Quantifying relationships between rooting traits and water uptake under drought in Mediterranean barley and durum wheat

In Mediterranean regions drought is the major factor limiting spring barley and durum wheat grain yields. This study aimed to compare spring barley and durum wheat root and shoot responses to drought and quantify relationships between root traits and water uptake under terminal drought.One spring barley（Hordeum vulgare L. cv. Rum） and two durum wheat Mediterranean cultivars（Triticum turgidum L. var durum cvs Hourani and Karim） were examined in soil‐column experiments under well watered and drought conditions. Root system architecture traits, water uptake, and plant growth were measured. Barley aerial biomass and grain yields were higher than for durum wheat cultivars in well watered conditions. Drought decreased grain yield more for barley（47%） than durum wheat（30%, Hourani）. Root‐to‐shoot dry matter ratio increased for durum wheat under drought but not for barley, and root weight increased for wheat in response todrought but decreased for barley. The critical root length density（RLD） and root volume density（RVD） for 90% available water capture for wheat were similar to（cv. Hourani） or lower than（cv. Karim） for barley depending on wheat cultivar. For both species, RVD accounted for a slightly higher proportion of phenotypic variation in water uptake under drought than RLD.     

Genomic dissection of drought resistance in durum wheat × wild emmer wheat recombinant inbreed line population

Drought is the major factor limiting wheat productivity worldwide. The gene pool of wild emmer wheat, Triticum turgidum ssp. dicoccoides , harbours a rich allelic repertoire for morpho-physiological traits conferring drought resistance. The genetic and physiological bases of drought responses were studied here in a tetraploid wheat population of 152 recombinant inbreed lines (RILs), derived from a cross between durum wheat (cv. Langdon) and wild emmer (acc# G18-16), under contrasting water availabilities. Wide genetic variation was found among RILs for all studied traits. A total of 110 quantitative trait loci (QTLs) were mapped for 11 traits, with LOD score range of 3.0–35.4. Several QTLs showed environmental specificity, accounting for productivity and related traits under water-limited (20 QTLs) or well-watered conditions (15 QTLs), and in terms of drought susceptibility index (22 QTLs). Major genomic regions controlling productivity and related traits were identified on chromosomes 2B, 4A, 5A and 7B. QTLs for productivity were associated with QTLs for drought-adaptive traits, suggesting the involvement of several strategies in wheat adaptation to drought stress. Fifteen pairs of QTLs for the same trait were mapped to seemingly homoeologous positions, reflecting synteny between the A and B genomes. The identified QTLs may facilitate the use of wild alleles for improvement of drought resistance in elite wheat cultivars.     

Photosynthetic characteristics and enzymatic antioxidant capacity of leaves from wheat cultivars exposed to drought

Two durum (Triticum durum L.), Barakatli-95 and Garagylchyg-2; and two bread (Triticum aestivum L.) wheat cultivars, Azamatli-95 and Giymatli-2/17 with different sensitivities to drought were grown in the field on a wide area under normal irrigation and severe water deficit. Drought caused a more pronounced inhibition in photosynthetic parameters in the more sensitive cvs Garagylchyg-2 and Giymatli-2/17 compared with the tolerant cvs Barakatli-95 and Azamatli-95. Upon dehydration, a decline in total chlorophyll and relative water content was evident in all cultivars, especially in later periods of ontogenesis. Potential quantum yield of PS II (F(v)/F(m) ratio) in cv Azamatli-95 was maximal during stalk emergency stage at the beginning of drought. This parameter increased in cv Garagylchyg-2, while in tolerant cultivar Barakatli-95 significant changes were not observed. Contrary to other wheat genotypes in Giymatli-2/17 drought caused a decrease in PS II quantum yield. Drought-tolerant cultivars showed a significant increase in CAT activity as compared to control plants. In durum wheat cultivars maximal activity of CAT was observed at the milk ripeness and in bread wheat cultivars at the end of flowering. APX activity also increased in drought-treated leaves: in tolerant wheat genotypes maximal activity occurred at the end of flowering, in sensitive ones at the end of ear formation. GR activity increased in the tolerant cultivars under drought stress at all stages of ontogenesis. SOD activity significantly decreased in sensitive cultivars and remained at the control level or increased in resistant ones. This article is part of a Special Issue entitled: Photosynthesis Research for Sustainability: from Natural to Artificial.     

