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1.  Role of adventitious roots in water relations of tamarack (Larix laricina) seedlings exposed to flooding 
BMC Plant Biology  2012;12:99.
Background
Flooding reduces supply of oxygen to the roots affecting plant water uptake. Some flooding-tolerant tree species including tamarack (Larix laricina (Du Roi) K. Koch) produce adventitious roots in response to flooding. These roots were reported to have higher hydraulic conductivity under flooding conditions compared with non-adventitious roots. In the present study, we examined structural and functional modifications in adventitious roots of tamarack seedlings to explain their flooding tolerance.
Results
Seedlings were subjected to the flooding treatment for six months, which resulted in an almost complete disintegration of the existing root system and its replacement with adventitious roots. We compared gas exchange parameters and water relations of flooded plants with the plants growing in well-drained soil and examined the root structures and root water transport properties. Although flooded seedlings had lower needle chlorophyll concentrations, their stomatal conductance, net photosynthesis rates and shoot water potentials were similar to non-flooded plants, indicative of flooding tolerance. Flooded adventitious roots had higher activation energy and a higher ratio of apoplastic to cell-to-cell water flow compared with non-flooded control roots as determined with the 1-hydroxypirene 3,6,8-trisulfonic acid apoplastic tracer dye. The adventitious roots in flooded plants also exhibited retarded xylem and endodermal development and accumulated numerous starch grains in the cortex. Microscopic examination of root sections treated with the PIP1 and PIP2 antibodies revealed high immunoreactivity in the cortex of non-flooded roots, as compared with flooded roots.
Conclusions
Structural modifications of adventitious roots suggest increased contribution of apoplastic bypass to water flow. The reduced dependence of roots on the hypoxia-sensitive aquaporin-mediated water transport is likely among the main mechanisms allowing tamarack seedlings to maintain water balance and gas exchange under flooding conditions.
doi:10.1186/1471-2229-12-99
PMCID: PMC3431261  PMID: 22738296
2.  Expression of ABA synthesis and metabolism genes under different irrigation strategies and atmospheric VPDs is associated with stomatal conductance in grapevine (Vitis vinifera L. cv Cabernet Sauvignon) 
Journal of Experimental Botany  2013;64(7):1907-1916.
The influence of different levels of irrigation and of variation in atmospheric vapour pressure deficit (VPD) on the synthesis, metabolism, and transport of abscisic acid (ABA) and the effects on stomatal conductance were examined in field-grown Cabernet Sauvignon grapevines. Xylem sap, leaf tissue, and root tissue were collected at regular intervals during two seasons in conjunction with measurements of leaf water potential (Ψleaf) and stomatal conductance (gs). The different irrigation levels significantly altered the Ψleaf and gs of the vines across both seasons. ABA abundance in the xylem sap was correlated with gs. The expression of genes associated with ABA synthesis, NCED1 and NCED2, was higher in the roots than in the leaves throughout and highest in the roots in mid January, a time when soil moisture declined and VPD was at its highest. Their expression in roots was also inversely related to the levels of irrigation and correlated with ABA abundance in the roots, xylem sap, and leaves. Three genes encoding ABA 8’-hydroxylases were isolated and their identities confirmed by expression in yeast cells. The expression of one of these, Hyd1, was elevated in leaves when VPD was below 2.0–2.5 kPa and minimal at higher VPD levels. The results provide evidence that ABA plays an important role in linking stomatal response to soil moisture status and that changes in ABA catabolism at or near its site of action allows optimization of gas exchange to current environmental conditions.
doi:10.1093/jxb/ert052
PMCID: PMC3638820  PMID: 23630325
Abscisic acid; ABA; ABA 8’-hydroxylase genes; grapevine; irrigation; NCED genes.
3.  Erythrina speciosa (Leguminosae-Papilionoideae) under soil water saturation: morphophysiological and growth responses 
Annals of Botany  2009;104(4):671-680.
Background and Aims
Erythrina speciosa is a Neotropical tree that grows mainly in moist habitats. To characterize the physiological, morphological and growth responses to soil water saturation, young plants of E. speciosa were subjected experimentally to soil flooding.
Methods
Flooding was imposed from 2 to 4 cm above the soil surface in water-filled tanks for 60 d. Non-flooded (control) plants were well watered, but never flooded. The net CO2 exchange (ACO2), stomatal conductance (gs) and intercellular CO2 concentration (Ci) were assessed for 60 d. Soluble sugar and free amino acid concentrations and the proportion of free amino acids were determined at 0, 7, 10, 21, 28 and 45 d of treatments. After 28, 45 and 60 d, dry masses of leaves, stems and roots were determined. Stem and root cross-sections were viewed using light microscopy.
Key Results
The ACO2 and gs were severely reduced by flooding treatment, but only for the first 10 d. The soluble sugars and free amino acids increased until the tenth day but decreased subsequently. The content of asparagine in the roots showed a drastic decrease while those of alanine and γ-aminobutyric increased sharply throughout the first 10 d after flooding. From the 20th day on, the flooded plants reached ACO2 and gs values similar to those observed for non-flooded plants. These events were coupled with the development of lenticels, adventitious roots and aerenchyma tissue of honeycomb type. Flooding reduced the growth rate and altered carbon allocation. The biomass allocated to the stem was higher and the root mass ratio was lower for flooded plants when compared with non-flooded plants.
Conclusions
Erythrina speciosa showed 100 % survival until the 60th day of flooding and was able to recover its metabolism. The recovery during soil flooding seems to be associated with morphological alterations, such as development of hypertrophic lenticels, adventitious roots and aerenchyma tissue, and with the maintenance of neutral amino acids in roots under long-term exposure to root-zone O2 deprivation.
doi:10.1093/aob/mcp159
PMCID: PMC2729630  PMID: 19581282
Erythrina speciosa; aerenchyma; amino acid content; biomass allocation; photosynthesis; flooding adaptations; stomatal conductance; O2 deficiency; γ-aminobutyric acid (GABA)
4.  Conditions Leading to High CO2 (>5 kPa) in Waterlogged–Flooded Soils and Possible Effects on Root Growth and Metabolism 
Annals of Botany  2006;98(1):9-32.
