to%
plant signalling) [7] and Fe%deficiency [17].
This response to signal occurs at a specific
developmental stage, namely when hairs are
terminating growth [14]. At the final stages of
the slow desiccation stress, the roots exhibit
curling of their distal end (Fig. 5B). The simi%
lar phenomenon was already observed [28] in
the experiments with gel drying. This may be
an indication that the drying of gel surface
increases its sticking capacity, which in its
turn is capable of slowing the movement of
A B
C D
Fig. 5. Root phenotypes. Bars: 1 mm in A and B and
100 µm in C and D.
A — Control plant. Regular distribution of root hairs
could be observed; B—D — plants grown under
stress conditions; B — bending of root tip. Root hairs
in the area of tip bending have peculiar shape, which
may be result of turgor loss in the distal part of the
root. Irregular distribution of root hairs and hairless
areas could also be observed; C — root hair with
vesicular extension at its distal end (indicated by
Arrow); D — root hair branching. Arrows indicate
position of root hairs with bifurcations
root tip, and causing deflections as a result of
elongation. In association with curling the
"crumpled" root hairs could be also observed,
which may be due to lower turgid pressure at
the distal end of the root.
In this work we presented a new method
of slow desiccation stress, which combines
the advantages of rapid [9, 26, 34] and pro%
longed [30] treatments. This method allowed
us to observe gradual retardation of growth
and other effects of drought stress action
(bifurcated root hairs, root tip bending, root
hair loss).
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laboratory to field. Using information from Arabidopsis
to engineer salt, cold, and drought tolerance in crops //
Plant Physiol. — 2004. — 135, N 1. — P. 1—7.
Recommended to publication by B.O. Levenko
Ю. Бобровницький
Інститут ботаніки ім. М.Г. Холодного НАН України,
Україна, м. Київ
МЕТОД ІМІТАЦІЇ ВОДНОГО СТРЕСУ
НА ARABIDOPSIS THALIANA L.
Автором розроблено метод з імітації водного стресу
на проростках Arabidopsis thaliana L., який дає змо%
гу знижувати вміст води в середовищі поступово і
рівномірно, вирощувати рослини в стерильних умо%
вах і вивчати кореневу систему під час росту. В
процесі експерименту відмічене поступове пригні%
чення росту рослин в умовах водного стресу. Вста%
новлено, що поділ та елонгація клітин є відпові%
дальними за пригнічення росту рослин порівняно з
контролем. Описано морфологічні зміни в коренях
рослин в умовах водного стресу.
Ю. Бобровницкий
Институт ботаники им. Н.Г. Холодного НАН Украины,
Украина, г. Киев
МЕТОД ИМИТАЦИИ ВОДНОГО СТРЕССА
НА ARABIDOPSIS THALIANA L.
Автором разработан метод имитации водного стрес%
са на проростках Arabidopsis thaliana L., позволяю%
щий снижать содержание воды в среде медленно и
равномерно, выращивать растения в стерильных
условиях и изучать корневую систему во время
роста. В процессе эксперимента отмечено посте%
пенное угнетение роста растений в условиях водно%
го стресса. Установлено, что деление и удлинение
клеток являются ответственными за угнетение рос%
та растений по сравнению с контролем. Описаны
морфологические изменения в корнях растений в
условиях водного стресса.
104 ISSN 1605�6574. Інтродукція рослин, 2006, № 1
JJ.. BBoobbrroowwnnyyzzkkyy
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| id | oai:ojs2.plantintroduction.org:article-905 |
| institution | Plant Introduction |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2025-07-17T12:46:55Z |
| publishDate | 2006 |
| publisher | M.M. Gryshko National Botanical Garden of the NAS of Ukraine |
| record_format | ojs |
| resource_txt_mv | wwwplantintroductionorg/0f/f505fe62aa1c37561dcbc3700a39290f.pdf |
| spelling | oai:ojs2.plantintroduction.org:article-9052019-12-28T11:07:04Z A method for the imitation of drought stress in Arabidopsis thaliana L. Метод імітації водного стресу на Arabidopsis thaliana L. Bobrownyzky, J. A method for the imitation of drought stress in Arabidopsis thaliana L. combining the advantages of rapid and prolonged treatments was developed. The proposed method allows decreasing water content in the growth medium gradually and homogenously, growing plants under sterile conditions, and observing a root system during the growth. In the course of experiment, gradual growth retardation of stressed plants was observed. The rates of cell division and cell elongation of the tap root were measured and it was defined that cell division and cell elongation are both responsible for growth retardation. Morphological changes in the roots of stressed plants were described. Автором розроблено метод з імітації водного стресу на проростках Arabidopsis thaliana L., який дає змогу знижувати вміст води в середовищі поступово і рівномірно, вирощувати рослини в стерильних умовах і вивчати кореневу систему під час росту. В процесі експерименту відмічене поступове пригнічення росту рослин в умовах водного стресу. Встановлено, що поділ та елонгація клітин є відповідальними за пригнічення росту рослин порівняно з контролем. Описано морфологічні зміни в коренях рослин в умовах водного стресу. M.M. Gryshko National Botanical Garden of the NAS of Ukraine 2006-03-01 Article Article application/pdf https://www.plantintroduction.org/index.php/pi/article/view/905 10.5281/zenodo.2567226 Plant Introduction; Vol 29 (2006); 98-104 Інтродукція Рослин; Том 29 (2006); 98-104 2663-290X 1605-6574 10.5281/zenodo.3377827 en https://www.plantintroduction.org/index.php/pi/article/view/905/868 http://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Bobrownyzky, J. Метод імітації водного стресу на Arabidopsis thaliana L. |
| title | Метод імітації водного стресу на Arabidopsis thaliana L. |
| title_alt | A method for the imitation of drought stress in Arabidopsis thaliana L. |
| title_full | Метод імітації водного стресу на Arabidopsis thaliana L. |
| title_fullStr | Метод імітації водного стресу на Arabidopsis thaliana L. |
| title_full_unstemmed | Метод імітації водного стресу на Arabidopsis thaliana L. |
| title_short | Метод імітації водного стресу на Arabidopsis thaliana L. |
| title_sort | метод імітації водного стресу на arabidopsis thaliana l. |
| url | https://www.plantintroduction.org/index.php/pi/article/view/905 |
| work_keys_str_mv | AT bobrownyzkyj amethodfortheimitationofdroughtstressinarabidopsisthalianal AT bobrownyzkyj metodímítacíívodnogostresunaarabidopsisthalianal AT bobrownyzkyj methodfortheimitationofdroughtstressinarabidopsisthalianal |