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��ࡱ�>��	�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������'`"���bjbj��;�r�r�6�^�������:):):):)z6z6z6>9tz|z|z|z|�^t�DTH��^<�<�<�<���B���WEYEYEYEYEYEYE$0Mh�O�}E�z6ɜ݅���ɜɜ}E:):)<�<��H������ɜ�K:)�	<�z6<�WE��ɜWE����._7H�2�z6�:<��p߮���z|����9@cB�$H0TH�9<P[�<P��:�:�<Pz6�;|!�������S����!�!�!�}E}Ey�!�!�!�THɜɜɜɜ�D�D�D�"Vg$�D�D�DVg�6�28:9:):):):):):)����Introduction  
      To understand how stability of plant ontogenesis is maintained in the variable environment remains a paramount task of theoretical and experimental biology, especially this century. There is no doubt that stability, i.e. realization of the genetically determined program of ontogenesis, is under genetic control. Nevertheless, plants survive and reproduce under the unfavorable fluctuations of ecological factors due to plasticity of their organization that is known as phenotypic plasticity, i.e. a genome ability to alter its expression and be realized in different phenotypes in response to various environmental impacts. Phenotypic manifestations of the changes in gene expression are already defined at the level of transcription efficiency and also RNA processing and translation [1] and include a very broad spectrum of ecologically important traits � physiological, biochemical, anatomical and morphological, peculiarities of developmental biology, time of transition to the reproductive stage, propagation systems and progeny development. The wide theoretical and experimental investigations of phenotypic plasticity at the population and interspecies levels have been carried out in order to elucidate its importance in evolution, specialization, population dynamics and a strategy of survival in the heterogenic environment [2�10]. 
      In response to drought, flooding, extreme temperature, soil salinity, high light intensity, infection with pathogenic agents, structural and metabolic processes, which counteract stress, occur in plants. The water content in soil  significantly varies in the sites of plant habitation depending on a soil type and topography, climatic and meteorological conditions. In the same time, as it is known, water accessibility for plants is a critical external factor for their growth and development, and drought is one of the major abiotic stresses [11, 12].  In this respect, the species of the same genera growing under different water regime in the biotope are suitable models to examine plasticity of different traits  in plant adaptation to the environment. Investigations of plasticity in the ecological context are important for understanding the mechanisms of plant responses to abiotic and biotic factors and plant interrelations with the changing environment [5, 7].    
     We performed a comparative study of  lipid peroxidation intensity and antioxidant activity, the phytohormone content,  levels of hot stress protein synthesis and PIP aquaporin gene expression  in aerial-aquatic Sium latifolium L. and terrestrial S. sisaroideum DC. plants (Apiaceae) in response to changes in the soil water content. The objective of this work was to check the hypothesis that plasticity of structural and metabolic traits may ensure the survival of these species under variable soil moisture " from water-logging to dry aerated soil.  
Material and Methods
Field collection 
Close related species S.�latifolium  L. .known by the common name  great water-parsnip  and S. sisaroideum DC. known by the common name  skirret  are native throughout Ukraine in the forest and forest-steppe zones. They are perennial herbs, flower in June�August, bear fruit in August�September. Small and pentamerous flowers with five sepals and five petals nearly always aggregated in terminal umbels, that may be simple or more commonly compound. Fruit dry, separating into 2  carpellary units, each unit is named as a�mericarp (Fig. 1).�We investigated aerial-aquatic S.�latifolium plants growing in coastal water of the river Psjol in Poltava region of Ukraine and terrestrial plants S. sisaroideum growing on the riverside in the broad-leaved forest. Our long-term (2005�2018 years) phenological observations of these species showed that they are able to sustain well the sudden fluctuations of soil moisture, from flooding to drought during the vegetative period as a result of snowmelt flood, weather changes, or human activity. As the distance between plants growing in water and on the steep bank was only 1�5 m, it is possible to sharply determine an action factor � the water content in soil. Material was collected in the natural conditions at the stages of vegetative growth, budding�flowering and flowering-fruiting.  Leaves were sampled from different plants (three and more) under dry weather at air temperature 25-300 C and 35�400 C, frozen in liquid nitrogen at the collecting ground, preserved at -70(! and analyzed in the laboratory later.
Flooding experiments 
Experiments were carried out in the field conditions with S. sisaroideum plants which were transplanted at the budding flowering stage in pots with soil from the collection site.  For flooding, pots with plants were placed into containers with water to submerge the root system and leaf samples were collected after flooding exposure for 8, 12 and 18 h, frozen in liquid nitrogen, and stored at "70 �! before an analysis of gene expression.