• Aims Soil waterlogging impedes gas exchange with the atmosphere, resulting in low PO2 and often high PCO2. Conditions conducive to development of high PCO2 (5–70 kPa) during soil waterlogging and flooding are discussed. The scant information on responses of roots to high PCO2 in terms of growth and metabolism is reviewed.
• Scope PCO2 at 15–70 kPa has been reported for flooded paddy-field soils; however, even 15 kPa PCO2 may not always be reached, e.g. when soil pH is above 7. Increases of PCO2 in soils following waterlogging will develop much more slowly than decreases in PO2; in soil from rice paddies in pots without plants, maxima in PCO2 were reached after 2–3 weeks. There are no reliable data on PCO2 in roots when in waterlogged or flooded soils. In rhizomes and internodes, PCO2 sometimes reached 10 kPa, inferring even higher partial pressures in the roots, as a CO2 diffusion gradient will exist from the roots to the rhizomes and shoots. Preliminary modelling predicts that when PCO2 is higher in a soil than in roots, PCO2 in the roots would remain well below the PCO2 in the soil, particularly when there is ventilation via a well-developed gas-space continuum from the roots to the atmosphere. The few available results on the effects of PCO2 at > 5 kPa on growth have nearly all involved sudden increases to 10–100 kPa PCO2; consequently, the results cannot be extrapolated with certainty to the much more gradual increases of PCO2 in waterlogged soils. Nevertheless, rice in an anaerobic nutrient solution was tolerant to 50 kPa CO2 being suddenly imposed. By contrast, PCO2 at 25 kPa retarded germination of some maize genotypes by 50 %. With regard to metabolism, assuming that the usual pH of the cytoplasm of 7·5 was maintained, every increase of 10 kPa CO2 would result in an increase of 75–90 mm HCO3− in the cytoplasm. pH maintenance would depend on the biochemical and biophysical pH stats (i.e. regulatory systems). Furthermore, there are indications that metabolism is adversely affected when HCO3− in the cytoplasm rises above 50 mm, or even lower; succinic dehydrogenase and cytochrome oxidase are inhibited by HCO3− as low as 10 mm. Such effects could be mitigated by a decrease in the set point for the pH of the cytoplasm, thus lowering levels of HCO3− at the prevailing PCO2 in the roots.
• Conclusions Measurements are needed on PCO2 in a range of soil types and in roots of diverse species, during waterlogging and flooding. Species well adapted to high PCO2 in the root zone, such as rice and other wetland plants, thrive even when PCO2 is well over 10 kPa; mechanisms of adaptation, or acclimatization, by these species need exploration.
doi:10.1093/aob/mcl076
PMCID: PMC3291891  PMID: 16644893
Acid load; aerenchyma; bicarbonate; carbon dioxide; cytochrome c; O2 deficiency; pH regulation; metabolism; respiration; waterlogging; wetland plants
5.  Expression of Tolerance for Meloidogyne graminicola in Rice Cultivars as Affected by Soil Type and Flooding 
Journal of Nematology  2000;32(3):309-317.
The effects of different water regimes on the pathogenicity of Meloidogyne graminicola on six rice cultivars were determined in two soil types in three greenhouse experiments. Two water regimes, simulating continuous flooding and intermittent flooding, were used with five of the cultivars. All cultivars were susceptible to the nematode, but IR72 and IR74 were more tolerant than IR20 and IR29 under intermittent flooding. All were tolerant under continuous flooding. UPLRi-5 was grown under multiple water regimes: no flooding; continuous flooding; flooding starting at maximum tillering, panicle initiation, or booting stage; and flooding from sowing until maximum tillering or booting. In sandy loam soil, M. graminicola reduced stem and leaf dry weight, root dry weight, and grain weight under all water regimes. In clay loam soil, the nematode reduced root weight when the soil was not flooded or flooded only for a short time, from panicle initiation, or booting to maturity, and from sowing to maximum tillering. In clay loam soil, stem and leaf dry weight, as well as grain weight, were reduced by the nematode under all water regimes except continuous flooding or when the soil was flooded from sowing to booting stage. These results indicate that rice cultivar tolerance of M. graminicola varies with water regime and that yield losses due to M. graminicola may be prevented or minimized when the rice crop is flooded early and kept flooded until a late stage of development.
PMCID: PMC2620461  PMID: 19270982
Meloidogyne graminicola; pathogenicity; rice; tolerance; water regime
6.  Measurement of Leaf Hydraulic Conductance and Stomatal Conductance and Their Responses to Irradiance and Dehydration Using the Evaporative Flux Method (EFM) 
Water is a key resource, and the plant water transport system sets limits on maximum growth and drought tolerance. When plants open their stomata to achieve a high stomatal conductance (gs) to capture CO2 for photosynthesis, water is lost by transpiration1,2. Water evaporating from the airspaces is replaced from cell walls, in turn drawing water from the xylem of leaf veins, in turn drawing from xylem in the stems and roots. As water is pulled through the system, it experiences hydraulic resistance, creating tension throughout the system and a low leaf water potential (Ψleaf). The leaf itself is a critical bottleneck in the whole plant system, accounting for on average 30% of the plant hydraulic resistance3. Leaf hydraulic conductance (Kleaf = 1/ leaf hydraulic resistance) is the ratio of the water flow rate to the water potential gradient across the leaf, and summarizes the behavior of a complex system: water moves through the petiole and through several orders of veins, exits into the bundle sheath and passes through or around mesophyll cells before evaporating into the airspace and being transpired from the stomata. Kleaf is of strong interest as an important physiological trait to compare species, quantifying the effectiveness of the leaf structure and physiology for water transport, and a key variable to investigate for its relationship to variation in structure (e.g., in leaf venation architecture) and its impacts on photosynthetic gas exchange. Further, Kleaf responds strongly to the internal and external leaf environment3. Kleaf can increase dramatically with irradiance apparently due to changes in the expression and activation of aquaporins, the proteins involved in water transport through membranes4, and Kleaf declines strongly during drought, due to cavitation and/or collapse of xylem conduits, and/or loss of permeability in the extra-xylem tissues due to mesophyll and bundle sheath cell shrinkage or aquaporin deactivation5-10. Because Kleaf can constrain gs and photosynthetic rate across species in well watered conditions and during drought, and thus limit whole-plant performance they may possibly determine species distributions especially as droughts increase in frequency and severity11-14.
