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Osmolyte accumulation: can it really help increase crop yield under drought conditions?

Osmolyte accumulation (OA) is frequently cited as a key putative mechanism for increasing yields of crops subjected to drought conditions. The hypothesis is that OA results in a number of benefits that sustain cell and tissue activity under water-deficit conditions. It has been proposed as an effective tolerance mechanism for water deficits, which could be enhanced in crops by traditional plant breeding, marker-assisted selection or genetic engineering, to generate drought-tolerant crops. However, field studies examining the association between OA and crop yield have tended to show no consistent benefit. The few, often-cited, investigations with positive associations were obtained under severe water deficits with extremely low yields or conditions with special water-supply scenarios when much of the benefit is plant survival. Under conditions where water deficits threaten crop survival, yields are so low that even large fractional yield gains offer little practical benefit to growers. Indeed, the often-cited benefit of turgor maintenance in cells is likely to result in crop behaviour that is exactly opposite to what is beneficial to crops. The one clear mechanism identified in this review for beneficial yield responses to OA is in the maintenance of root development in order to reach water that may be available deeper in the soil profile.

Journal Article↗

Calcium signalling in Arabidopsis thaliana responding to drought and salinity.

Changes in cytosolic free calcium concentration ([Ca2+]cyt) in response to mannitol (drought) and salt treatments were detected in vivo in intact whole Arabidopsis seedlings. Transient elevations of [Ca2+]cyt to around 1.5 microM were observed, and these were substantially inhibited by pretreatment with the calcium-channel blocker lanthanum and to a lesser extent, the calcium-chelator EGTA. The expression of three genes, p5cs, which encodes delta(1)-pyrroline-5-carboxylate synthetase (P5CS), the first enzyme of the proline biosynthesis pathway, rab18 and Iti78 which both encode proteins of unknown function, was induced by mannitol and salt treatments. The induction of all three genes by mannitol was inhibited by pretreatment with lanthanum. Salt-induced p5cs, but not rab18 and Iti78, expression was also inhibited by lanthanum. Induction of p5cs by mannitol was also inhibited by the calcium channel-blockers gadolinium and verapamil and the calcium chelator EGTA, further suggesting the involvement of calcium signalling in this response. Mannitol induced greater levels of p5cs gene expression than an isoosmolar concentration of salt, at both relatively high and low concentrations. However, calcium transients were of a similar magnitude and duration in response to both mannitol and isoosmolar concentrations of salt, suggesting that a factor other than calcium is involved in the discrimination between drought and salinity signals in Arabidopsis. In order to gauge the involvement of the vacuole as an intracellular calcium store in the response of Arabidopsis to mannitol, [Ca2+]cyt was measured at the microdomain adjacent to the vacuolar membrane. The results obtained were consistent with a significant calcium release from the vacuole contributing to the overall mannitol-induced [Ca2+]cyt response. Data obtained by using inhibitors of inositol signalling suggested that this release was occurring through IP3-dependent calcium channels.

1-Pyrroline-5-Carboxylate Dehydrogenase↗

OsDREB genes in rice, Oryza sativa L., encode transcription activators that function in drought-, high-salt- and cold-responsive gene expression.

The transcription factors DREBs/CBFs specifically interact with the dehydration-responsive element/C-repeat (DRE/CRT) cis-acting element (core motif: G/ACCGAC) and control the expression of many stress-inducible genes in Arabidopsis. In rice, we isolated five cDNAs for DREB homologs: OsDREB1A, OsDREB1B, OsDREB1C, OsDREB1D, and OsDREB2A. Expression of OsDREB1A and OsDREB1B was induced by cold, whereas expression of OsDREB2A was induced by dehydration and high-salt stresses. The OsDREB1A and OsDREB2A proteins specifically bound to DRE and activated the transcription of the GUS reporter gene driven by DRE in rice protoplasts. Over-expression of OsDREB1A in transgenic Arabidopsis induced over-expression of target stress-inducible genes of Arabidopsis DREB1A resulting in plants with higher tolerance to drought, high-salt, and freezing stresses. This indicated that OsDREB1A has functional similarity to DREB1A. However, in microarray and RNA blot analyses, some stress-inducible target genes of the DREB1A proteins that have only ACCGAC as DRE were not over-expressed in the OsDREB1A transgenic Arabidopsis. The OsDREB1A protein bound to GCCGAC more preferentially than to ACCGAC whereas the DREB1A proteins bound to both GCCGAC and ACCGAC efficiently. The structures of DREB1-type ERF/AP2 domains in monocots are closely related to each other as compared with that in the dicots. OsDREB1A is potentially useful for producing transgenic monocots that are tolerant to drought, high-salt, and/or cold stresses.