Lipid peroxidation (LP) and antioxidant activity (AOA)
Registration of reactive oxygen species (ROS) generation      
 Production of ROS was measured by registration of luminol-dependent chemiluminescence (LDCL) (chemiluminometer XLM1C-01, Russia). Leaves frozen in liquid nitrogen (1 g) were ground with a precooked mortar and pestle and then 5 ml of cold 50 mM sodium phosphate buffer (pH 7.2) was added. The homogenate was filtering through four layers of cheesecloth and centrifugation at 3000 � g during 7 min. Then, the supernatant (0.5 ml) was input to a chemiluminometer cuvette with 0.8 ml of 10-2 M luminol and the chemiluminescence intensity during 5 min was recorded. The level of ROS was calculated as impulses per minute. 
Determination of LP intensity   
 The LP level was determined by measuring the content of thiobarbituric acid-reacting substances (TBARS) according to [13]. Leaf samples (0.5 g) were ground in a precooked mortar and pestle after treating with liquid nitrogen and suspended in 5 ml of 0.1% trichloroacetic acid. The homogenate was centrifuged at 10,000 � g for 10 min at 4oC and then to 1 ml of supernatant, 4 ml 20% trichloroacetic acid containing 0.5% thiobarbithuric acid was added. The mixture was heated at 95oC for 30 min and then quickly coaled. After centrifuged at 10,000 � g for 10 min, the absorbance of the supernatant at 532 and at 600 nm was read on a SP-2000 spectrophotometer (Russia). TBARS content was calculated using extinction coefficient of 155 mM-1Am-1 and expressed as nmol g"1 fresh weight. 
Assessment of total AOA
AOA was determined by the diphenylpicrylhydrasine reaction according to Glevind [14].  Leaf homogenate was prepared as for ROS determination. Then, to 0.6 ml filtered homogenate, 1.8 ml of ethanol was added and antioxidant extraction was carry out during 5 min at 95oC on water bath. Following quickly cooled extract was centrifuged at 10,000 � g for 10 min and 1.5 ml of supernatant was input to spectrophotometer cuvette with 2 ml of work solution of diphenylpicrylhydrasine. The absorbance at 517 nm was recorded. tAOA was calculated as relative unit g-1 fresh weight.
Activity test of superoxide dismutase (SOD)  
 SOD (EC 1.15.1.1) activity was assayed by measuring its ability to inhibit the photochemical reduction of NBT according to [15]. Leaf samples (0.5 g) were ground in liquid nitrogen and then 5 ml of 50 mM sodium phosphate buffer (pH 7.0) containing 1% (w/v) polyvinylpyrrolidone was added. The homogenate was filtered through four layers of cheesecloth and centrifugation at 10,000 � g for 10 min at 4oC. The supernatant was collected and 50 �l of obtained enzyme extract was input to tubes with 3 ml reaction mixture contained 63 �M nitroblue tetrasolium  (NBT), 1.3 �M riboflavin, 0.1 mM EDTA, 13 mM methionine and 50 mM phosphate buffer (pH 7.8). Then tubes were placed under fluorescent lamp at 4000 lux during 10 min and the absorbance at 560 nm was recorded. Enzyme amount corresponding to 50% inhibition of the rate of NBT reduction was considered as one enzyme unit. The SOD activity was calculated as unit g-1 fresh weight.
Detection of the lipofuscin content 
Lipofuscin content was studied according to [16]. Lipids were extracted in the chloroform-methanol-water solvent system according to [17]. Chloroform layer with lipids was collected. For separation of lipids from plant pigments, the phase division between equal volumes of hexane and 80% water methanol were carry out. After, the water methanol phase was collected and the chloroform and water were added for volume relation the chloroform-methanol-water (8:4:3). After phase separation, the chloroform layer with lipofuscins was extracted and used in experiments. Lipofuscins were determined in a %-  spectrofluorometer (Japan) with an excitation wavelength of 355 nm and an emission wavelength of 400 nm. The lipofuscin content was calculated as relative units g-1 fresh weight.
Statistical analysis      
The results were based on three biological and analytical replications of every studied stages of plant development. Student�s t-test with a probability of 95% (P = 80.05) was used for statistical significance. The correlative analysis between ROS production and LP levels was carried out according to [18].
Expression of plasma membrane intrinsic protein (PIP) aquaporins
RNA isolation  
     Total RNA was extracted using TRI-REAGENT (Sigma) according to manufacturer protocol. All used solutions were treated with diethyl pyrocarbonate and autoclaved. 100 mg of plant material was homogenized with mortar and pestle in liquid nitrogen, diluted in 1ml of TRI-REAGENT and centrifuged (12000g, 10 min, 40C). After centrifugation, supernatant was removed and pellet was washed with 1 ml of 75% ethanol. Sample was vortexed briefly, then centrifuged at 7500g for 5 minutes at 4�C. RNA pellet was air dried for 5�10 minutes and resuspended in RNAse-free water RNA. Yield was determined using absorbance of RNA at 260 nm.  