We present a simple method for simultaneous determination of Kleaf and gs on excised leaves. A transpiring leaf is connected by its petiole to tubing running to a water source on a balance. The loss of water from the balance is recorded to calculate the flow rate through the leaf. When steady state transpiration (E, mmol • m-2 • s-1) is reached, gs is determined by dividing by vapor pressure deficit, and Kleaf by dividing by the water potential driving force determined using a pressure chamber (Kleaf= E /- Δψleaf, MPa)15.
This method can be used to assess Kleaf responses to different irradiances and the vulnerability of Kleaf to dehydration14,16,17.
doi:10.3791/4179
PMCID: PMC3577864  PMID: 23299126
Plant Biology; Issue 70; Molecular Biology; Physiology; Ecology; Biology; Botany; Leaf traits; hydraulics; stomata; transpiration; xylem; conductance; leaf hydraulic conductance; resistance; evaporative flux method; whole plant
7.  Root attributes affecting water uptake of rice (Oryza sativa) under drought 
Journal of Experimental Botany  2012;63(13):4751-4763.
Lowland rice roots have a unique physiological response to drought because of their adaptation to flooded soil. Rice root attributes that facilitate growth under flooded conditions may affect rice response to drought, but the relative roles of root structural and functional characteristics for water uptake under drought in rice are not known. Morphological, anatomical, biochemical, and molecular attributes of soil-grown rice roots were measured to investigate the genotypic variability and genotype×environment interactions of water uptake under variable soil water regimes. Drought-resistant genotypes had the lowest night-time bleeding rates of sap from the root system in the field. Diurnal fluctuation predominated as the strongest source of variation for bleeding rates in the field and root hydraulic conductivity (Lp r) in the greenhouse, and was related to expression trends of various PIP and TIP aquaporins. Root anatomy was generally more responsive to drought treatments in drought-resistant genotypes. Suberization and compaction of sclerenchyma layer cells decreased under drought, whereas suberization of the endodermis increased, suggesting differential roles of these two cell layers for the retention of oxygen under flooded conditions (sclerenchyma layer) and retention of water under drought (endodermis). The results of this study point to the genetic variability in responsiveness to drought of rice roots in terms of morphology, anatomy, and function.
doi:10.1093/jxb/ers150
PMCID: PMC3427995  PMID: 22791828
Aquaporin; drought; rice; root anatomy; root hydraulic conductivity; suberin
8.  Hydraulic Lift Increases Herbivory by Diaprepes abbreviatus Larvae and Persistence of Steinernema riobrave in Dry Soil 
Journal of Nematology  2001;33(2-3):142-146.
Citrus seedlings were grown in double pots that separated the root systems into discrete lower and upper zones to test the hypothesis that hydraulic lift affects persistence and efficacy of entomopathogenic nematodes. Three treatments were established: (i) both pots were irrigated at water potential ≤ -15 kPa (no drought); (ii) only the bottom pot was irrigated (partial drought); (iii) neither pot was irrigated (complete drought). Steinernema riobrave infective juveniles (IJ) were added to the soil in the top pots of all treatments. During 27 days, the water potential in soil in the top pots of both the partial and complete drought treatments declined to ca. -160 kPa. A greater number of nematodes (P ≤ 0.01) persisted in soil as motile IJ under conditions of partial drought (143/pot) than under no drought (6.1/pot) or complete drought (4.4/pot). A second experiment was initiated with the same treatments as the first, except that only half of the 20 replicates in each moisture regime were inoculated with nematodes. After 15 days, all top pots were irrigated and two larvae of the insect Diaprepes abbreviatus were added to all of the top pots in each treatment. Irrigation regimes were reinstituted until water potential in the top pots under partial and complete drought had again declined to ca. -150 kPa and the experiment was terminated. In the absence of nematodes, the damage to tap roots caused by D. abbreviatus feeding under partial drought and complete drought was 80% and 32%, respectively, of that under no drought. Numbers of motile IJ in soil were greater under conditions of partial drought (736/pot) than under complete drought (2.0/pot) or no drought (7.2/pot). Survival of D. abbreviatus and insect damage to roots were reduced by the presence of S. riobrave to a greater extent under partial drought as compared to other treatments. Hydraulic lift from the lower to the upper rhizosphere appears to have modulated the effect of dry soil conditions on feeding behavior of D. abbreviatus and created favorable conditions for persistence and efficacy of the entomopathogenic nematode.
PMCID: PMC2638139  PMID: 19266011
biological control; citrus; Diaprepes abbreviatus; drought; hydraulic lift; soil moisture; Steinernema riobrave; water potential
9.  Rapid and long-term effects of water deficit on gas exchange and hydraulic conductance of silver birch trees grown under varying atmospheric humidity 
BMC Plant Biology  2014;14:72.
Background
Effects of water deficit on plant water status, gas exchange and hydraulic conductance were investigated in Betula pendula under artificially manipulated air humidity in Eastern Estonia. The study was aimed to broaden an understanding of the ability of trees to acclimate with the increasing atmospheric humidity predicted for northern Europe. Rapidly-induced water deficit was imposed by dehydrating cut branches in open-air conditions; long-term water deficit was generated by seasonal drought.
Results
The rapid water deficit quantified by leaf (ΨL) and branch water potentials (ΨB) had a significant (P < 0.001) effect on gas exchange parameters, while inclusion of ΨB in models resulted in a considerably better fit than those including ΨL, which supports the idea that stomatal openness is regulated to prevent stem rather than leaf xylem dysfunction. Under moderate water deficit (ΨL≥-1.55 MPa), leaf conductance to water vapour (gL), transpiration rate and leaf hydraulic conductance (KL) were higher (P < 0.05) and leaf temperature lower in trees grown in elevated air humidity (H treatment) than in control trees (C treatment). Under severe water deficit (ΨL<-1.55 MPa), the treatments showed no difference. The humidification manipulation influenced most of the studied characteristics, while the effect was to a great extent realized through changes in soil water availability, i.e. due to higher soil water potential in H treatment. Two functional characteristics (gL, KL) exhibited higher (P < 0.05) sensitivity to water deficit in trees grown under increased air humidity.