Adaptation, Physiological↗

Subcellular effects of drought stress in Rosmarinus officinalis.

The use of Rosmarinus officinalis, and other wild plant species, in the Mediterranean area is an interesting solution in order to avoid the desertification and rapid soil erosion, because of their good resistance to environmental conditions. Previous articles have described experiments designed to determine the impact of water stress at the plant level in this species, but more knowledge is required at the subcellular and ultrastructural levels. An anatomic and ultrastructural study of the leaves was conducted on Rosmarinus officinalis plants growing under different water treatments. In the leaves of water-stressed plants, the leaf water potential and turgor decreased, and leaf osmotic potential became more negative with respect to control plants. The anatomic investigations showed that both the mesophyll intercellular spaces and the epidermal cell size were reduced significantly under the more intense drought stress conditions. At the subcellular level, chloroplasts accumulated plastoglobuli and lipid bodies, and cuticle thickness was increased under water stress. In our experiment, the anatomic and ultrastructural modifications of Rosmarinus officinalis could be considered an additional adaptation to drought stress together with physiological and biochemical modifications as antioxidant accumulation.

Acclimatization↗

Fire cycles in North American interior grasslands and their relation to prairie drought.

High-resolution analyses of a late Holocene core from Kettle Lake in North Dakota reveal coeval fluctuations in loss-on-ignition carbonate content, percentage of grass pollen, and charcoal flux. These oscillations are indicative of climate-fuel-fire cycles that have prevailed on the Northern Great Plains (NGP) for most of the late Holocene. High charcoal flux occurred during past moist intervals when grass cover was extensive and fuel loads were high, whereas reduced charcoal flux characterized the intervening droughts when grass cover, and hence fuel loads, decreased, illustrating that fire is not a universal feature of the NGP through time but oscillates with climate. Spectral and wavelet analyses reveal that the cycles have a periodicity of approximately = 160 yr, although secular trends in the cycles are difficult to identify for the entire Holocene because the periodicity in the early Holocene ranged between 80 and 160 yr. Although the cycles are evident for most of the last 4,500 yr, their occasional muting adds further to the overall climatic complexity of the plains. These findings clearly show that the continental interior of North America has experienced short-term climatic cycles accompanied by a marked landscape response for several millennia, regularly alternating between dual landscape modes. The documentation of cycles of similar duration at other sites in the NGP, western North America, and Greenland suggests some degree of regional coherence to climatic forcing. Accordingly, the effects of global warming from increasing greenhouse gases will be superimposed on this natural variability of drought.

Carbonates↗

Up-regulation of a H+-pyrophosphatase (H+-PPase) as a strategy to engineer drought-resistant crop plants.

Engineering drought -resistant crop plants is a critically important objective. Overexpression of the vacuolar H(+)-pyrophosphatase (H(+)-PPase) AVP1 in the model plant Arabidopsis thaliana results in enhanced performance under soil water deficits. Recent work demonstrates that AVP1 plays an important role in root development through the facilitation of auxin fluxes. With the objective of improving crop performance, we expressed AVP1 in a commercial cultivar of tomato. This approach resulted in (i) greater pyrophosphate-driven cation transport into root vacuolar fractions, (ii) increased root biomass, and (iii) enhanced recovery of plants from an episode of soil water deficit stress. More robust root systems allowed transgenic tomato plants to take up greater amounts of water during the imposed water deficit stress, resulting in a more favorable plant water status and less injury. This study documents a general strategy for improving drought resistance of crops.

Arabidopsis↗

Messenger RNA is conserved during drying of the drought-tolerant moss Tortula ruralis.