Reverse transcription-polymerase chain reaction 
     RT-PCR was carried on the FERMENTAS protocol (Lithuana) with an amplificator Terzik (DNA-technology). To obtain cDNA, total RNA with 100 pmol of oligo(dT) primer was incubated for 5 min at 650C, then to the reaction was added: 4 �l of 5x reaction buffer, 1 �l of 10 m dNTP mix, 1 �l of RNAse inhibitor,  and the mixture was incubated for 5 min at 370!. After incubation 1 �l of M-MuLV reverse transcriptase was added and the reaction was incubated for 1 hour at 420!. Reaction was terminated by heating up to 700! for 10 min. Obtained cDNA was used for PCR and 
 sequence analysis. 
     PCR was carried using the obtained cDNA and constructed degenerate primers with 20-21 nucleotides:  
Forward 5� CAC ATT AAC CCG GCG GTG AC-3�
Reverse  5� CGC AGA GAA GAC GGT GTA GAC -3�
25 �l 2E PCR buffer, 0,5 �M each primer, 1 �g cDNA were poured in each test tube and  a volume of the reactionary mixture was brought to 50 <:; with sterile water.  25 �l mineral oil  for PCR was applied over.  Amplification was performed under the following conditions: (950!   30 s, 570!   30 s, 720C   1 min) 25 cycles, 720!   10 min, 100C   preservation.
The PCR products were separated on a 2% agarous gel, visualized in ultraviolet light and photographed using the system of gel visualization Bio-Vision. Software Image Mater Total Lab"! was used to estimate the quantity of products. The obtained data were reliable under pd"0,05  Student s test. 
Selection of degenerate primers 
    Degenerate primers were firstly constructed to study the expression of PIP aquaporin genes in Sium species on the basis of known mRNA sequences of aquaporins in 10 species of dicotyledonous plants from the data in NCBI and IDT�DNA ( HYPERLINK "http://www.ncbi.nih.gov" http://www.ncbi.nih.gov/;  HYPERLINK "http://www.idtdna.com/analyzer" http://www.idtdna.com/analyzer). An analysis of mRNA sequences was carried by means of the programs BLAST and ClustalW ( HYPERLINK "http://www.ebi.ac.uk/clustalw/)." http://www.ebi.ac.uk/clustalw/).  
   Testing the homology of degenerate primers 
        The obtained cDNA was incorporated in a plasmid pJET 1.2/blunt cloning vector (FERMENTAS) and cloned by means of  the E. coli strain DH10b. After cloning,  kDNA was purified by means of the  set FlexiPrep (GE Helthcare). Nucleotide sequence  was  determined by using the Sanger�s method [19] with an automatic sequenator IBI Prism 3130 Genetic Analyser (Applied Biosystems, USA).                                                                                                                                                                                                                                                                     
The comparison of the kDNA encoding sequence with known sequences of aquaporins in database NCBI revealed its similarity with mRNA of PIP aquaporins of other plants:  
Glycyrrhiza uralensis PIP � similarity  80%. 
Schizolobium parahyba PIP2 � similarity 77%
Juglans regia PIP1;2� similarity 77%
Tulipa gesneriana TgPIP1;2 � similarity 78%
T. gesneriana TgPIP1;1 � similarity 77%
Thus, it has been proved that used primers were complementary to the most conservative patches of PIP aquaporin mRNA of dicotyledonous plants. 
Protein assays
    For Western-blot-analysis of heat shock proteins (HSPs), frozen leaf samples were homogenized with extraction buffer (200��l / 100�mg) containing 25 mM Tris-HCl pH 8.0, 1�mM EDTA, 20�mM NaCl, 1�mM phenylmethylsulfonyl fluoride with a chilled porcelain mortar. The homogenate was centrifuged at 14000 g for 5 min at 4�C. A part of the extract was used to determine protein concentration according to [20]. Another part was mixed with a buffer 0,125� "ris-!l (pH 6,8), 4�% sodium dodecylsulfate, 20�% glycerine and 5�% �-mercaptoethanol in the ratio 1:1. Before electrophoresis protein samples were heated 2 min under 90�! and aliquots contained 30��g of protein were applied on gel. SDS-polyacrylamide gel electrophoresis (SDS-PAGE) was performed in 10�% gel according to [21] using Hoeffer SE 400 (Amersham). Then the gels were electroblotted to nitrocellulose membrane (Amersham) at 0.3 A for 1 h. Immunodetection were carried using monoclonal mouse antibodies against HSP70 (H5147, Sigma) and HSP90 (H1775, Sigma), biotin-conjugated antimouse IgG (Sigma) as described earlier [22]. Protein bands were visualized with the ExtAvidin�peroxidase system using the SIGMA FASTTM DAB system (Sigma). Protein quantity and transfer was contro lled by membrane staining with Ponseau S.