Genetic mapping of the gene affecting polyphenol oxidase activity in tetraploid durum wheat

The quality of durum wheat (Triticum turgidum ssp. durum) is influenced by polyphenol oxidase(PPO) activity and its corresponding substrates. A saturated molecular-marker linkage map was constructed previously by using a set of recombinant inbred (RI) lines, derived from a cross between durum wheat cultivars Jennah Khetifa and Cham 1. Quantitative trait loci (QTL) for PPO activity in seeds were mapped in this population. PPO activity in seeds of the parents and 110 RI lines was measured spectrophotometrically. The PPO activity of Cham 1 was significantly lower than that of Jennah Khetifa. QTL analysis of these data indicated that most of PPO activity was associated with major loci on the long arm of chromosome 2A. The trait was found to be strongly associated with the SSR marker Xgwm312@2A. With this knowledge, marker-assisted selection can be used to select genotypes with lower PPO activity in durum wheat populations.     

Effects of in vivo copper treatment on the photosynthetic apparatus of two Triticum durum cultivars with different stress sensitivity

The effect of copper on the photosynthetic apparatus of two cultivars of durum wheat ( Triticum durum cvs Adamello and Ofanto) with different sensitivity to drought and nickel stress were investigated. Plants were grown in nutrient solution or in nutrient solution further supplemented with CuSO₄ to achieve final concentrations of 3.6 μM and 20 μM Cu. Several fluorescence analyses were performed, in presence or absence of DCMU, and with varying light intensities. Furthermore, light and electron microscopic investigations were carried out. In vivo treatment using 3.6 μ Cu produced a marked reduction in growth of the Cu-treated plants, but only mild effects on the fluorescence-related parameters. The Cu-induced reduction in the area above the fluorescence induction curve and in the time needed to reach the maximum of chlorophyll fluorescence (F_max) were more pronounced. These results favour the hypothesis that under such conditions copper affects photosynthesis mainly in an indirect way, causing a slowing down of the electron transport as a consequence of the reduced requirement for photosynthesis products. The morphological analyses corroborate this hypothesis, showing toxic effects on the chloroplast structure due to Cu treatment. The differences between the two cultivars were not as pronounced as reported in the case of nickel or drought stresses; nevertheless, cv. Ofanto seemed to be less sensitive also to Cu stress than cv. Adamello.     

Photosynthetic performance of two coffee species under drought

Coffea arabica cv. Red Catuaí and C. canephora cv. Kouillou were grown in pots beneath a plastic shelter. When they were 14 months old, irrigation was withheld until the leaf pre-dawn water potential was about -1.5 and -2.7 MPa (designated mild and severe water stress, respectively). Under mild stress, net photosynthetic rate (PN) decreased mainly as a consequence of stomatal limitations in Kouillou, whereas such decreases were dominated by non-stomatal limitations in Catuaí. Under severe drought, further decreases in PN and apparent quantum yield were not associated to any changes in stomatal conductance in either cultivar. Decreases were much more pronounced in Catuaí than in Kouillou, the latter maintained carbon gain at the expense of water conservation. In both cultivars the initial chlorophyll (Chl) fluorescence slightly increased with no changes in the quantum efficiency of photosystem 2. In response to rapidly imposed drought, the Chl content did not change while saccharide content increased and starch content decreased. Photoinhibition and recovery of photosynthesis, as evaluated by the ratio of variable to maximum fluorescence and by the photosynthetic O2 evolution, were unaffected by mild drought stress. Photoinhibition was enhanced under severe water deficit, especially in Catuaí. In this cultivar the O2 evolution did not resume upon reversal from photoinhibition, in contrast to the complete recovery in Kouillou. This revised version was published online in June 2006 with corrections to the Cover Date.     

