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PubMed2006

Gene networks involved in drought stress response and tolerance.

Plants respond to survive under water-deficit conditions via a series of physiological, cellular, and molecular processes culminating in stress tolerance. Many drought-inducible genes with various functions have been identified by molecular and genomic analyses in Arabidopsis, rice, and other plants, including a number of transcription factors that regulate stress-inducible gene expression. The products of stress-inducible genes function both in the initial stress response and in establishing plant stress tolerance. In this short review, recent progress resulting from analysis of gene expression during the drought-stress response in plants as well as in elucidating the functions of genes implicated in the stress response and/or stress tolerance are summarized. A description is also provided of how various genes involved in stress tolerance were applied in genetic engineering of dehydration stress tolerance in transgenic Arabidopsis plants.

Abscisic Acid↗
PubMed2006

The drought environment: physical, biological and agricultural perspectives.

'Drought' has many meanings in relation to crop production. These range from: statistical (say, the lowest decile of annual rainfall) to a meteorologist; through yield being limited by too little water to an agronomist; to sudden severe water deficits to many molecular biologists. To a farmer, the corresponding management issues, respectively, are risk management (how best to manage a meteorologically drought-prone farm over several years), how best to match cultivar and agronomic operations to the developing growing season, and how best to minimize possible major damage to (say) floral fertility induced by severe water deficits during flowering. All these definitions and the issues they imply are relevant to improving crop production when water is limiting. How can scientists best help? The answers depend on the scales (temporal and spatial) being addressed. Agronomists and breeders, interacting, can help improve components of seasonal water balance in the field, for example, minimizing evaporative losses from the soil surface by better matching the development of a crop to its environment. Physiologists, biochemists, and molecular biologists can help by identifying ways of improving the competence of particular organs. A promising target is floral infertility resulting from water deficits, which results from lesions in tissue, and cellular and molecular processes. Choosing problems whose solutions will have implications in the field and be attractive to farmers requires knowledge of what is important in the field.

Agriculture↗
PubMed2006

Marker-assisted selection to improve drought adaptation in maize: the backcross approach, perspectives, limitations, and alternatives.

A number of different marker-assisted selection (MAS) approaches do exist for the improvement of polygenic traits. Results of a marker-assisted backcross (MABC) selection experiment aimed at improving grain yield under drought conditions in tropical maize are presented and compared with alternative MAS strategies. The introgression of favourable alleles at five target regions involved in the expression of yield components and flowering traits increased grain yield and reduced the asynchrony between male and female flowering under water-limited conditions. Eighty-five per cent of the recurrent parent's genotype at non-target loci was recovered in only four generations of MABC by screening large segregating populations (2200 individuals) for three of the four generations. Selected MABC-derived BC(2)F(3) families were crossed with two testers and evaluated under different water regimes. Mean grain yield of MABC-derived hybrids was consistently higher than that of control hybrids (crosses from the recurrent parent to the same two testers as the MABC-derived families) under severe water stress conditions. Under those conditions, the best five MABC-derived hybrids yielded, on average, at least 50% more than control hybrids. Under mild water stress, defined as resulting in <50% yield reduction, no difference was observed between MABC-derived hybrids and the control plants, thus confirming that the genetic regulation for drought tolerance is dependent on stress intensity. MABC conversions involving several target regions are likely to result in partial rather than complete line conversion. Simulations were conducted to assess the utility of such partial conversions, i.e. containing favourable donor alleles at non-target regions, for subsequent phenotypic selection. The results clearly showed that selecting several genotypes (10-20) at each MABC cycle was most efficient. In the light of these results, alternative approaches to MABC are discussed, including recurrent selection, illustrated by an example of improving the adaptation of maize to low temperatures. Given the current approaches for MAS and the choices of marker technologies available now and potential for future developments, the use of MAS techniques in further improving grain yield under abiotic stresses in maize appears very promising.

Crosses, Genetic↗
PubMed2006

Drought-adaptive traits derived from wheat wild relatives and landraces.