Conclusions
The experiment supported the hypothesis that physiological traits in trees acclimated to higher air humidity exhibit higher sensitivity to rapid water deficit with respect to two characteristics - leaf conductance to water vapour and leaf hydraulic conductance. Disproportionate changes in sensitivity of stomatal versus leaf hydraulic conductance to water deficit will impose greater risk of desiccation-induced hydraulic dysfunction on the plants, grown under high atmospheric humidity, in case of sudden weather fluctuations, and might represent a potential threat in hemiboreal forest ecosystems. There is no trade-off between plant hydraulic capacity and photosynthetic water-use efficiency on short time scale.
doi:10.1186/1471-2229-14-72
PMCID: PMC3976162  PMID: 24655599
Betula pendula; Branch water potential; Climate change; Hydraulic conductance; Leaf water potential; Net photosynthesis; Silver birch; Stomatal conductance; Water-use efficiency
10.  Aerobic rice genotypes displayed greater adaptation to water-limited cultivation and tolerance to polyethyleneglycol-6000 induced stress 
Water scarcity and drought have seriously threatened traditional rice cultivation practices in several parts of the world including India. In the present investigation, experiments were conducted to see if the water-efficient aerobic rice genotypes developed at UAS, Bangalore (MAS25, MAS26 and MAS109) and IRRI, Philippines (MASARB25 and MASARB868), are endowed with drought tolerance or not. A set of these aerobic and five lowland high-yielding (HKR47 and PAU201, Taraori Basmati, Pusa1121 and Pusa1460) indica rice genotypes were evaluated for: (i) yield and yield components under submerged and aerobic conditions in field, (ii) root morphology and biomass under aerobic conditions in pots in the nethouse, (iii) PEG-6000 (0, −1, −2 and −3 bar) induced drought stress at vegetative stage using a hydroponic culture system and (iv) polymorphism for three SSR markers associated with drought resistance traits. Under submerged conditions, the yield of aerobic rice genotypes declined by 13.4–20.1 % whereas under aerobic conditions the yield of lowland indica/Basmati rice varieties declined by 23–27 %. Under water-limited conditions in pots, aerobic rice genotypes had 54–73.8 % greater root length and 18–60 % higher fresh root biomass compared to lowland indica rice varieties. Notably, root length of MASARB25 was 35 % shorter than MAS25 whereas fresh and dry root biomass of MASARB25 was 10 % and 64 % greater than MAS25. The lowland indica were more sensitive to PEG-stress with a score of 5.9–7.6 for Basmati and 6.1–6.7 for non-aromatic indica rice varieties, than the aerobic rice genotypes (score 2.7–3.3). A set of three microsatellite DNA markers (RM212, RM302 and RM3825) located on chromosome 1 which has been shown to be associated with drought resistance was investigated in the present study. Two of these markers (RM212 and RM302) amplified a specific allele in all the aerobic rice genotypes which were absent in lowland indica rice genotypes.
doi:10.1007/s12298-011-0094-2
PMCID: PMC3550527  PMID: 23573038
Aerobic; Drought; Indica; Polyethyleneglycol 6000; Root traits; SSR
11.  Association of water spectral indices with plant and soil water relations in contrasting wheat genotypes 
Journal of Experimental Botany  2010;61(12):3291-3303.
Spectral reflectance indices can be used to estimate the water status of plants in a rapid, non-destructive manner. Water spectral indices were measured on wheat under a range of water-deficit conditions in field-based yield trials to establish their relationship with water relations parameters as well as available volumetric soil water (AVSW) to indicate soil water extraction patterns. Three types of wheat germplasm were studied which showed a range of drought adaptation; near-isomorphic sister lines from an elite/elite cross, advanced breeding lines, and lines derived from interspecific hybridization with wild relatives (synthetic derivative lines). Five water spectral indices (one water index and four normalized water indices) based on near infrared wavelengths were determined under field conditions between the booting and grain-filling stages of crop development. Among all water spectral indices, one in particular, which was denominated as NWI-3, showed the most consistent associations with water relations parameters and demonstrated the strongest associations in all three germplasm sets. NWI-3 showed a strong linear relationship (r2 >0.6–0.8) with leaf water potential (ψleaf) across a broad range of values (–2.0 to –4.0 MPa) that were determined by natural variation in the environment associated with intra- and inter-seasonal affects. Association observed between NWI-3 and canopy temperature (CT) was consistent with the idea that genotypes with a better hydration status have a larger water flux (increased stomatal conductance) during the day. NWI-3 was also related to soil water potential (ψsoil) and AVSW, indicating that drought-adapted lines could extract more water from deeper soil profiles to maintain favourable water relations. NWI-3 was sufficiently sensitive to detect genotypic differences (indicated by phenotypic and genetic correlations) in water status at the canopy and soil levels indicating its potential application in precision phenotyping.
doi:10.1093/jxb/erq156
PMCID: PMC2905199  PMID: 20639342
Canopy reflectance; canopy water content; leaf water potential; root growth; water index
12.  Risk-taking plants 
Plant Signaling & Behavior  2012;7(7):767-770.
Water scarcity is a critical limitation for agricultural systems. Two different water management strategies have evolved in plants: an isohydric strategy and an anisohydric strategy. Isohydric plants maintain a constant midday leaf water potential (Ψleaf) when water is abundant, as well as under drought conditions, by reducing stomatal conductance as necessary to limit transpiration. Anisohydric plants have more variable Ψleaf and keep their stomata open and photosynthetic rates high for longer periods, even in the presence of decreasing leaf water potential. This risk-taking behavior of anisohydric plants might be beneficial when water is abundant, as well as under moderately stressful conditions. However, under conditions of intense drought, this behavior might endanger the plant. We will discuss the advantages and disadvantages of these two water-usage strategies and their effects on the plant’s ability to tolerate abiotic and biotic stress. The involvement of plant tonoplast AQPs in this process will also be discussed.