Determination of phytohormones 
Fresh leaf samples (10 g) were placed in 80�% methanol solution immediately after  collection and then homogenized during 3 min using electrical homogenizer (Mechanika Precyzyjna, Poland) in. Cytokinins and abscisic acid (ABA) were extracted from homogenized samples  (Mechanika Precyzyjna, Poland) with 80�% methanol (10 ml/g)  and purified as follows: centrifugation �! fractionation with n-butanol (1:1 v/v) or diethyl ether respectively�! ion-exchange chromatography on Dowex 50Wx8 for cytokinins (Serva, Germany) �! thin layer chromatography on Silicagel 60 F254 (Merk, Germany). Detection and quantification of phytohormones were performed using HPLC-MS system (Agilent 1200, USA). Solid samples were dissolved in 200 �l of mobile phase and 5�l aliquot was injected into Agilent Zorbax Eclipse XDB-C18 column (4,6x250 mm, 5 �m). The column was eluted with an isocratic solvents system methanol:water:acetic acid (37:62.9:0.1 v/v/v) at a flow rate of 0.5 ml/min and column temperature +30�C. The fractions eluted were directly passed through mass spectrometer (Agilent 6120 Quadrupole LC/MS) in a combined regime �multi mode� (electrospray and chemical ionization at atmosphere pressure) of positive ionization. Data were analyzed and processed using the software Agilent ChemStation, version .03.01 on line. Concentrations were calculated on the basis of the peak areas for the endogenous compounds relative to those determined for the internal standards. The data were processed according to the standard methods of variation statistics using Microsoft Excel 2007 program. Values of P < 0,05 were considered to be significant.
Micromorphological study 
     Anthers and mericarps were isolated from peripheral flowers of simple umbels in a compound umbel, as a size of these flowers was bigger and more aligned.
Confocal microscopy
To study the callose deposits, anthers were fixed in Carnoy's solution and embedded in paraffin after dehydration. Sections of 12�14 �m in thickness were made with a sledge microtome (Reichert, Germany), treated with 0.05�% aniline in 0,06� phosphate buffer, @ 8.4, and embedded in  65.0�% sucrose [23]. The localization and content of callose were determined with a confocal microscope LSM-5 Pascal (Carl Zeiss, Germany) under excitation wave-length 405 nm and emission wave-length 530 nm with filters LP530 and BP420-490. Statistical treatment of the obtained data was carried out by means of S""IS"! 5.0. 
Light microscopy   
Mericarps at the successive phases of development were fixed by the Navashin s solution (1% chromic acid :16% formalin : glacial acetic acid, 10 : 4 : 1) Fixed material was dehydrated in graded concentrations of ethanol solutions and in toluol  and embedded in paraf�n with the wax addition. Permanent preparations of 12-14 � m in thickness were made with a sledge microtome (Reichert, Germany), stained with basic fuchsin and light green, embedded in the permount (Permount, Fisher Chemical), analyzed and photographed using microscopes   Stemy SV-6 and Axioscope (Carl Zeiss, Germany). 
 Results
LP and AOA 
Reactive oxygen species production 
ROS production studied by registration of LDCL intensity in leaves of S. latifolium and S, sisaroideum increased from the vegetative phase till fruiting (Fig. 2, i). Since the luminescence intensity is proportionate to the concentration of react components and quantity characterizes the relative content of ROS [24], the enhancement of LDCL intensity evidenced an increase in ROS production.  More remarkable increasing the ROS levels from one phase to another was detected in terrestrial plants. This tendency was maintained in different years, though there were fluctuations in the ROS levels, especially in depending on  weather conditions (Fig. 2, ii).  ROS production in S. sisaroideum increased the most considerably at air temperature of 37�400C.
LP intensity 
LP intensity measured in terms of TBARS content in leaves of both species was higher in S. sisaroideum compared to S. latifolium at the successive phases of plant development (Fig. 2, iii). Similarly to ROS production, the levels of TBARS increased in drought hot weather, especially in terrestrial plants (Fig. 2, iiii).
Using a correlative analysis allowed to demonstrate the positive correlation between ROS production and LP intensity in S. latifolium during the whole vegetation period: a correlative coefficient (r) was 0.97 � 0.03 at the vegetative phase, 0.98 � 0.02 at budding, 0.96,6 � 0.02 at flowering, and 0.97 � 0.03 at fruiting at   d" 0,05. However, in S. sisaroideum, the positive correlation took place only till fruiting: r = 0.98 � 0.01,  0.96 � 0.03 and  = 0.95 � 0.02 respectively  at   d" 0,05.  At the fruiting phase, TBAS accumulation was lower than ROS production. 