Photosynthetic performance of two coffee species under drought

Coffea arabica cv. Red Catuaí and C. canephora cv. Kouillou were grown in pots beneath a plastic shelter. When they were 14 months old, irrigation was withheld until the leaf pre-dawn water potential was about -1.5 and -2.7 MPa (designated mild and severe water stress, respectively). Under mild stress, net photosynthetic rate (PN) decreased mainly as a consequence of stomatal limitations in Kouillou, whereas such decreases were dominated by non-stomatal limitations in Catuaí. Under severe drought, further decreases in PN and apparent quantum yield were not associated to any changes in stomatal conductance in either cultivar. Decreases were much more pronounced in Catuaí than in Kouillou, the latter maintained carbon gain at the expense of water conservation. In both cultivars the initial chlorophyll (Chl) fluorescence slightly increased with no changes in the quantum efficiency of photosystem 2. In response to rapidly imposed drought, the Chl content did not change while saccharide content increased and starch content decreased. Photoinhibition and recovery of photosynthesis, as evaluated by the ratio of variable to maximum fluorescence and by the photosynthetic O2 evolution, were unaffected by mild drought stress. Photoinhibition was enhanced under severe water deficit, especially in Catuaí. In this cultivar the O2 evolution did not resume upon reversal from photoinhibition, in contrast to the complete recovery in Kouillou.     

Water deficit during root development: effects on the growth of roots and osmotic water permeability of isolated root protoplasts

The effect of low water potentials on root growth of flax (Linum usitatissimum L. cv. Ariane), rape (Brassica napus L. de Candolle, cv. Bristol), hard wheat (Triticum turgidum L. cv. Cham1) and soft wheat (Triticum aestivum L. cv. Ritmo) was studied by measuring the osmotic water permeability (Pos) of root protoplasts and the protein abundance of PIP1 and PIP2 aquaporins. These different species require more or less water, the most sensitive to water deficit being flax and rape. Ritmo, is a cultivar of wheat adapted to temperate zones, while the other cultivar Cham1 is adapted to low-rainfall areas. The seedlings were germinated and grown in water, salt or sugar solutions at different water potentials. The values of Pos for flax, rape and Chaml wheat were normally distributed and could be characterized by mean +/- SD. Root protoplasts from water-grown seedlings had Pos values of 485+/-159 microm s(-1) (flax), 582+/-100 microm s(-1) (rape), and 6.3+/-3.5 microm s(-1) (Cham1). At the same age, the protoplasts from Ritmo exhibited a much wider range of values than the protoplasts of Cham1. When seedlings were grown under conditions of osmotic or salt stress, the mass of the roots was reduced for all species. With 0.25 mol kg(-1) sorbitol or 0.125 M NaCl, the Pos for flax, rape and Cham1 remained constant or slightly increased, while for Ritmo the reduction in the mass of the roots was paralleled by a reduction in Pos. Only Cham1 and Ritmo were able to germinate at a lower potential (0.5 mol kg(-1) sorbitol). For Ritmo the reduction in the mass of the roots was paralleled by a reduction in Pos when grown in this stress condition and both wheats exhibited low Pos values. The expression of the PIP1 and PIP2 aquaporins families was also studied by immunoblotting. We did not observe any difference in protein expression for any of the species, whatever the growing conditions. We suggest that the high Pos values for flax and rape could play a role in the sensitivity of these plants at low water potential. The low native Pos for Cham1 in spite of the expression of both families of aquaporins may reflect its adaptation to low-rainfall conditions by a functional regulation of the water channels. For a similar reason, the low-water-potential-induced Pos of Ritmo may also correspond to a down-regulation of the aquaporins, reflecting adaptation of this wheat to water-deficit conditions.     

Pressure-volume curves and drought resistance in two wheat genotypes

The water relations of two durum wheat cultivars ( Triticum durum Desf.) were studied throughout the growing season. Irrigated and unirrigated plants were compared from booting to milk stage; a period where water stress occurred naturally in the field. Modulus of elasticity (ε), turgid weight/dry weight ratio (TW/DW), relative water content at zero turgor (RWC_o) and osmotic potential at full turgor (ε) declined throughout the season while average turgor (ψ_p) increased. Water stress induced a further decrease in ψ_π¹⁰⁰ and the TW/DW ratio. The elastic modulus varied greatly. During the first stages of growth, cv. Appulo (the more resistant cultivar) showed lower ε values than cv. Valforte. At the milk stage, ε was lower for the unirrigated than the irrigated plants. Correlation coefficients between the TW/DW ratio and the osmotic potential were significant for both cultivars. In cv. Valforte, TW/DW was also correlated with the average turgor and the bulk modulus of elasticity. Structural changes that affect the TW/DW ratio seem to be important factors influencing water relations and drought tolerance in durum wheat.     