doi:10.4161/psb.20505
PMCID: PMC3583960  PMID: 22751307
abiotic stress; anisohydric; aquaporins; biotic stress; isohydric; leaf water potential; relative water content
13.  Effect of Depth of Flooding on the Rice Water Weevil, Lissorhoptrus oryzophilus, and Yield of Rice 
The rice water weevil, Lissorhoptrus oryzophilus (Kuschel) (Coleoptera: Curculionidae), is a semi-aquatic pest of rice and is the most destructive insect pest of rice in the United States. Adults oviposit after floods are established, and greenhouse studies have shown that plants exposed to deep floods have more eggs oviposited in leaf sheaths than plants exposed to a shallow flood. Experiments were conducted in three mid-southern states in the USA to determine if the depth of flooding would impact numbers of L. oryzophilus on rice plants under field conditions. Rice was flooded at depths of approximately 5 or 10 cm in Arkansas in 2007 and 2008 and Louisiana in 2008, and at depths between 0–20 cm in Missouri in 2008. Plants were sampled three and four weeks after floods were established in all locations, and also two weeks after flood in Missouri. On all sampling dates in four experiments over two years and at three field sites, fewer L. oryzophilus larvae were collected from rice in shallow-flooded plots than from deep-flooded plots. The number of L. oryzophilus was reduced by as much as 27% in shallow-flooded plots. However, the reduction in insect numbers did not translate to a significant increase in rice yield. We discuss how shallow floods could be used as a component of an integrated pest management program for L. oryzophilus.
doi:10.1673/031.013.6201
PMCID: PMC3740922  PMID: 23906324
cultural control; Oryza sativa; rice water weevil management
14.  The effects of water regime on phosphorus responses of rainfed lowland rice cultivars 
Annals of Botany  2008;103(2):211-220.
Background and Aims
Soil phosphorus (P) solubility declines sharply when a flooded soil drains, and an important component of rice (Oryza sativa) adaptation to rainfed lowland environments is the ability to absorb and utilize P under such conditions. The aim of this study was to test the hypothesis that rice cultivars differ in their P responses between water regimes because P uptake mechanisms differ.
Methods
Six lowland rice cultivars (three considered tolerant of low P soils, three sensitive) were grown in a factorial experiment with three water regimes (flooded, moist and flooded-then-moist) and four soil P levels, and growth and P uptake were measured. Small volumes of soil were used to maximize inter-root competition and uptake per unit root surface. The results were compared with the predictions of a model allowing for the effects of water regime on P solubility and diffusion.
Key Results
The plants were P stressed but not water stressed in all the water regimes at all P levels except the higher P additions in the flooded soil. The cultivar rankings scarcely differed between the water regimes and P additions. In all the treatments, the soil P concentrations required to explain the measured uptake were several times the concentration of freely available P in the soil.
Conclusions
The cultivar rankings were driven more by differences in growth habit than specific P uptake mechanisms, so the hypothesis cannot be corroborated with these data. Evidently all the plants could tap sparingly soluble forms of P by releasing a solubilizing agent or producing a greater root length than measured, or both. However, any cultivar differences in this were not apparent in greater net P uptake, possibly because the restricted rooting volume meant that additional P uptake could not be converted into new root growth to explore new soil volumes.
doi:10.1093/aob/mcn199
PMCID: PMC2707314  PMID: 18945744
Oryza sativa; rainfed lowland; phosphorus efficiency; root morphology; solubilization; rice cultivar
15.  Rice Performance and Water Use Efficiency under Plastic Mulching with Drip Irrigation 
PLoS ONE  2013;8(12):e83103.
Plastic mulching with drip irrigation is a new water-saving rice cultivation technology, but little is known on its productivity and water-saving capacity. This study aimed to assess the production potential, performance, and water use efficiency (WUE) of rice under plastic mulching with drip irrigation. Field experiments were conducted over 2 years with two rice cultivars under different cultivation systems: conventional flooding (CF), non-flooded irrigation incorporating plastic mulching with furrow irrigation (FIM), non-mulching with furrow irrigation (FIN), and plastic mulching with drip irrigation (DI). Compared with the CF treatment, grain yields were reduced by 31.76–52.19% under the DI treatment, by 57.16–61.02% under the FIM treatment, by 74.40–75.73% under the FIN treatment, which were mainly from source limitation, especially a low dry matter accumulation during post-anthesis, in non-flooded irrigation. WUE was the highest in the DI treatment, being 1.52–2.12 times higher than with the CF treatment, 1.35–1.89 times higher than with the FIM treatment, and 2.37–3.78 times higher than with the FIN treatment. The yield contribution from tillers (YCFTs) was 50.65–62.47% for the CF treatment and 12.07–20.62% for the non-flooded irrigation treatments. These low YCFTs values were attributed to the poor performance in tiller panicles rather than the total tiller number. Under non-flooded irrigation, root length was significantly reduced with more roots distributed in deep soil layers compared with the CF treatment; the DI treatment had more roots in the topsoil layer than the FIM and FIN treatments. The experiment demonstrates that the DI treatment has greater water saving capacity and lower yield and economic benefit gaps than the FIM and FIN treatments compared with the CF treatment, and would therefore be a better water-saving technology in areas of water scarcity.
doi:10.1371/journal.pone.0083103
PMCID: PMC3858349  PMID: 24340087
16.  Waterlogging-induced changes in root architecture of germplasm accessions of the tropical forage grass Brachiaria humidicola 
AoB Plants  2014;6:plu017.
Brachiaria humidicola, a tropical forage grass, develops aerenchyma in nodal roots to adapt to waterlogging. A large body of work has focused on the functional role of aerenchyma in nodal roots under waterlogged soil conditions. On the other hand, quantification of responses of lateral roots to waterlogging has been often overlooked in past work. Our data indicated that although waterlogging reduced the overall proportion of lateral roots, its proportion significantly increased in the top 10 cm of the soil. This suggests that soil flooding increases lateral root proliferation of B. humidicola in upper soil layers. This may compensate the reduction of root surface area brought by the restriction of root growth at depths below 30 cm into waterlogged soil.