Total AOA
tAOA, which reflects a pull of alcohol-dissolve low-molecular-weight antioxidants such as glutathione and ascorbate, increased parallely to higher ROS production and TBRAS accumulation, except for the fruiting phase, where AOA increased fewer than ROS production and LP intensity, especially in S. sisaroideum. A comparative analysis has shown the higher tAOA in terrestrial plants than in aerial-aquatic plants at all phases of plant development (Fig. 3, i). 
SOD activity 
Dynamics of changes in the SOD activity in S. latifolium and S. sisaroideum was analogous to that in tAOA during all phases of plant growth and development.  The SOD activity increased from phase to phase in both species and it was higher in S. sisaroideum, especially under very drought weather (Fig. 3, ii) but less than elevation of ROS production and LP intensity.  
Lipofuscin content 
Lipofuscins, so-called senescence pigments [25], have been detected in leaves of both species at the flowering and fruiting stages, especially in terrestrial plants.  At the fruiting stage, it was < 1.8 times more in S. sisaroideum than in S. latifolium.
Aquaporins
It was shown that accumulation of PIP mRNA in leaves of investigated species was usually more or less similar during the successive stages of ontogenesis but it was significantly lower in S. latifolium aeriel-aquatic plants in comparison wuth S. sisaroideum terrestrial plants, when leaves have been collected at 25�300 C (Fig. 4, a).  Especially clear differences in transcript abundance between terrestrial and aerial�aquatic plants have been observed at air temperature 35�400C (Fig. 4, b). Expression of S. sisaroideum aquaporin PIP genes remained approximately at the same level during ontogenesis, while it was reduced in S. latifolium at the flowering-fruiting phase in comparison with that at the vegetation and budding-flowering phases. In the experiments with growing terrestrial plants at the budding�flowering stage in the overwetting soil as aerial-aquatic ones, the level of PIP mRNA was reduced. Dynamics of gene expression in both investigated species in the daytime was similar: the most level was observed at eight o�clock in the morning, it decreased to noon and it was the lowest at six o'clock in the afternoon.
HSP
     Testing the HSP70 and HSP90 level in leaves of aerial-aquatic S. latifolium and terrestrial S. sisaroideum has been performed at the phase of budding-flowering. Western- blotting revealed the essential variability in the activity of synthesis of both proteins between individuals (Fig. 5, a), while it was higher in terrestrial plants, especially HSP70. The HSP level increased at air temperature of 35-400 C and was more in S. sisaroideum (Fig. 5, b). The HSP90 level in investigated terrestrial and aerial-aquatic plants was more stable between individuals.
Phytohormones
Cytokinins analysis revealed the presence of trans-zeatin (t-Z) in S. latifolium and S. sisaroideum leaves during the vegetative phase and inflorescence formation (Table 1). The largest content of this cytokinin was detected in young plants at the beginning of vegetation. Its level reduced in leaves almost twice during the formation of inflorescences but and it remained quite high in young inflorescences. Zeatin-O-glucoside (ZOG) appeared both in leaves and inflorescences at the flowering�fruiting phase. Seeds contained the high level of free and bound zeatin (Table 1). The obtained data showed that leaves, inflorescences and seeds of aerial-aquatic plants contained 1,5-2 times more trans-zeatin than terrestrial ones, whereas the content of zeatin-O-glucoside in them was similar. 
       The content of abscisic acid (ABA) did not change significantly in plants during development (Table 1). The maximal content of this hormone was detected in inflorescences and seeds. It was about 14-30 % higher in S. sisaroideum compared with S. latifolium. The ratio trans-zeatin/ABA was also higher in S. sisaroideum in comparison with that in S. latifolium during the whole vegetation period, including seeds. 
       Indole-3-acetic acid (IAA) concentrations were higher both in leaves and inflorescences of S. latifolium and S. sisaroideum than other phytohormones (Table 1). The IAA content in S. latifolium leaves increased to the end of vegetation, whereas such tendency was not observed in S. sisaroideum. Significantly increased levels of this phytohormone in S. latifolium compared with those in S. sisaroideum were determined in vegetative and generative organs during plant ontogenesis. 