小麦叶片光合作用与其渗透调节能力的关系

在缓慢干旱条件下,小麦叶片渗透调节能力在一定范围内随胁迫程度的加剧而增加,而在快速干旱下,渗透调节能力丧失。小麦叶片通过渗透调节使光合速率和气孔导度对水分胁迫的敏感性降低,叶片维持较高的电子传递能力、RuBP羧化酶活性和叶绿体光合能量转换系统活性,并推迟了小麦叶片光合速率受气孔因素限制向叶肉细胞光合活性限制转变的时间。     

A new reduced height gene found in the tetraploid semi-dwarf wheat landrace Aiganfanmai

Aiganfanmai is a dwarf tetraploid wheat landrace (Triticum turgidum var. turgidum) that stably produces the semi-dwarf trait. Plant height varies from 80-105 cm under cultivation. Compared with tall durum wheat (T. turgidum var. durum) variety Langdon, we found it to have short spikes and seeds, besides a semi-dwarf character. We crossed Aiganfanmai with Langdon to analyze the genetic basis of the semi-dwarf trait. The F(2) population segregated at a 1:3 ratio for the short trait to the normal, which demonstrates that Aiganfanmai carries a recessive reduced height (Rht) gene. This gene was found to be located between the molecular markers Xgwm471 and Xgwm350 on chromosome arm 7AS by microsatellite analysis. No Rht gene had been reported from this chromosome; we designated it as Rht22. Rht 22, unlike other previously reported Rht genes, does not reduce internodal cell length. Reduced cell numbers might explain the short stem trait.     

The influence of the mycorrhiza Glomus etunicatum on drought acclimation in safflower and wheat

A study was done to determine the effects of vesicular‐arbuscular mycorrhizal (VAM) colonization on drought acclimation of host plants. Safflower ( Carthamus tinctorius L. cv. S555) and wheat ( Triticum aestivum L. cv. Anza) were grown under environmentally controlled conditions with or without the VAM fungus, Glomus etunicatum Becker and Gerd., and were either acclimated (by pre‐exposing plants to a 10–11 day drought period) or unacclimated to drought. Plants from all treatments were then exposed to drought for 9 days, and plant water status and root water uptake were measured. To minimize interactions between drought and P uptake, growth periods were adjusted so that acclimated and unacclimated plants were similar in size when measurements were made. When wheat was acclimated to drought, osmotic adjustment occurred (leaf solute potentials of leaf tissue rehydrated to full turgor were approximately 0.5 MPa lower in acclimated than unacclimated plants); in safflower, osmotic adjustment was minimal when plants were acclimated. Consequently, acclimated wheat plants were able to tolerate drought better than unacclimated plants, and maintained higher leaf water potentials and relative water contents as soil water was depleted. For both safflower and wheat, acclimated plants had higher water use efficiency, and therefore produced more biomass when water availability was limited, than unacclimated plants. However, mycorrhizal colonization did not affect osmotic adjustment, plant water status, water use efficiency or water uptake in either plant species, and therefore had no effect on drought acclimation or resistance.     

Performance of wheat crops with different chromosome ploidy: root-sourced signals, drought tolerance, and yield performance

Pot-culture experiments were carried out to estimate the role of non-hydraulic root signals (nHRS) and the relation of these signals to drought tolerance and grain yield formation under drought stress in six wheat varieties. These were two modern hexaploid wheat (Triticum aestivum L., AABBDD) Plateau602 and Longchun8139-2, two diploid wheat (Triticum monococcum L., AB) MO1 and MO4, and two tetraploid wheat (Triticum dicoccum Schuebl L., AABB) DM22 and DM31. In the two diploid relatives, the nHRS was switched on and off at a soil water content (SWC) of approximately 53–45% field water capacity (FWC). In contrast, in the modern hexaploid varieties, Longchun8139-2 and Plateau602 the nHRS occurred between a SWC of about 71 and 35% FWC, a much wider soil moisture range. The two tetraploid relatives, DM22 and DM31, were generally intermediate. The nHRS threshold range in SWC also narrowed as all six varieties went through successive developmental stages from shooting to grain filling. The two hexaploid wheat varieties had the longest duration of survival after the water supply ceased, and the best yield stability under drought stress, similar to with tetraploid wheat varieties; the diploid wheat varieties were least robust. These two parameters were both significantly correlated with the nHRS soil moisture threshold range (r=0.9456** and 0.8608*, respectively). Based on these patterns, we propose a ‘triple Z’ model to describe the features of non-hydraulic stomatal sensitivity versus soil drought in wheat growth.     