Waterlogging is one of the major factors limiting the productivity of pastures in the humid tropics. Brachiaria humidicola is a forage grass commonly used in zones prone to temporary waterlogging. Brachiaria humidicola accessions adapt to waterlogging by increasing aerenchyma in nodal roots above constitutive levels to improve oxygenation of root tissues. In some accessions, waterlogging reduces the number of lateral roots developed from main root axes. Waterlogging-induced reduction of lateral roots could be of adaptive value as lateral roots consume oxygen supplied from above ground via their parent root. However, a reduction in lateral root development could also be detrimental by decreasing the surface area for nutrient and water absorption. To examine the impact of waterlogging on lateral root development, an outdoor study was conducted to test differences in vertical root distribution (in terms of dry mass and length) and the proportion of lateral roots to the total root system (sum of nodal and lateral roots) down the soil profile under drained or waterlogged soil conditions. Plant material consisted of 12 B. humidicola accessions from the gene bank of the International Center for Tropical Agriculture, Colombia. Rooting depth was restricted by 21 days of waterlogging and confined to the first 30 cm below the soil surface. Although waterlogging reduced the overall proportion of lateral roots, its proportion significantly increased in the top 10 cm of the soil. This suggests that soil flooding increases lateral root proliferation of B. humidicola in the upper soil layers. This may compensate for the reduction of root surface area brought about by the restriction of root growth at depths below 30 cm. Further work is needed to test the relative efficiency of nodal and lateral roots for nutrient and water uptake under waterlogged soil conditions.
doi:10.1093/aobpla/plu017
PMCID: PMC4038435  PMID: 24876299
Lateral root proportion; oxygen deficiency; rooting depth; root length; soil flooding; vertical root distribution.
17.  Population Changes of Tylenchulus semipenetrans Under Localized Versus Uniform Drought in the Citrus Root Zone 
Journal of Nematology  1996;28(3):360-368.
Population development of Tylenchulus semipenetrans in dry soil was investigated in a greenhouse study. Citrus seedlings were grown in sandy soil in vertical tubes with upper and lower sections. Nematode population densities in the upper tubes were measured at 16, 23, and 37 days, post-treatment. Three treatments consisted of i) irrigating both tubes when soil water potential reached -1 5 kPa (non-drought), ii) irrigating only the bottom tube (local drought), and iii) no irrigation (uniform drought). Soil water potential in the upper tubes did not differ under local and uniform drought during the first 16 days post-treatment, when it approached - 125 kPa. Thereafter, the water potential of soil under uniform drought continued to decrease, while that under local drought stabilized at approximately -150 kPa. Treatments had no consistent effects on female T. semipenetrans counts from soil or roots. However, after 37 days, numbers of eggs, juvenile, and male nematodes per gram of root under local drought were more than 2.4-fold greater than those under non-drought or uniform drought. Numbers of juvenile and male nematodes in soil were 6.5 times higher under local drought than under non-drought after 37 days. Nematodes did not survive in soil under uniform drought. Most of the eggs recovered on each date, from roots under local and non-drought, hatched within 35 days. Sixteen days of uniform drought reduced cumulative egg hatch to 51%, and almost no eggs hatched after 23 and 37 days of uniform drought. Thus, the response of T. semipenetrans to dry soil is fundamentally different, depending on whether all or part of the rhizosphere experiences drought. These data and field observations suggest that hydraulic lift via the root xylem may prolong the activity of some nematodes and possibly other rhizosphere-inhabiting organisms in dry soil.
PMCID: PMC2619702  PMID: 19277154
carbohydrate; citrus; citrus nematode; ecology; drought; hydraulic lift; soil moisture; survival; Tylenchulus semipenetrans
18.  Physiological and molecular responses to drought in Petunia: the importance of stress severity 
Journal of Experimental Botany  2012;63(18):6335-6345.
Plant responses to drought stress vary depending on the severity of stress and the stage of drought progression. To improve the understanding of such responses, the leaf physiology, abscisic acid (ABA) concentration, and expression of genes associated with ABA metabolism and signalling were investigated in Petunia × hybrida. Plants were exposed to different specific substrate water contents (θ = 0.10, 0.20, 0.30, or 0.40 m3·m–3) to induce varying levels of drought stress. Plant responses were investigated both during the drying period (θ decreased to the θ thresholds) and while those threshold θ were maintained. Stomatal conductance (gs) and net photosynthesis (A) decreased with decreasing midday leaf water potential (Ψleaf). Leaf ABA concentration increased with decreasing midday Ψleaf and was negatively correlated with gs (r = –0.92). Despite the increase in leaf ABA concentration under drought, no significant effects on the expression of ABA biosynthesis genes were observed. However, the ABA catabolism-related gene CYP707A2 was downregulated, primarily in plants under severe drought (θ = 0.10 m3∙m–3), suggesting a decrease in ABA catabolism under severe drought. Expression of phospholipase Dα (PLDα), involved in regulating stomatal responses to ABA, was enhanced under drought during the drying phase, but there was no relationship between PLDα expression and midday Ψleaf after the θ thresholds had been reached. The results show that drought response of plants depends on the severity of drought stress and the phase of drought progression.
doi:10.1093/jxb/ers285
PMCID: PMC3504489  PMID: 23077204
abscisic acid; acclimation; automated irrigation; soil moisture sensor; stomatal conductance; substrate water content
19.  Contrasting physiological effects of partial root zone drying in field-grown grapevine (Vitis vinifera L. cv. Monastrell) according to total soil water availability 
Journal of Experimental Botany  2012;63(11):4071-4083.
