A comprehensive investigation was carried out to determine the changes that occurred in water-stressed cucumber (Cucumis sativus L.) in response to melatonin treatment. We examined the potential roles of melatonin during seed germination and root generation and measured its effect on reactive oxygen species (ROS) levels, antioxidant enzyme activities, and photosynthesis. Melatonin alleviated polyethylene glycol induced inhibition of seed germination, with 100 μm melatonin-treated seeds showing the greatest germination rate. Melatonin stimulated root generation and vitality and increased the root:shoot ratio; therefore, melatonin may have an effect on strengthening cucumber roots. Melatonin treatment significantly reduced chlorophyll degradation. Seedlings treated with 100 μm melatonin clearly showed a higher photosynthetic rate, thus reversing the effect of water stress. Furthermore, the ultrastructure of chloroplasts in water-stressed cucumber leaves was maintained after melatonin treatment. The antioxidant levels and activities of the ROS scavenging enzymes, i.e., superoxide dismutase, peroxidase, and catalase, were also increased by melatonin. These results suggest that the adverse effects of water stress can be minimized by the application of melatonin.

Kochia sieversiana (Pall.) C. A. M., a naturally alkali-resistant halophyte, was chosen as the test organism for our research. The seedlings of K. sieversiana were treated with varying (0-400 mM) salt stress (1:1 molar ratio of NaCl to Na 2 SO 4 ) and alkali stress (1:1 molar ratio of NaHCO 3 to Na 2 CO 3 ). The concentrations of various solutes in fresh shoots, including Na + , K + , Ca 2+ , Mg 2+ , Cl À , SO 4 2À , NO 3 À , H 2 PO 3 À , betaine, proline, soluble sugar (SS), and organic acid (OA), were determined. The water content (WC) of the shoots was calculated and the OA components were analyzed. Finally, the osmotic adjustment and ion balance traits in the shoots of K. sieversiana were explored. The results showed that the WC of K. sieversiana remained higher than 6 [g g À1 Dry weight (DW)] even under the highest salt or alkali stress. At salinity levels >240 mM, proline concentrations increased dramatically, with rising salinity. We proposed that this was not a simple response to osmotic stress. The concentrations of Na + and K + all increased with increasing salinity, which implies that there was no competitive inhibition for absorption of either in K. sieversiana. Based on our results, the osmotic adjustment feature of salt stress was similar to that of alkali stress in the shoots of K. sieversiana. The shared essential features were that the shoots maintained a state of high WC, OA, Na + , K + and other inorganic ions, accumulated largely in the vacuoles, and betaine, accumulated in cytoplasm.On the other hand, the ionic balance mechanisms under both stresses were different. Under salt stress, K. sieversiana accumulated OA and inorganic ions to maintain the intracellular ionic equilibrium, with close to equal contributions of OA and inorganic ions to anion. However, under alkali stress, OA was the dominant factor in maintaining ionic equilibrium. The contribution of OA to anion was as high as 84.2%, and the contribution of inorganic anions to anion was only 15.8%. We found that the physiological responses of K. sieversiana to salt and alkali stresses were unique, and that mechanisms existed in it that were different from other naturally alkaliresistant gramineous plants, such as Aneurolepidium chinense, Puccinellia tenuiflora.

Effects of salt and alkali stresses on growth, osmotic adjustment and ionic balance of Suaeda glauca (Bge.), an alkali-resistant succulent halophyte, were compared. The results showed that alkali stress clearly inhibited the growth of S. glauca. Moreover, the concentrations of Na + and K + both increased with increasing salinity under both stresses, suggesting no competitive inhibition between absorptions of Na + and K + . The mechanism underlying osmotic adjustment during salt stress was similar to alkali stress in shoots. The shared essential features were that organic acids, betaine and inorganic ions (dominated by Na + ) mostly accumulated. On the other hand, the mechanisms governing ionic balance under both stresses were different. Under salt stress, S. glauca accumulated organic acids and inorganic anions to maintain the intracellular ionic equilibrium, but the anion contribution of inorganic ions was greater than that of organic acids. However, the concentrations of inorganic anions under alkali stress were significantly lower than those under salt stress of the same intensity, suggesting that alkali stress might inhibit uptake of anions, such as NO 3 -and H 2 PO 4 -. Under alkali stress, organic acids were the dominant factor in maintaining ionic equilibrium. The contribution of organic acids to anions was 74.1%, while that of inorganic anions was only 25.9%. S. glauca enhanced the synthesis of organic acids, dominated by oxalic acid, to compensate for the shortage of inorganic anions.