On rehydration after complete drying, the drought-tolerant moss Tortula ruralis (Hedw.) Gaertn., Meyer, and Scherb reforms its polyribosomes and can resume protein synthesis without new RNA synthesis. Results obtained with the double-label ratio technique coupled with polyacrylamide disc gel electrophoresis show that (i) the proteins synthesized on rehydration are largely similar to those synthesized at the time of dehydration and (ii) inhibition of RNA synthesis does not alter the rate or the pattern of protein synthesis during rehydration of slowly dried moss. Poly(A)-rich RNA has been isolated from fresh and dried moss by chromatography on oligo(dT)-cellulose and has been translated in vitro in the cell-free wheat germ system. It is concluded that mRNA is conserved during complete drying of T. ruralis and supports protein synthesis on subsequent rehydration. This ability to conserve mRNA is a characteristic held in common by vegetative drought-tolerant tissue of T. ruralis and air-dried seeds of higher plants.

Journal Article↗

Up-regulation of glutathione metabolism and changes in redox status involved in adaptation of reed (Phragmites communis) ecotypes to drought-prone and saline habitats.

The glutathione (GSH) metabolic characteristics and redox balance in three ecotypes of reed (Phragmites communis), swamp reed (SR), dune reed (DR), and heavy salt meadow reed (HSMR), from different habitats in desert regions of northwest China were investigated. The DR possessed the highest rate of GSH biosynthesis and metabolism with the lowest levels of total and reduced GSH and its biosynthetic precursors, gamma-glutamylcysteine (gamma-EC) and cysteine (Cys), of the three reed ecotypes. This suggests that a higher rate of GSH biosynthesis and metabolism, but not GSH accumulation, might be involved in the adaptation of this terrestrial reed ecotype to its dry habitat. The HSMR shared this profile although it exhibited the highest reduced thiol levels of the three ecotypes. Two key enzymes in the Calvin-cycle possessing exposed sulfhydryl groups, NADP(+)-dependent glyceraldehydes-3-phosphate dehydrogenase (G3PD) and fructose-1,6-bisphosphatase (FBPase), and other two key enzymes in the pentose-phosphate pathway (PPP), glucose-6-phosphate dehydrogenase (G6PDH) and 6-phosphogluconate dehydrogenase (6-PGD), had very similar activities in the three reed ecotypes. Compared to the SR, the DR and HSMR had higher ratios of NADPH/NADP+ and NADH/NAD+, indicating that a more reduced redox status in the plant cells might be involved in the survival and adaptation of the two terrestrial reed ecotypes to long-term drought and salinity, respectively. These results suggest that changes of GSH metabolism and redox balance were important components of the adaptation of reed, a hydrophilic plant, to more extreme dune and saline habitats. The coordinated up-regulations of the rate of GSH biosynthesis and metabolism and reduction state of redox status of plant cells, conferred on the plant high resistance or tolerance to long-term drought and salinity.

Adaptation, Physiological↗

Genotypic variation in the photosynthetic competence of Sorghum bicolor seedlings subjected to polyethylene glycol-mediated drought stress.

Eleven varieties of Sorghum bicolor, subjected to PEG-mediated drought stress were compared for their photosynthetic performance. The varieties differed in their relative water content over a range of PEG concentrations (0-25%). CO2 assimilation, stomatal conductance and the quantum yield of PSII electron transport decreased with increasing PEG concentrations in all varieties. However the intercellular CO2 concentration showed a nonlinear PEG concentration-dependent change. At lower PEG concentrations there was a decrease in the levels of intercellular CO2 concentration in all varieties that could be attributed to stomatal closure. At higher PEG concentrations, some varieties showed an increase in the intercellular CO2 concentration, indicating an inhibition of photosynthetic activity due to non-stomatal effects, while others did not. It was seen that the varieties differed in the stress thresholds at which stomatal and metabolic limitations to photosynthesis occur. These differences in the photosynthetic adaptation of Sorghum varieties could be useful in identifying genotypes showing large differences in photosynthetic adaptation, which could be useful in mapping photosynthetic traits for drought stress tolerance.

Adaptation, Physiological↗

Dietary pattern and state of nutrition among children in drought-prone areas of southern Ethiopia.