Exotic parents are being used to increase allelic diversity in bread wheat breeding through (i) interspecific hybridization of the ancestral genomes to produce so-called synthetic derived (SYN-DER) wheat, and (ii) crossing with landrace accessions, originating in abiotically stressed environments, that have become isolated from mainstream gene pools. Evaluation of the inherent genetic diversity encompassed by drought-adapted landraces compared with checks using DNA fingerprinting confirmed that some landraces were not only distant from checks but also showed significant diversity among each other. Improvement in performance of SYN-DER lines compared with recurrent parents was not associated with a larger overall investment in root dry weight, but rather an increased partitioning of root mass to deeper soil profiles (between 60 cm and 120 cm) and increased ability to extract moisture from those depths. The best Mexican landraces showed superior ability in terms of water extraction from soil depth, as well as increased concentration of soluble carbohydrates in the stem shortly after anthesis. Although it can be argued that inferring theoretical yield gains from the over-expression of any of these traits is questionable, since compensatory mechanisms may be at work, the fact remains that mechanistic or genetic linkages among physiological traits remain largely un-established. In the meantime, trait information is being used to make strategic crosses based on the theoretical combination of useful stress-adaptive traits with the possibility of realizing additive gene action in selected progeny. Candidates for crossing with elite check cultivars include landraces identified that showed relatively high biomass under drought combined with favourable expression of physiological traits such as stem carbohydrates, water extraction characteristics, and transpiration efficiency.

Adaptation, Physiological↗
PubMed2002

Identification and characterization of a gene encoding drought-inducible protein localizing in the bundle sheath cell of sugarcane.

We have identified a drought-inducible gene, designated as SoDip22, in sugarcane leaves. The cDNA encoded a hydrophilic protein with a calculated molecular mass of 15.9 kDa and the amino acid sequence was similar to that of ABA, stress and ripening-inducible protein from various plant species. ABA or mannitol-treatment of the detached leaves also induced SoDip22 expression. Stepwise homogenization of the stressed leaves showed that SoDip22 is localized in bundle sheath cells. These results suggest that SoDip22 functions to adapt to drought stress in the bundle sheath cell, and that the signaling pathway for the induction is, at least in a part, mediated by ABA.

Abscisic Acid↗
PubMed2005

Co-expression of cytochrome b561 and ascorbate oxidase in leaves of wild watermelon under drought and high light conditions.

Despite carrying out C3 photosynthesis, wild watermelon (Citrullus lanatus sp.) exhibits exceedingly good tolerance to severe drought at high light intensities. However, the mechanism(s) by which this plant protects itself from photodamage has yet to be elucidated. In this study, we characterized wild watermelon cytochrome b561 (cyt b561), which potentially mediates regeneration of apoplastic ascorbate by transferring electrons from cytosolic ascorbate across the plasma membrane. Two cDNA species for wild watermelon cyt b561, designated CLb561A and CLb561B, were isolated. Levels of both CLb561A mRNA and protein were significantly elevated in the leaves during drought at a light intensity of 700 micromol photons m(-2) s(-1). The transcript of CLb561B was detected to a much lesser extent, but no CLb561B protein was produced under any condition used in this study. A transient expression assay with the CLb561A::green fluorescent protein fusion construct showed clear fluorescence on the plasma membrane of onion epidermal cells. The CLb561A protein was enriched in the plasma membrane fraction in leaves of transgenic tobacco expressing CLb561A. Moreover, the high activity of apoplastic ascorbate oxidase (AO), which was able to dispose of cyt b561-transferred reducing equivalents, increased in leaves of wild watermelon grown at high light intensity, but not lower light intensities. Taken together, these observations suggest the occurrence of a novel pathway for excess light energy dissipation in wild watermelon leaves, where excessive energy absorbed by chloroplasts can be transported to and dissipated safely in the apoplasts through the cooperative action of cyt b561 and AO.

Amino Acid Sequence↗
PubMed2006

Characterization of transgenic Arabidopsis plants overexpressing high mobility group B proteins under high salinity, drought or cold stress.