We compared the effects of salt-stresses (SS, 1 : 1 molar ratio of NaCl to Na 2 SO 4 ) and alkali-stresses (AS, 1 : 1 molar ratio of NaHCO 3 to Na 2 CO 3 ) on the growth, photosynthesis, solute accumulation, and ion balance of barley seedlings, to elucidate the mechanism of AS (high-pH) damage to plants and the physiological adaptive mechanism of plants to AS. The effects of SS on the water content, root system activity, membrane permeability, and the content of photosynthetic pigments were much less than those of AS. However, AS damaged root function, photosynthetic pigments, and the membrane system, led to the severe reductions in water content, root system activity, content of photosynthetic pigments, and net photosynthetic rate, and a sharp increase in electrolyte leakage rate. Moreover, with salinity higher than 60 mM, Na + content increased slowly under SS and sharply under AS. This indicates that high-pH caused by AS might interfere with control of Na + uptake in roots and increase intracellular Na + to a toxic level, which may be the main cause of some damage emerging under higher AS. Under SS, barley accumulated organic acids, Cl − , SO 4 2− , and NO 3 − to balance the massive influx of cations, the contribution of inorganic ions to ion balance was greater than that of organic acids. However, AS might inhibit absorptions of NO 3 -and Cl -, enhance organic acid synthesis, and SO 4 2− absorption to maintain intracellular ion balance and stable pH.

Hexaploid bread wheat (Triticum aestivum L., genome BBAADD) is generally more salt tolerant than its tetraploid wheat progenitor (Triticum turgidum L.). However, little is known about the physiological basis of this trait or about the relative contributions of allohexaploidization and subsequent evolutionary genetic changes on the trait development. Here, we compared the salt tolerance of a synthetic allohexaploid wheat (neo-6x) with its tetraploid (T. turgidum; BBAA) and diploid (Aegilops tauschii; DD) parents, as well as a natural hexaploid bread wheat (nat-6x). We studied 92 morphophysiological traits and analyzed homeologous gene expression of a major salt-tolerance gene High-Affinity K + Transporter 1;5 (HKT1;5). We observed that under salt stress, neo-6x exhibited higher fitness than both of its parental genotypes due to inheritance of favorable traits like higher germination rate from the 4x parent and the stronger root Na + retention capacity from the 2x parent. Moreover, expression of the D-subgenome HKT1;5 homeolog, which is responsible for Na + removal from the xylem vessels, showed an immediate transcriptional reprogramming following allohexaploidization, i.e., from constitutive high basal expression in Ae. tauschii (2x) to salt-induced expression in neo-6x. This phenomenon was also witnessed in the nat-6x. An integrated analysis of 92 traits showed that, under salt-stress conditions, neo-6x resembled more closely the 2x than the 4x parent, suggesting that the salt stress induces enhanced expressivity of the D-subgenome homeologs in the synthetic hexaploid wheat. Collectively, the results suggest that condition-dependent functionalization of the subgenomes might have contributed to the wide-ranging adaptability of natural hexaploid wheat.transcriptional rewiring | Na + homeostasis | salinity tolerance P olyploidy or whole genome duplication (WGD) is a pervasive, driving force in plant and vertebrate evolution, which has fascinated biologists for more than a century (1, 2). The common occurrence of WGD suggests an evolutionary advantage of having multiple genomes at least in certain circumstances, which might have enabled the polyploid organisms to be better adapted to some adverse environmental conditions than their diploid progenitors (3, 4). Polyploidy can instantaneously develop novel features that allow them to invade new territories or expand their parental niche (3). Polyploids may also exhibit higher evolvability than their diploid progenitors, which allows them to adapt to capricious environmental conditions (4). Thus, polyploidy has been demonstrated as a process that may lead to saltational speciation especially when novel ecological niches are available for colonization.Although abrupt genome duplication often produces adverse effects on physiology at both cellular and organismal levels (4, 5), it has been shown that polyploidy in plants may result in favorable physiological consequences such as increased photosynthetic capacity and enhanced tolerance to biotic and abiotic stresses, which ...