To assess dietary habits and nutritional state in drought-prone areas of southern Ethiopia, we studied 334 households in a pastoral and 282 in an agricultural community. Milk and cereals were the main sources of food among children of the pastoral Boran in Dubluk, while cereals with limited supplements of animal products or legumes formed the main sources of food among children of the agricultural population of Elka in the Rift valley. Of the children in Elka, 54.9% were stunted, as compared with 19.5% among children in Dubluk. Also, stunting occurred at an earlier age among the Elka children. Prevalences of wasting were less than 5% in both communities. Improvement in the state of nutrition of the pastoral children followed soon after the main rains, but occurred later and after the main harvest among the agricultural children. In contrast to arm circumference, the weight-for-height measure showed marked seasonal variation. Socio-economic factors, such as family wealth and crowding, significantly influenced the state of nutrition among the children. Nutritional recovery following the prolonged drought among the agricultural children was slow and associated with families acquiring more wealth.

Anthropometry↗

Identification of a rice zinc finger protein whose expression is transiently induced by drought, cold but not by salinity and abscisic acid.

A C2H2-type zinc finger protein gene ZFP245 was cloned by RT-PCR approach from cold treated rice seedlings. ZFP245 is 712 bp in length and encodes a 24.5 KDa protein, which has 35% identity in amino acid with SCOF-1, a cold-inducible zinc finger protein from soybean. By database search, ZFP245 was mapped on chromosome 7 and clustered together with another C2H2 zinc finger gene. Tissue expression analysis showed that ZFP245 was constitutively expressed in various rice tissues including roots, stems, leaves and spikes. The semi-quantitative-RT-PCR assay revealed ZFP245 was strongly induced after 6 h exposure to cold and drought stresses, and then reduced to the baseline. However, ZFP245 was not regulated by high salt or abscisic acid treatment. The promoter sequence of 1017 nucleotides, upstream of ZFP245, was cloned directly by PCR approach. Sequence analysis revealed a CRT/DRE element was found within the ZFP245 promoter region. Taken together, ZFP245, as the first identified C2H2-type zinc finger protein involved in stress response in monocots probably plays a role as a transcription regulator in plant cold and drought responses through an ABA-independent pathway.

Abscisic Acid↗

Drought and oxidative load in the leaves of C3 plants: a predominant role for photorespiration?

Although active oxygen species are produced at high rates in both the chloroplasts and peroxisomes of the leaves of C3 plants, most attention has focused on the potentially damaging consequences of enhanced chloroplastic production in stress conditions such as drought. This article attempts to provide quantitative estimates of the relative contributions of the chloroplast electron transport chain and the glycolate oxidase reaction to the oxidative load placed on the photosynthetic leaf cell. Rates of photorespiratory H2O2 production were obtained from photosynthetic and photorespiratory flux rates, derived from steady-state leaf gas exchange measurements at varying irradiance and ambient CO2. Assuming a 10% allocation of photosynthetic electron flow to the Mehler reaction, photorespiratory H2O2 production would account for about 70% of total H2O2 formed at all irradiances measured. When chloroplastic CO2 concentration rates are decreased, photorespiration becomes even more predominant in H2O2 generation. At the increased flux through photorespiration observed at lower ambient CO2, the Mehler reaction would have to account for more than 35% of the total photosynthetic electron flow in order to match the rate of peroxisomal H2O2 production. The potential signalling role of H2O2 produced in the peroxisomes is emphasized, and it is demonstrated that photorespiratory H2O2 can perturb the redox states of leaf antioxidant pools. We discuss the interactions between oxidants, antioxidants and redox changes leading to modified gene expression, particularly in relation to drought, and call attention to the potential significance of photorespiratory H2O2 in signalling and acclimation.

Acclimatization↗

The effects of drought on human nutrition in an Ethiopian province.

In mid 1974 the authors conducted a survey of Harerghe Province, Ethiopia in order to verify reports of serious effects of drought six months earlier. Information was obtained on anthropometric nutritional status, mortality, livestock, and market prices from a random sample of 62 villages in four ecological zones - the highlands, the northern and southern rangelands and an intermediate zone of mixed economy. The rangelands and intermediate areas had experienced heavy human and livestock losses during the drought but this had not occurred in the highlands. Although a high prevalence of malnutrition was found in every area studied, the geographical and height distribution of malnutrition within population groups suggested that this could only be clearly attributed to the effects of unusual food shortage in one part of the southern rangelands.

Adolescent↗

Dehydrin transcript fluctuations during a day/night cycle in drought-stressed sunflower.