High mobility group B (HMGB) proteins found in the nuclei of higher eukaryotes play roles in various cellular processes such as replication, transcription and nucleosome assembly. The Arabidopsis thaliana genome contains eight genes encoding HMGB proteins, the functions of which remain largely unknown in the transcriptional regulation of plant stress responses. To understand better the functions of HMGB proteins in the responses of plants to environmental stimuli, we examined the effect of various abiotic stresses on germination and growth of transgenic Arabidopsis plants that overexpress a single isoform of HMGB. The expression of HMGB2, HMGB3 and HMGB4 was up-regulated by cold stress, whereas the expression of HMGB2 and HMGB3 was markedly down-regulated by drought or salt stress. Under salt or drought stress, the transgenic Arabidopsis plants that overexpress HMGB2 displayed retarded germination and subsequent growth compared with wild-type plants. Overexpression of HMGB4 had no impact on seed germination and seedling growth of the plants under the stress conditions tested. In contrast to no significant stress-related phenotypes of HMGB5-overexpressing plants, loss-of-function mutants of HMGB5 displayed retarded germination and subsequent growth compared with wild-type plants under stress conditions. Although transcript levels of various stress-responsive genes were not modulated by the expression of HMGB2, expression of several germination-responsive genes was modulated by HMGB2 under salt stress. Taken together, these results provide a novel basis for understanding the biological functions of HMGB protein family members that differently affect germination and seedling growth of Arabidopsis plants under various stress conditions.

Arabidopsis↗
PubMed1993

Effects of elevated carbon dioxide and drought on the growth and physiology of clonal Sitka spruce plants (Picea sitchensis (Bong.) Carr.).

Two-year-old Sitka spruce (Picea sitchensis (Bong.) Carr.) plants from four clones were grown in naturally lit growth chambers for 6 months at either ambient (350 ppm) or ambient + 250 ppm (600 ppm) CO(2) concentration. Plants were grown in large boxes filled with peat, in a system that allowed the roots of individual plants to be harvested easily at the end of the growing season. Half of the boxes were kept well watered and half were allowed to dry out slowly over the summer. Plants growing in elevated CO(2) showed a 6.9% increase in mean relative growth rate compared to controls in the drought treatment and a 9.8% increase compared to controls in the well-watered treatment, though there was considerable variation in response among the different clones and water treatments. Rates of net CO(2) assimilation were higher and stomatal conductances were lower in plants grown in elevated CO(2) than in ambient CO(2) in both the well-watered and drought treatments. Both of these factors contributed to the doubling of instantaneous water use efficiency. The partitioning of biomass to roots was unaffected by elevated CO(2), but the ratio of needle mass/stems + branches mass decreased. Together with reduced stomatal conductance, this probably caused the observed increases in xylem pressure potentials with elevated CO(2).

Journal Article↗
PubMed1994

Effects of soil freezing and drought stress on abscisic acid content of sugar maple sap and leaves.

In 1991 and 1992, mature maple trees (Acer saccharum Marsh.) were freeze-stressed or drought-stressed by preventing precipitation (snow or rain) from reaching the forest floor under selected trees. Lack of snow cover caused a decrease in soil temperature to well below 0 degrees C from December to April and a lowering of the soil water content to 10%. The abscisic acid (ABA) concentration in the spring sap of deep-soil frost-stressed trees was significantly higher than in control or drought-stressed trees. The increase in ABA concentration in the xylem sap in the spring of 1991 and 1992 preceded symptoms of canopy decline and a decrease in leaf area that were observed during the summers of 1991 and 1992. These results suggest a role for ABA in root-to-shoot communication in response to environmental stress. The largest differences in ABA concentration induced by the treatments was found in sap collected at the end of sap flow. The increase in ABA concentration in spring sap at the end of the sap flow could be used as an early indicator of stress suffered by trees during the winter. Not only did the increase in ABA concentration occur before any visible symptoms of tree decline appeared, but the trees that showed the most evident decline had the highest ABA concentrations in the spring sap. Leaf ABA concentration was not a good indicator of induced stress.

Journal Article↗
PubMed1994

Vulnerability to drought-induced cavitation of riparian cottonwoods in Alberta: a possible factor in the decline of the ecosystem?