The seedlings of wheat were treated by salt-stress (SS, molar ratio of NaCl : Na 2 SO 4 = 1 : 1) and alkali-stress (AS, molar ratio of NaHCO 3 : Na 2 CO 3 = 1 : 1). Relative growth rate (RGR), leaf area, and water content decreased with increasing salinity, and the extents of the reduction under AS were greater than those under SS. The contents of photosynthetic pigments did not decrease under SS, but increased at low salinity. On the contrary, the contents of photosynthetic pigments decreased sharply under AS with increasing salinity. Under SS, the changes of net photosynthetic rate (P N ), stomatal conductance (g s ), and transpiration rate (E) were similar and all varied in a single-peak curve with increasing salinity, and they were lower than those of control only at salinity over 150 mM. Under AS, P N , g s , and E decreased sharply with rising salinity. The decrease of g s might cause the obvious decreases of E and intercellular CO 2 concentration, and the increase of water use efficiency under both stresses. The Na + content and Na + /K + ratio in shoot increased and the K + content in shoot decreased under both stresses, and the changing extents under AS were greater than those under SS. Thus SS and AS are two distinctive stresses with different characters; the destructive effects of AS on the growth and photosynthesis of wheat are more severe than those under SS. High pH is the key feature of the AS that is different from SS. The buffer capacity is essentially the measure of high pH action on plant. The deposition of mineral elements and the intracellular unbalance of Na + and K + caused by the high pH at AS might be the reason of the decrease of P N and g s and of the destruction of photosynthetic pigments.

BackgroundIt is well known that salt stress has different effects on old and young tissues. However, it remains largely unexplored whether old and young tissues have different regulatory mechanism during adaptation of plants to salt stress. The aim of this study was to investigate whether salt stress has different effects on the ion balance and nitrogen metabolism in the old and young leaves of rice, and to compare functions of both organs in rice salt tolerance.ResultsRice protected young leaves from ion harm via the large accumulation of Na+ and Cl− in old leaves. The up-regulation of OsHKT1;1, OsHAK10 and OsHAK16 might contribute to accumulation of Na+ in old leaves under salt stress. In addition, lower expression of OsHKT1;5 and OsSOS1 in old leaves may decrease frequency of retrieving Na+ from old leaf cells. Under salt stress, old leaves showed higher concentration of NO3− content than young leaves. Up-regulation of OsNRT1;2, a gene coding nitrate transporter, might contribute to the accumulation of NO3− in the old leaves of salt stressed-rice. Salt stress clearly up-regulated the expression of OsGDH2 and OsGDH3 in old leaves, while strongly down-regulated expression of OsGS2 and OsFd-GOGAT in old leaves.ConclusionsThe down-regulation of OsGS2 and OsFd-GOGAT in old leaves might be a harmful response to excesses of Na+ and Cl−. Under salt stress, rice might accumulate Na+ and Cl− to toxic levels in old leaves. This might influence photorespiration process, reduce NH4+ production from photorespiration, and immediately down-regulate the expression of OsGS2 and OsFd-GOGAT in old leaves of salt stressed rice. Excesses of Na+ and Cl− also might change the pathway of NH4+ assimilation in old leaves of salt stressed rice plants, weaken GOGAT/GS pathway and elevate GDH pathway.

Subgenome integrity in bread wheat (Triticum aestivum; BBAADD) makes possible the extraction of its BBAA component to restitute a novel plant type. The availability of such a ploidy-reversed wheat (extracted tetraploid wheat [ETW]) provides a unique opportunity to address whether and to what extent the BBAA component of bread wheat has been modified in phenotype, karyotype, and gene expression during its evolutionary history at the allohexaploid level. We report here that ETW was anomalous in multiple phenotypic traits but maintained a stable karyotype. Microarray-based transcriptome profiling identified a large number of differentially expressed genes between ETW and natural tetraploid wheat (Triticum turgidum), and the ETW-downregulated genes were enriched for distinct Gene Ontology categories. Quantitative RT-PCR analysis showed that gene expression differences between ETW and a set of diverse durum wheat (T. turgidum subsp durum) cultivars were distinct from those characterizing tetraploid cultivars per se. Pyrosequencing revealed that the expression alterations may occur to either only one or both of the B and A homoeolog transcripts in ETW. A majority of the genes showed additive expression in a resynthesized allohexaploid wheat. Analysis of a synthetic allohexaploid wheat and diverse bread wheat cultivars revealed the rapid occurrence of expression changes to the BBAA subgenomes subsequent to allohexaploidization and their evolutionary persistence.