To investigate environmental stimuli involved in the modulation of drought-induced gene expression, the influence of the day/night cycle on the expression of two dehydrin genes (HaDhn1 and HaDhn2) in leaves of sunflowers subjected to mild or severe drought stress has been studied. It was observed that the HaDhn1, but not HaDhn2, transcript oscillated in a diurnal fashion. In severely stressed plants, the peak of HaDhn1 mRNA accumulation occurred at midday.

Circadian Rhythm↗

Oxidative stress, heat shock and drought differentially affect expression of a tobacco protein phosphatase 2C.

A protein phosphatase 2C (PP2C)-homologous cDNA was isolated from Nicotiana tabacum (NtPP2C1). The deduced protein sequence of 416 amino acids showed the highest degree of similarity to the PP2C of Arabidopsis thaliana (AtPP2CA) implicated in abscisic acid signalling. The expression of NtPP2C1 was strongly induced by drought, but repressed by oxidative stress and heat shock. It is suggested that NtPP2C1 operates at the junction of drought, heat shock and oxidative stress.

Amino Acid Sequence↗

Drought differentially affects expression of a PR-10 protein, in needles of maritime pine (Pinus pinaster Ait.) seedlings.

A full length cDNA encoding a PR-10 protein was isolated from maritime pine drought-stressed seedlings. The predicted protein contained 150 amino acids, has a molecular mass of 16.7 kDa and an isoelectric point of 5.28. The transcript level of PR-10 displayed a transient accumulation in needles of drought-stressed plants, and was not detectable in root and stem tissues.

Amino Acid Sequence↗

Senescence- and drought-related changes in peroxidase and superoxide dismutase isoforms in leaves of Ramonda serbica.

Ramonda sp. (Gesneriaceae) is an endemic and relic plant in a very small group of poikilohydric angiosperms that are able to survive in an almost completely dehydrated state. Senescence- and drought-related changes in the activity of peroxidase (POD; EC 1.11.1.7), ascorbate peroxidase (EC 1.11.1.11), and superoxide dismutase (SOD; EC 1.15.1.1) were determined in leaves of different age and relative water content. The results indicate that different POD isoforms were stimulated during senescence and dehydration. Two of the soluble POD isoforms were anionic with pI 4.5, and two were cationic with pI 9.3 and 9.0. The activity of ascorbate peroxidase remained unchanged either by drought or senescence. For the first time, SOD isoforms have now been determined in this resurrection plant. Several SOD isoforms, all of the Mn type, were found to be anionic with pI 4 and a few others had pI from 5 to 6, while one band of FeSOD with a lower molecular weight was neutral. Rehydration brought about a remarkable decrease over the first hour in the activity of all the antioxidant enzymes examined but activity recovered 1 d after rehydration. The results confirmed that dehydration and senescence caused disturbance in the redox homeostasis of Ramonda leaves, while inducing different POD isoforms. A physiological role of peroxidase reaction with hydroxycinnamic acids in conservation and protection of cellular constituents of desiccated Ramonda leaves is suggested.

Aging↗

Barley transcript profiles under dehydration shock and drought stress treatments: a comparative analysis.

A microarray including 1654 cDNAs, mainly derived from dehydration-shocked barley leaf tissues, was utilized to monitor expression changes in leaves of barley plants subjected to slow drying conditions (7 d and 11 d: 7d-WS and 11d-WS) in soil and after rehydration. The results were compared with those obtained under shock-like conditions imposed with a 6 h dehydration treatment. A total number of 173 transcripts (approximately 10% of all transcripts profiled) were declared up- or down-regulated in at least one of the conditions tested. The majority of the transcripts were regulated by only one of the drought treatments, with 57% of the differentially expressed transcripts exclusively affected in the dehydration shock treatment, 6% at 7d-WS, 14% at 11d-WS, and 6% after rehydration. Irrespective of the low percentage of transcripts (10%) with similar expression changes between shock and slow stress treatments, a sizeable portion of these transcripts shared a common expression trend under the different drought treatment conditions, as evidenced by low but significant correlations between the fast occurring and the 7d-WS and 11d-WS treatments (r=0.32 and 0.41, P=0.001, respectively). These results are discussed with respect to the merit of different dehydration treatments in the investigation of the changes in transcript profiling.

Desiccation↗