Vulnerability of xylem to loss of hydraulic conductivity caused by drought-induced cavitation was determined for three riparian cottonwood species in Lethbridge, Alberta: Populus deltoides Bartr., P. balsamifera L., and P. angustifolia James. These species suffered 50% loss of hydraulic conductivity in one-year-old stem segments when xylem pressure potential fell to -0.7 MPa for P. deltoides and -1.7 MPa for P. balsamifera and P. angustifolia, making them the three most vulnerable tree species reported so far in North America. The possible contribution of drought-induced xylem dysfunction to the decline of riparian ecosystems in dammed rivers is discussed.

Journal Article↗
PubMed1996

High temperature and drought stress effects on survival of Pinus ponderosa seedlings.

We studied the effects of high temperature and drought on the survival, growth and water relations of seedlings of Pinus ponderosa (Dougl.) Lawson, one of few coniferous tree species that can successfully colonize drought-prone sites with high soil surface temperatures. Temperature profiles were measured with 0.07-mm thermocouples in a sparse ponderosa pine forest in northern Idaho. The soil surface and the adjacent 5 mm of air reached maximum temperatures exceeding 75 degrees C, well above the lethal temperature threshold for most plants. Air temperatures 50 mm above the soil surface (seedling needle height) rarely exceeded 45 degrees C. Pinus ponderosa seedlings that survived maintained basal stem temperatures as much as 15 degrees C lower than the surrounding air. The apparent threshold temperature at the seedling stem surface resulting in death was approximately 63 degrees C for less than 1 min. No correlation between seedling mortality and needle temperature was found, although some needles reached temperatures as high as 60 degrees C for periods of </= 1 min. Surviving seedlings had significantly higher stomatal conductance than seedlings that did not survive until fall. Transpiration rates, calculated from measured needle temperatures, stomatal conductance and evaporative demand, were high (up to 16.7 mmol m(-2) s(-1)), indicating that water transport through seedling stems may have acted as a heat transfer mechanism, cooling the stem below the lethal threshold temperature. Heat exchange calculations showed that rapid water flow through seedling stems can absorb sufficient energy to reduce stem temperature by 30 degrees C during peak sunlight hours.

Journal Article↗
PubMed2000

Observations of evapotranspiration in a break of slope plantation susceptible to periodic drought stress.

Break of slope (BOS) plantations are advocated as a means of water table control in areas where groundwater flows through colluvial deposits overlying low permeability bedrock. It is also believed that BOS plantations can supplement their water use requirements by exploiting shallow groundwater at the breaks in topographic slope. We compared measurements of BOS plantation and pasture evapotranspiration during spring, when the weather was warm and soils moist, and late summer when drought conditions prevailed. Microlysimeters and ventilated chambers were used to determine pasture and plantation floor evaporation, and heat pulse sensors were used to determine transpiration of the plantation. In spring, pasture evapotranspiration was 65% of that of the plantation, whereas, in summer, pasture evapotranspiration was equivalent to only 35% of that of the plantation. Rainfall interception by the canopy of the plantation was twice that of the pasture, reinforcing the notion that trees can help reduce groundwater recharge and alleviate dryland salinity and waterlogging. During the summer drought period, daily plantation transpiration was only 20% of that measured during spring, suggesting that the plantation was not utilizing groundwater supplies but was instead drawing from soil water supplies. This hypothesis was supported by the comparison of relative abundances of the isotopes of water ((2)H and (18)O) in soil and wood samples. We conclude that the BOS plantation is not behaving in the manner predicted, and our findings raise doubts about the predicted advantages of establishing plantations in break of slope positions.

Journal Article↗
PubMed2001

Response to seasonal drought in three cultivars of Ceratonia siliqua: leaf growth and water relations.