Micromorpological study
The anthers of S. latifolium aerial-aquatic plants and S. sisaroideum terrestrial plants were more or less similar in size but differed in color � light green in the first species and light violet in the second (Fig. 6, a, b). Mature pollen grains were three-apertured and three-celled, i.e. they consist of a vegetative cell and two male gametes.  Callose was absent in the envelopes of archesporial cells in anthers and it was revealed in envelopes of microspore mother cells (MMCs) at the beginning the transition of archesporial cells to meiosis. In the prophase I, callose was observed in the envelopes of MMCs   by way of the solid but inhomogeneous on thickness layer in S. sisaroideum, while it was observed in the form of separate agglomerations (Fig. 6, c, d). The level of callose fluorescence in MMC envelopes fluctuated in broad limits in S. sisaroideum due to callose uneven deposits, from  25 rel. un. till 230 rel. un., whereas in S. latifolium  � from 25 rel. un. till 53 rel. un.    Callose maximal accumulation in the MMC envelopes was observed in tetrads of microspores and the level of callose fluorescence was higher in S. sisaroideum in comparison with that in S. latifolium:  75,3���2,4 rel. un. and 61,4���2,6 rel. un. respectively. 
There were differences in the rate of embryogenesis between the investigated species, which slowed down in S. latVfolium compared with S. sisaroideum, beginning since 8-10-day old embryos. Consequently,   the embryos in ripe mericarps of S. sisaroideum reached a length of 1.3-1.4 mm, embryos of S. latVfolium   0.7-0.8 mm (Fig. 7).  Embryos consisted of two cotyledons and a root in both species, and a little plumule in S. sisaroideum, and only a meristem protuberance in S. latVfolium. Mericarps in S. sisaroideum matured earlier on 10 15 days than in S. latVfolium 
 Discussion
     The obtained results revealed that levels of ROS formation and LP intensity in leaves of S. latifolium and S. sisaroideum depended on the ontogenesis stage and weather conditions. Increased ROS accumulation and LP intensification have been reported in various plant species under drought conditions depending on stress intensity and duration, plant species and age [26-28]. It is well known that ROS can cause an oxidative damage of proteins, lipids, and DNA [26, 29, 30] and at the same time, they may function also as signals for triggering adaptive/defense responses [31-35]. There are many reports that plant tolerance to different stresses, including water deficit, has been linked to raised activity and/or quantities of antioxidant system components [36-39]. In our work, ROS increasing production and LP intensity in leaves of S. latifolium and S. sisaroideum during plant ontogenesis was accompanied by corresponding enhancement of tAOA and SOD activity that was higher in S. sisaroideum, especially under long-term dry weather.  
      Significantly higher oxidative metabolism in investigated species at the fruiting phase, especially in terrestrial plants, may be explained by the development of plant senescence, as ROS contribute to a process of oxidative deterioration of cellular macromolecules that leads to cell death [40-43]. We hypothesize that LP lower  intensity (measured as TBARS content) in comparison with ROS production in S. sisaroideum at the fruiting phase arised from the formation of  lipofuscins, which generate within plant cells by the interaction of malondialdehyde, that is prominent among TBARS, and hydroxynonenal with amino acids of different macromolecules [25, 44]. There is an evidence that lipofuscins are not detectable at the early stages of plant development and may emerge at the beginning of senescence [16, 45]. Thus, the considerable accumulation of lipofuscins in S. sisaroideum terrestrial plants in comparison with S. latifolium aerial-aquatic onces in flowering � fruiting may indicate an earlier and more intensive senescence of terrestrial plants in conditions of the riverside. Our results are in agreement with an idea of Foyer & Noctor [31] that increased ROS production, instead of as negative term �oxidative stress� in some cases may be described as �oxidative signaling� to the changes in gene expression, metabolism and physiology, that  allows plants to grow and develop in the fluctuated environment.  
         We suppose that the increased level of aquaporin PIP mRNA in leaves of S. sisaroideum in comparison with that in S. latifolium may indicate the aquaporin increased level in the plasmalemma that enchances water inflow in plants and favors the maintainance of water balance   in plants in the riverside conditions. Water transport through the plasmalemma is well known to be ensured by aquaporins, which are membrane proteins forming the water channels [46-49]. Aquaporins of the plasmalemma (PIP) is the most numerous family in plants, that is divided on 2 subfamilies:   1 and   2, the latter are charatterized   by significantly more capability of water tranport in comparison with PIP1 [50]. 
         It has been shown that water deficit impacts the expression of plant PIP genes and a role of aquaporins in plant drought tolerance is widely considered [12, 49]. For example, ZmPIP1;1 in transgenic Arabidopsis thaliana is supposed to enchance plant drought tolerance by inducing stress responsive genes and AOA activity [51]. Expression of PIPs in Solanum tuberosum is assumed to enhance the whole plant tolerance to drought improving overall water relations in plants [52]. We showed the higher activity of tAOA and SOD simultaneously with more accumulation of PIP mRNA in leaves of S. sisaroideum terrestrial plants as compared with  S. latifolium, especially in very dry and hot weather. In addition, PIP expression decreased in leaves of S. sisaroideum   in response to waterlogging, i.e. hypoxia conditions, which are routine for aerial-aquatic plants. It is suggested a possible pivotal role of certain aquaporins in Sorghum bicolor tolerance to waterlogging stress [53]. The available literature data on a main role of aquaporins in water transport  and maintaining water balance in plants allow to consider the regulation of AQP activity and gene expression as a part of the adaptation mechanisms to stress conditions [49].  Our investigations of PIP expression in close related species S. latifolium and S. sisaroideum growing in waterlogging and aerated soil corroborate this idea and demonstrate high lability of PIP expression in response to changes in the soil water content in the biotope.