Different spatial distributions of soil moisture were imposed on field-grown grapevines by applying the same irrigation volumes to the entire (DI; deficit irrigation) or part of the (PRD; partial root zone drying) root zone. Five treatments were applied: controls irrigated at 60% ETc (crop evapotranspiration) for the whole season (308 mm year−1); DI-1 and PRD-1 that received the same irrigation as controls before fruit set, 30% ETc from fruit set to harvest and 45% ETc post-harvest (192 mm year−1); and DI-2 and PRD-2 that were the same, except that 15% ETc was applied from fruit set to harvest (142 mm year−1). Compared with DI-1, PRD-1 maintained higher leaf area post-veraison and increased root water uptake, whole-plant hydraulic conductance, leaf transpiration, stomatal conductance, and photosynthesis, but decreased intrinsic gas exchange efficiency without causing differences in leaf xylem abscisic acid (ABA) concentration. Compared with DI-2, PRD-2 increased leaf xylem ABA concentration and decreased root water uptake, whole-plant hydraulic conductance, leaf transpiration, stomatal conductance, and photosynthesis, mainly at the beginning of PRD cycles. Distinctive PRD effects (e.g. greater stomatal closure) depended on the volumetric soil water content of the wet root zone, as predicted from a model of root-to-shoot ABA signalling.
doi:10.1093/jxb/ers088
PMCID: PMC3398444  PMID: 22451721
ABA signalling; deficit irrigation; leaf water relations; partial root zone drying; root water uptake; soil water content
20.  Cropping practices, soil properties, pedotransfer functions and organic carbon storage at Kuanria canal command area in India 
SpringerPlus  2013;2:631.
Effects of cropping practices on soil properties viz. particle size distribution, pH, bulk density (BD), field capacity (FC, -33 kPa), permanent wilting point (PWP, -1500 kPa), available water capacity (AWC) and soil organic carbon (SOC) were assessed. The pedotransfer functions (PTFs) were developed for saturated hydraulic conductivity (Ks), water retention at FC and PWP of soils for different sites under major cropping system in a canal irrigated area. The results revealed that the soils are mainly composed of sand and clay with the clay contents ranging from 29.6 to 48.8%, BD of 1.44-1.72 Mg m-3, and 0.34 to 0.95% SOC. The Ks decreased, and water retention at FC, PWP and AWC increased significantly with soil depth due to greater clay contents in lower soil depths. The PTFs were best represented as the power functions for prediction of Ks with clay content as predictor variable; whereas the PTFs for water retention at FC and PWP were better represented as the exponential functions. SOC content was higher under rice-sugarcane crop rotation compared to other systems. SOC storage in the surface layer was higher in rice-sugarcane rotation (18.90-20.53 Mg ha-1) than other sites. The developed PTFs would be very useful in soil and water management strategies for the study area or elsewhere having similar soil and cropping practices. The information on SOC storage in the Kuanria region would help for better soil and crop planning in future.
doi:10.1186/2193-1801-2-631
PMCID: PMC3843505  PMID: 24303340
Cropping; Soil properties; Pedotransfer functions; SOC storage
21.  Do the Rich Always Become Richer? Characterizing the Leaf Physiological Response of the High-Yielding Rice Cultivar Takanari to Free-Air CO2 Enrichment 
Plant and Cell Physiology  2014;55(2):381-391.
The development of crops which are well suited to growth under future environmental conditions such as higher atmospheric CO2 concentrations ([CO2]) is essential to meeting the challenge of ensuring food security in the face of the growing human population and changing climate. A high-yielding indica rice variety (Oryza sativa L. cv. Takanari) has been recently identified as a potential candidate for such breeding, due to its high productivity in present [CO2]. To test if it could further increase its productivity under elevated [CO2] (eCO2), Takanari was grown in the paddy field under season-long free-air CO2 enrichment (FACE, approximately 200 µmol mol−1 above ambient [CO2]) and its leaf physiology was compared with the representative japonica variety ‘Koshihikari’. Takanari showed consistently higher midday photosynthesis and stomatal conductance than Koshihikari under both ambient and FACE growth conditions over 2 years. Maximum ribulose-1,5-bisphosphate carboxylation and electron transport rates were higher for Takanari at the mid-grain filling stage in both years. Mesophyll conductance was higher in Takanari than in Koshihikari at the late grain-filling stage. In contrast to Koshihikari, Takanari grown under FACE conditions showed no decrease in total leaf nitrogen on an area basis relative to ambient-grown plants. Chl content was higher in Takanari than in Koshihikari at the same leaf nitrogen level. These results indicate that Takanari maintains its superiority over Koshihikari in regards to its leaf-level productivity when grown in elevated [CO2] and it may be a valuable resource for rice breeding programs which seek to increase crop productivity under current and future [CO2].
doi:10.1093/pcp/pcu009
PMCID: PMC3913450  PMID: 24443497
Ecophysiology; FACE; Global change; Photosynthesis
22.  Does night-time transpiration contribute to anisohydric behaviour in a Vitis vinifera cultivar? 
Journal of Experimental Botany  2009;60(13):3751-3763.
The hypothesis that vines of the Semillon wine grape variety show anisohydric behaviour was tested, i.e. that tissue hydration is unstable under fluctuating environmental conditions. Stomatal conductance and transpiration rates from leaves were measured during the day and at night. Leaf water potential (Ψl) in Semillon was negatively correlated to vapour pressure deficit (VPD) both predawn and during the day. Furthermore, Ψl fell to significantly lower values than in any of the nine other varieties examined. Night-time values of stomatal conductance (gn) and transpiration (En) in Semillon were up to four times higher than in other varieties; plants enclosed in plastic bags overnight to reduce En resulted in better plant–soil equilibration so that predawn Ψl in Semillon was the same as in Grenache. These data indicate that the hypothesis is supported, and that night-time transpiration contributes significantly to the low Ψl values in Semillon during warm, dry nights. The other contributing factor is daytime stomatal conductance (gday), which in Semillon leaves was higher than in other varieties, although the decline in gday with increasing VPD was greater in Semillon than in Shiraz or Grenache. The high values of gday were associated with high rates of transpiration (Eday) by Semillon through a day when VPD reached 4.5 kPa. When compared to other varieties, Semillon was not unusual in terms of root length density, stomatal density, xylem sap abscisic acid, or leaf electrolyte leakage. Night-time and daytime water loss and insufficient stomatal regulation therefore account for the tendency to anisohydric behaviour shown by Semillon.
doi:10.1093/jxb/erp217
PMCID: PMC2736890  PMID: 19584116
Drought responses; evaporative demand; Semillon; stomatal regulation; water relations
23.  Zn uptake, translocation and grain Zn loading in rice (Oryza sativa L.) genotypes selected for Zn deficiency tolerance and high grain Zn 
Journal of Experimental Botany  2013;64(10):2739-2751.