Responses of leaf growth and leaf water relations to seasonal drought were monitored during two successive years in three cultivars (Galhosa, Espargal and Mulata) of Ceratonia siliqua L. growing in the field in southern Portugal. Leaf water relations of fully expanded leaves were characterized by pressureâ volume analysis, and morphometric measurements of petiolar xylem were made. The three cultivars differed with respect to onset of leaf initiation. In Galhosa, there was a sharp peak of leaf initiation in June that was immediately followed by a period when leaf expansion rates were highest. The onset of leaf growth occurred earlier in Espargal and Mulata than in Galhosa, and both cultivars continued producing new leaves throughout the summer period. The diurnal pattern of water relations in recently expanded leaves indicated that, during midsummer, Galhosa was the only cultivar in which leaf water potential did not fall below the turgor loss point and leaf relative water content remained above 90%. The occurrence of osmotic adjustment in recently expanded leaves of Galhosa was not demonstrated conclusively. However, during the dry season following leaf formation, a seasonal decrease in osmotic potential sufficient to maintain turgor was detected in 1-year-old leaves of Galhosa but not in 1-year-old leaves of the other cultivars. Among cultivars, Galhosa petioles had the widest xylem conduits, which may partly explain why midday leaf water potential in Galhosa never decreased below -2.0 MPa even at the end of the summer drought.

Fabaceae↗
PubMed2001

Response of stomatal conductance to drought in ponderosa pine: implications for carbon and ozone uptake.

To gain insight into the limitations imposed by a typical Mediterranean-climate summer drought on the uptake of carbon and ozone in the ponderosa pine (Pinus ponderosa Dougl. ex Laws.) ecosystem, we compared diurnal trends in leaf physiology of young trees in a watered and a control plot located in the Sierra Nevada Mountains, CA, USA (Blodgett Forest, 38 degrees 53' N, 120 degrees 37' W, 1315 m elevation). Predawn water potential of trees in the watered plot remained above -0.3 MPa throughout the growing season, whereas it dropped in the control plot from -0.24 to -0.52 MPa between late May and mid-August. Photosynthesis and stomatal conductance of trees in the watered plot were relatively insensitive to atmospheric vapor pressure deficit (VPD), whereas gas exchange of trees in the control plot varied with changes in soil water, VPD and temperature. Although the 1998 growing season was abnormally wet, we saw a pronounced drought effect at the control site. Over the 2 months following the onset of watering, carbon and ozone uptake were measured on three days at widely spaced intervals. Carbon uptake per unit leaf area by 1-year-old foliage of trees in the control plot was 39, 35 and 30% less, respectively, than in the watered plot, and estimated ozone deposition per unit leaf area (ozone concentration times stomatal conductance) was 36, 46 and 41% less.

California↗
PubMed2002

Enhanced tolerance of photosynthesis to high-light and drought stress in Pseudotsuga menziesii seedlings grown in ultraviolet-B radiation.

We investigated the effects of an ambient dose of ultraviolet-B (UV-B) radiation on chamber-grown Pseudotsuga menziesii var. glauca (Beissn.) Franco (Douglas-fir) seedlings, to determine if the presence of UV-B radiation in the growth light regime induces tolerance to environmental stresses such as high light and drought. Douglas-fir seedlings were grown without UV-B radiation or with 6 kJ m-2 day-1 of biologically effective UV-B, which is ambient for the intermountain regions of Idaho. Non-stressed seedlings grown with UV-B radiation had 35% lower seedling dry mass, 36% higher concentrations of UV-B absorbing compounds per unit leaf area, 30% lower stomatal frequencies, 25% lower light-saturated photochemical efficiencies of Photosystem II and 45% lower light-saturated stomatal conductance than non-stressed seedlings grown without UV-B radiation. After 4 days of high-light stress, seedlings grown with UV-B radiation had 32% higher light-saturated carbon assimilation rates (A(CO2)) than seedlings grown without UV-B radiation. After water was withheld from the seedlings for up to 15 days, seedlings grown with UV-B radiation had 50% higher A(CO2) and 40% higher seedling water potentials than seedlings grown without UV-B radiation. The results support the hypothesis that UV-B radiation can act as an environmental signal to induce tolerance to high-light and drought stress in Douglas-fir seedlings. Possible mechanisms for the enhanced stress tolerance are discussed.

Dehydration↗
PubMed2003

Osmotic adjustment in three-year-old seedlings of five provenances of maritime pine (Pinus pinaster) in response to drought.