    Early we established high metabolic and structural plasticity of  root systems in S. latifolium and S. sisaroideum, which is under control of soil moisture [54]. Lowering of the water level (dry and hot weather) induced secondary growth of the cortex of available adventitious roots in S. latifolium. Under soil water-logging (long-term rainy weather), new adventitious roots with alcohol dehydrogenase (ADH)  and aerenchyma initiated rapidly in S. sisaroideum. 
      We showed the presence of HSP70 and HSP90, which are capable to bind with antibodies for conservative areas of these proteins, in S. latifolium and S.�sisaroideum plants from natural populations. Under water-logging in the natural and experimental conditions, HSP70 accumulation in leaves positively correlated with the ADH level, especially such correlation was pronounced in S.�sisaroideum [54]. Synthesis of HSPs is well known to be induced by unfavorable changes of ecological factors and it is one of the main components of plant stress-reaction [55-60]. The enhanced HSP70 synthesis in leaves under hypoxia may be induced by the ROS increased content and  ethanol accumulation.  Simultaneous rapid induction of HSP70 and ADH in response to anoxia has been shown in A.�thaliana seedlings [61].  HSP70  is considered to bind non-native conformations of proteins and occupy the central position in work of the cell chaperon system [62]. A less variability in the HSP90 content in leaves of S. latifolium and S. sisaroideum may be explained by the HSP90 significance in cell metabolism in the normal conditions. Its expression in response to unfavorable effects changes considerably less than expression of other HSPs, e.g. HSP70 [63]. Our data confirmed that adaptation of investigated species to changes in their habitats  is connected with the induction of HSP70 synthesis under the maintenance of the sufficiently regular HSP90 level. 
     Cytokinins increase a rate of shoot growth, induce the opening of stomachs, i.e. their actions are the opposite of the effects of water deficiency. The assumption that cytokinins are involved in the drought adaptation is evidenced not only by changes in their qualitative composition and quantitative content, but also by the fact that transgenic plants with modified signaling of cytokinins change water regime and stress tolerance [64]. Reducing cytokinins level caused the decrease in growth rate, reduction in leaf surface and as a sequence the decline in the transpiration [65]. Consequently, the reduced level of endogenous cytokinins in S. sisaroideum can help to overcome the negative effects of limited moisture, regulating the water status of plants. It should be noted that distribution of cytokinins between vegetative and generative organs of both species was similar, that is possibly allows to maintain them fertility at the equal level.
     ABA is a crucial hormone controlling plants responses to different stresses including water deficiency [66]. Plants regulate the stomatal closure and control the transpirational water loss under drought by changes in ABA synthesis. The ABA increased content is common for plants with a greater ability to adapt to water lack [67]. Accumulation of the ABA higher levels in terrestrial S. sisaroideum compared with aerial-aquatic S. latifolium during ontogenesis is assumed to prevent the tissue dehydration. The balance of hormones is known to be maintained by the ability of each of them to affect the content of another, changing its synthesis, transport or metabolism [68]. Changes in the concentrations of antagonistic hormones in the opposite direction may cause an increase in the same effect, e.g. stomata closure under heat stress [69]. Probably, the ratio of cytokinin/ABA in plant organs reflects the type of their way to respond to stress [70]. Unlike aerial-aquatic S. latifolium, the ABA prevalence over cytokinins in terrestrial S. sisaroideum may be attributed to the adaptive mechanisms to the tiverside conditions. 
A role of IAA in adaptation to drought is contradictory. IAA has been reported to induce leaf epinasty and to stimulate the root formation as a water-stress defense mechanism [71]. IAA is suggested to modulate the crosstalk between endogenous hormones and polyamines, improving the antioxidant defense under water stress [72]. The IAA dominance at all developmental phases in S. latifolium as compared with S. sisaroideum indicates the participation of this phytohormone in regulation of these plants tolerance to water-limited conditions.