Effect of excess Cu on the photosynthetic apparatus of runner bean leaves treated at two different growth stages

Variations in water binding strength, water relations and the accumulation of solutes during water stress of three durum wheat ( Triticum durum Desf.) cultivars are reported and discussed. Water binding strength was determined by constructing adsorption isotherms at 5 and 20°C and by calculating the differential enthalpy (ΔH) after van't Hoff.
Reducing sugars, proline, K⁺ and Cl⁻ were the major contributors to osmotic adjustment. Solutes, such as quaternary ammonium compounds, and non-reducing sugars contributed to osmotic potential at full turgor, but did not increase in proportion to water stress. Genotypic differences have been observed for K⁺ accumulation capability, the water-stressed leaves of cv. Capeiti 8 showing the largest increase. The same cultivar demonstrated the most negative ΔH values, indicative of strongly bound water, and the highest integrated enthalpy (ΔH_inf) values for leaf moisture below 0.1 g H₂O per g dry weight, i.e. in the isotherm region where water was presumably chemisorbed to the charged groups of macromolecules. The accumulation of ions (Cl⁻, K⁺) and proline was concurrent with an increase in the binding-strength of tightly and weakly bound water, respectively.     

Characterization of drought resistance in a wild relative of wheat,Triticum kotschyi

Wild relatives of wheat have served as a genetic source for economically useful traits. A better understanding of the mechanisms underlying such traits may be useful in the genetic transfer and selection processes. Research was undertaken to compare the effects of controlled water stress on photosynthetic parameters in Triticum kotschyi, a drought resistant wild wheat and Triticum aestivum cv. Lakhish, a drought sensitive wheat cultivar. During stress development, the leaf water potential decreased at a slower rate, and the quantum yield of oxygen evolution, measured photoacoustically in vivo, decreased to a smaller extent in the drought resistant wild wheat than in the wheat cultivar. The decrease in quantum yield at water potentials from –0.9 Mpa down to –2.3 Mpa was not accompanied by damage to PS II reaction centers as there was no change in variable fluorescence. Below –2.3 Mpa the fluorescence yield of both species decreased indicating loss of intrinsic efficiency of PS II. The osmotic potential of cell sap was found to decrease at the same rate in both species at high hydration states. Proline accumulated to a much greater extent in the wild wheat as compared to the cultivated wheat as a result of water stress. Drought resistance was also examined in relation to thylakoid membrane fluidity measured by fluorescence polarization. Thylakoid membrane fluidity was fully maintained in the wild wheat, but decreased substantially in the wheat cultivar, at equal tissue water potentials below –1.9 Mpa. One mechanism for maintaining the higher quantum yield of oxygen evolution during severe stress (at water potentials below –1.9 Mpa), may involve the greater stability of thylakoid membrane fluidity in the wild wheat.Abbreviations DPH 1,6-diphenyl-1,3,5-hexatriene - LHC II light-harvesting chlorophyll-protein a/b complex - LWP leaf water potential - PS I, PS II Photosystem I, II - RUBPcase ribulose 1,5-bisphosphate carboxylase     