Three-year-old seedlings of five provenances of Pinus pinaster Ait. that differed in climatic conditions at their geographical origin were subjected to decreasing soil water availability. The degree of needle osmotic adjustment (OA) was estimated based on logarithmic plots of needle relative water content (RWC) against needle osmotic potential (Psi(pi)); i.e., lnRWC versus -ln(-Psi(pi)). There were significant differences among provenances in active OA (0.13 to 0.30 MPa for a decrease in RWC to 80%), and a clear negative relationship was found between OA and precipitation (650 to 1280 mm of mean annual rainfall) at the geographical origins of the provenances. A high osmoregulatory capacity contributes to the maintainance of positive turgor at low water potentials. We conclude that OA is one of the mechanisms underlying adaptation to drought in P. pinaster. Solute accumulation was about 2.3 times higher in the provenance from the driest site than in the provenance from the wettest site. The contribution of osmotic adjustment to differences in drought tolerance mechanisms among provenances is discussed.

Adaptation, Physiological↗
PubMed2004

Stomatal limitation to CO2 assimilation and down-regulation of photosynthesis in Quercus ilex resprouts in response to slowly imposed drought.

Holm oak (Quercus ilex L.) is native to hot, dry Mediterranean forests where limited water availability often reduces photosynthesis in many species, and forest fires are frequent. Holm oaks resprout after a disturbance, with improved photosynthetic activity and water relations compared with unburned plants. To better understand the role of water availability in this improvement, watering was withheld from container-grown plants, either intact (controls) or resprouts after excision of the shoot, to gradually obtain a wide range of soil water availabilities. At high water availability, gas exchange rates did not differ between controls and resprouts. At moderate soil dryness, net photosynthesis of control plants decreased as a result of increased stomatal limitation, whereas gas exchange rates of resprouts, which had higher midday and predawn leaf water potentials, were unchanged. Under severe drought, resprouts showed a less marked decline in gas exchange than controls and maintained photosystem II integrity, as indicated by chlorophyll fluorescence measurements. Photosynthesis was down-regulated in both plant types in response to reduced CO2 availability caused by high stomatal limitation. Lower non-stomatal limitations in resprouts than in control plants, as evidenced by higher carboxylation velocity and the capacity for ribulose-1,5-bisphosphate regeneration, conferred greater drought resistance under external constraints similar to summer conditions at midday.

Carbon Dioxide↗
PubMed2005

Responses of deciduous forest trees to severe drought in Central Europe.

In 2003, Central Europe experienced the warmest summer on record combined with unusually low precipitation. We studied plant water relations and phenology in a 100-year- old mixed deciduous forest on a slope (no ground water table) near Basel using the Swiss Canopy Crane (SCC). The drought lasted from early June to mid September. We studied five deciduous tree species; half of the individuals were exposed to elevated CO(2) concentration ([CO(2)]) (530 ppm) using a free-air, atmospheric CO(2)-enrichment system. In late July, after the first eight weeks of drought, mean predawn leaf water potential about 30 m above ground was -0.9 MPa across all trees, dropping to a mean of -1.5 MPa in mid-August when the top 1 m of the soil profile had no plant accessible moisture. Mean stomatal conductance and rates of maximum net photosynthesis decreased considerably in mid-August across all species. However, daily peak values of sap flow remained surprisingly constant over the whole period in Quercus petraea (Matt.) Liebl., and decreased to only about half of the early summer maxima in Fagus sylvatica L. and Carpinus betulus L. (stomatal down- regulation of flux). Although we detected no differences in most parameters between CO(2)-treated and control trees, predawn leaf water potential tended to be less negative in trees exposed to elevated [CO(2)]. Leaf longevity was greater in 2003 compared with the previous years, but the seasonal increase in stem basal area reached only about 75% of that in previous years. Our data suggest that the investigated tree species, particularly Q. petraea, did not experience severe water stress. However, an increased frequency of such exceptionally dry summers may have a more serious impact than a single event and would give Q. petraea a competitive advantage in the long run.

Betulaceae↗