       The callose  increased content in the envelopes of both MMCs during microsporogenesis and pollen grains in S. sisaroideum may be considered to favour the moisture retention in envelopes and maintenance of  pollen vital activity during its transfer and germination on the stigma, a water potential of which is lower than in S. latifolium. Callose   � -1,3-glucan, plant polysaccharide is the necessary carbohydrate component of the envelopes of MMCs, that protects them from the water loss under lowering the water content in soil [73, 74]. In addition, flowers of terrestrial plants have the styles 2 times shorter than those in aerial-aquatic plants and larger stigmas that facilitates more pollen hit on stigmas and growth of pollen tubes by a shorter way to approach ovules and, thus, ensures pollination and fertilization of terrestrial plants. The presence of the callose increased content in the envelopes of microspore tetrads and pollen grains in the S. sisaroideum terrestrial plants corresponds to generally accepted view on a callose positive role in the male gametophyte development, especially in the soil water deficiency [75].  
       Earlier maturation of mericarps in terrestrial plants confirms the reported data on the acceleration of plant reproductive development under the soil limited content.  Plant resistance to changes in the soil moisture is known to be conditioned by the phytohormonal complex, especially ABA [76, 77]. It is of interest the content of this hormone in inflorescences and seeds of S. sisaroideum was about 14-30 % higher compared with S. latifolium. The ABA higher level under water deficiency induces dehydrine expression [78, 79], that leads to  more rapid water loss by seeds, i.e.  seed maturation is  to a certain extent controlled by ABA [80]. 
Stylopodia in S. sisaroideum differ from those in S. latVfolium by more flexuous edges which  cover the upper part of an ovary yet at the early stages of style formation. This stylopodia s in terrestrial plants along with their longer existence after fertilization is assumed to maintain the moisture level in growing mericarps. Stylopodia of aerial-aquatic plants wither greatly erlier. The presence of a completely differentiated embryo in mature mericarps ensures their rapid germination during short periods of sufficient soil moisture in spring.
         Thus, our study of certain micromorphological, physiological and molecular traits  of closely related but ecologically different Sium species " aerial-aquatic S. latifolium and terrestrial S. sisaroideum showed their plasticity in plant adaptation to different soil moisture, that helps to explain widespread distribution of these species in nature. The increased levels of free-radical oxidation during ontogenesis is supposed to play an important role in  S. sisaroideum drought-resistance. Being well-adjusted to the moisture of dry aerated soil, terrestrial plants retain their ability to adaptive respond on flooding. These findings showed an adaptive role of phenotypic plasticity in plant successful growth, development and reproduction under soil moisture fluctuations. The investigated species are suitable objects for further research  of plant ontogenetic plasticity in nature.
Acknowledgments
We are grateful to Valentina V. Baranenko and Antonina F. Popova for their major contribution to implementation of this research. The work was funded by the National Academy of Sciences of Ukraine.
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Legendes to Figures
Fig. 1.  Sium latifolium aerial-aquatic plants (a) and S. sisaroideum terrestrial plants (b)   
Fig. 2. Lipid peroxidation (LP) in leaves of S. latifolium � 1 and S. sisaroideum � 2. Intensity of luminol-dependent chemiluminescence (impulses in minute � 103) at the successive phases of ontogenesis (i) and at the flowering phase at air temperature of 300�250C  and 350�400C  (ii). Thiobarbituric acid-reacting substances (TBARS) content (nmol g"1 fresh weight) at the successive phases of ontogenesis (iii) and at the flowering phase at air temperature of 300 250C and 350 400C (iiii). a   vegetation phase, b   budding, c   flowering, d    fruiting.
Fig. 3. Total antioxidant activity (AOA) (i) and superoxide dismutase (SOD) activity (ii)  (rel. unit g-1 fresh weight) in leaves of S. latifolium � 1 and S. sisaroideum � 2 at the successive phases of ontogenesis:  a � vegetation phase, b � budding, c �  flowering, d �  fruiting. 
Fig. 4. Agarous gel � electrophoresis with the products of RT�PCR, mRNA of PIP aquaporin gene. A � aerial-aquatic plants, T � terrestrial plants. M � DNA-marker from 100 pn till 400 pn. PCR-product near 350 pn. a - t � 25-300C; b - t � 35-400C.
Fig. 5. Immunoblots of HSP70 and HSP90 in leaves of S. latifolium aerial-aquatic plants growing in coastal water and S. sisaroideum terrestrial plants growing on the riverside at air temperature of  250�300C (a) and 400 C (b).
Fig. 6. Flowers of S. sisaroideum (a) and S. latifolium (b), tetrads of microspores in anthers of S. sisaroideum (c) and S. latifolium (d).  A white line indicates the direction of scanning vector of the callose fluorescence �������	�����(�)�,�-�7�8�A�B�D�E�H�I�R�S�U�V�Z�[�\�^�e�g�o�q�s�t�y�z���������)�*�ܢݢ����2�f�g���������`�a�����s�t�G�H���E�F�ȩɩj�k�o�����������������������������������������hS�hS�mH	sH	h�
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