水分胁迫及复水过程中小麦抗氧化酶的变化

对两个抗旱性不同的小麦品种进行水分胁迫和复水处理,研究其抗氧化酶活性的响应。在水分胁迫下,陇春-20的相对含水量高于优鉴-24,复水24h后,优鉴-24的相对含水量恢复较快且高于陇春-20。水分胁迫下,优鉴-24中H2O2含量增加迅速,而且各阶段含量均高于陇春-20,复水后两个品种的H2O2含量都下降,这表明优鉴-24在水分胁迫时受到更严重的氧化胁迫。采用温和胶电泳结合抑制剂实验发现小麦有3条Mn—SOD,一条Fe—SOD和Cu／Zn-SOD同工酶带,CAT同工酶有3条谱带。在水分胁迫和复水期间,优鉴-24的SOD和CAT活性高于陇春-20,随着水分胁迫程度的增加,两个品种的SOD和CAT活性都增强,复水后,优鉴-24的SOD活性继续增强,而陇春-20的Mn—SOD—3活性略微降低,Fe—SOD和Cu／Zn—SOD活性略微升高,陇春-20的CAT活性降低。水分胁迫诱导了Mn—SOD—1在优鉴-24及Mn—SOD-2和Fe—SOD在陇春-20中的表达。     

Wheat grain yield on saline soils is improved by an ancestral Na⁺ transporter gene

The ability of wheat to maintain a low sodium concentration ([Na(+)]) in leaves correlates with improved growth under saline conditions. This trait, termed Na(+) exclusion, contributes to the greater salt tolerance of bread wheat relative to durum wheat. To improve the salt tolerance of durum wheat, we explored natural diversity in shoot Na(+) exclusion within ancestral wheat germplasm. Previously, we showed that crossing of Nax2, a gene locus in the wheat relative Triticum monococcum into a commercial durum wheat (Triticum turgidum ssp. durum var. Tamaroi) reduced its leaf [Na(+)] (ref. 5). Here we show that a gene in the Nax2 locus, TmHKT1;5-A, encodes a Na(+)-selective transporter located on the plasma membrane of root cells surrounding xylem vessels, which is therefore ideally localized to withdraw Na(+) from the xylem and reduce transport of Na(+) to leaves. Field trials on saline soils demonstrate that the presence of TmHKT1;5-A significantly reduces leaf [Na(+)] and increases durum wheat grain yield by 25% compared to near-isogenic lines without the Nax2 locus.     

Rapid Osmotic Adjustment in Detached Wheat Leaves

Osmotic adjustment is induced in detached wheat leaves by rapiddrying to a relative water content below 0·65, followedby re-saturation. Quantitatively, the response to this treatmentis comparable to the maximum of adjustment obtained with pottedplants at the same developmental stage. Low temperatures duringdrying and re-saturation of the leaves reduce the adjustmentresponse. We conclude that drought stress serves as a triggeronly, while the metabolic events lowering the osmotic potentialare favoured by high or intermediate water contents. Triticum durum L., durum wheat, rapid dehydration, osmotic adjustment, pressure-volume curves     

Ear of durum wheat under water stress: water relations and photosynthetic metabolism

The photosynthetic characteristics of the ear and flag leaf of well-watered (WW) and water-stressed (WS) durum wheat (Triticum turgidum L. var. durum) were studied in plants grown under greenhouse and Mediterranean field conditions. Gas exchange measurements simultaneously with modulated chlorophyll fluorescence were used to study the response of the ear and flag leaf to CO₂ and O₂ during photosynthesis. C₄ metabolism was identified by assessing the sensitivity of photosynthetic rate and electron transport to oxygen. The presence of CAM metabolism was assessed by measuring daily patterns of stomatal conductance and net CO₂ assimilation. In addition, the histological distribution of Rubisco protein in the ear parts was studied by immunocytochemical localisation. Relative water content (RWC) and osmotic adjustment (osmotic potential at full turgor) were also measured in these organs. Oxygen sensitivity of the assimilation rate and electron transport, the lack of Rubisco compartmentalisation in the mesophyll tissues and the gas-exchange pattern at night indicated that neither C₄ nor CAM metabolism occurs in the ear of WW or WS plants. Nevertheless, photosynthetic activity of the flag leaf was more affected by WS conditions than that of the ear, under both growing conditions. The lower sensitivity under water stress of the ear than of the flag leaf was linked to higher RWC and osmotic adjustment in the ear bracts and awns. We demonstrate that the better performance of the ear under water stress (compared to the flag leaf) is not related to C₄ or CAM photosynthesis. Rather, drought tolerance of the ear is explained by its higher RWC in drought. Osmotic adjustment and xeromorphic traits of ear parts may be responsible.