Zn deficiency is a widespread problem in rice (Oryza sativa L.) grown under flooded conditions, limiting growth and grain Zn accumulation. Genotypes with Zn deficiency tolerance or high grain Zn have been identified in breeding programmes, but little is known about the physiological mechanisms conferring these traits. A protocol was developed for growing rice to maturity in agar nutrient solution (ANS), with optimum Zn-sufficient growth achieved at 1.5 μM ZnSO4.7H2O. The redox potential in ANS showed a decrease from +350 mV to −200 mV, mimicking the reduced conditions of flooded paddy soils. In subsequent experiments, rice genotypes contrasting for Zn deficiency tolerance and grain Zn were grown in ANS with sufficient and deficient Zn to assess differences in root uptake of Zn, root-to-shoot Zn translocation, and in the predominant sources of Zn accumulation in the grain. Zn efficiency of a genotype was highly influenced by root-to-shoot translocation of Zn and total Zn uptake. Translocation of Zn from root to shoot was more limiting at later growth stages than at the vegetative stage. Under Zn-sufficient conditions, continued root uptake during the grain-filling stage was the predominant source of grain Zn loading in rice, whereas, under Zn-deficient conditions, some genotypes demonstrated remobilization of Zn from shoot and root to grain in addition to root uptake. Understanding the mechanisms of grain Zn loading in rice is crucial in selecting high grain Zn donors for target-specific breeding and also to establish fertilizer and water management strategies for achieving high grain Zn.
doi:10.1093/jxb/ert118
PMCID: PMC3697949  PMID: 23698631
Agar nutrient solution; grain Zn; grain Zn loading; remobilization; rice; Zn deficiency tolerance; Zn efficiency; Zn translocation; Zn uptake.
24.  Root signals and stomatal closure in relation to photosynthesis, chlorophyll a fluorescence and adventitious rooting of flooded tomato plants 
Annals of Botany  2008;103(2):313-323.
Background and Aims
An investigation was carried out to determine whether stomatal closure in flooded tomato plants (Solanum lycopersicum) results from decreased leaf water potentials (ψL), decreased photosynthetic capacity and attendant increases in internal CO2 (Ci) or from losses of root function such as cytokinin and gibberellin export.
Methods
Pot-grown plants were flooded when 1 month old. Leaf conductance was measured by diffusion porometry, the efficiency of photosystem II (PSII) was estimated by fluorimetry, and infrared gas analysis was used to determine Ci and related parameters.
Key Results
Flooding starting in the morning closed the stomata and increased ψL after a short-lived depression of ψL. The pattern of closure remained unchanged when ψ`L depression was avoided by starting flooding at the end rather than at the start of the photoperiod. Raising external CO2 concentrations by 100 µmol mol−1 also closed stomata rapidly. Five chlorophyll fluorescence parameters [Fq′/Fm′, Fq′/Fv′, Fv′/Fm′, non-photochemical quenching (NPQ) and Fv/Fm] were affected by flooding within 12–36 h and changes were linked to decreased Ci. Closing stomata by applying abscisic acid or increasing external CO2 substantially reproduced the effects of flooding on chlorophyll fluorescence. The presence of well-aerated adventitious roots partially inhibited stomatal closure of flooded plants. Allowing adventitious roots to form on plants flooded for >3 d promoted some stomatal re-opening. This effect of adventitious roots was not reproduced by foliar applications of benzyl adenine and gibberellic acid.
Conclusions
Stomata of flooded plants did not close in response to short-lived decreases in ψL or to increased Ci resulting from impaired PSII photochemistry. Instead, stomatal closure depressed Ci and this in turn largely explained subsequent changes in chlorophyll fluorescence parameters. Stomatal opening was promoted by the presence of well-aerated adventitious roots, implying that loss of function of root signalling contributes to closing of stomata during flooding. The possibility that this involves inhibition of cytokinin or gibberellin export was not well supported.
doi:10.1093/aob/mcn208
PMCID: PMC2707317  PMID: 19001430
Root to shoot communication; flooding stress; stomatal closure; photosynthesis; chlorophyll fluorescence; gas exchange; adventitious roots; plant hormones; abscisic acid; cytokinins; gibberellic acid
25.  Mechanisms associated with tolerance to flooding during germination and early seedling growth in rice (Oryza sativa) 
Annals of Botany  2008;103(2):197-209.
Background and Aims
Flooding slows seed germination, imposes fatalities and delays seedling establishment in direct-seeded rice. This study describes responses of contrasting rice genotypes subjected to flooding or low oxygen stress during germination and discusses the basis of tolerance shown by certain cultivars.
Methods
In one set of experiments, dry seeds were sown in soil and either watered normally or flooded with 10 cm of water. Seedling survival and shoot and root growth were assessed and seed portions of germinating seedlings were assayed for soluble sugars and starch concentrations. The whole germinating seedlings were assayed for amylase and peroxidase activities and for ethylene production. Activities of enzymes associated with anaerobic respiration were examined and gene expression was analysed separately with seeds germinating under different amounts of dissolved oxygen in dilute agar.
Key Results
Flooding during germination reduced survival but to a lesser extent in tolerant genotypes. Starch concentration in germinating seeds decreased while sugar concentration increased under flooding, but more so in tolerant genotypes. Amylase activity correlated positively with elongation (r = 0·85 for shoot and 0·83 for root length) and with plant survival (r = 0·92). Tolerant genotypes had higher amylase activity and higher RAmy3D gene expression. Ethylene was not detected in seeds within 2 d after sowing, but increased thereafter, with a greater increase in tolerant genotypes starting 3 d after sowing. Peroxidase activity was higher in germinating seeds of sensitive genotypes and correlated negatively with survival.
Conclusions
Under low oxygen stress, tolerant genotypes germinate, grow faster and more seedlings survive. They maintain their ability to use stored starch reserves through higher amylase activity and anaerobic respiration, have higher rates of ethylene production and lower peroxidase activity as germinating seeds and as seedlings. Relevance of these traits to tolerance of flooding during germination and early growth is discussed.
doi:10.1093/aob/mcn211
PMCID: PMC2707318  PMID: 19001425
Amylase; anoxia; crop establishment; direct-seeded rice; ethylene; flooding; germination; hypoxia; Oryza sativa

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