The influence of redox conditions in soils on extraplasmatic Fe-loading of plant roots

Loading of extraplasmatic Fe, as a potential storage pool for Fe nutrition, was studied in roots of maize grown under hypoxic conditions in soil culture. Extraplasmatic Fe loading was investigated depending on (i) duration of flooding (0, 1, 2 or 4 days) and (ii) microbial activity as affected by graduated addition of carbon sources (0, 2 or 10 g each starch and cellulose kg^?1 soil). Maize plants were grown in a soil culture system with root systems enclosed in membrane bags to avoid Fe contamination of the root surface by soil particles. Due to the high redox buffer capacity of the Haplic Luvisol employed for the experiments, flooding treatments induced only moderately reducing conditions (～?300 mV) and a slight increase of extraplasmatic Fe loading (41\to165 mg kg^?1 d.m.). Strongly reducing conditions (?100 mV) associated with a high Fe²⁺ concentration in the soil solution and a significant increase of extraplasmatic Fe (1190 mg kg^?1 d.m.) were obtained only after application of high amounts of organic carbon (10 g starch and 10 g cellulose kg^?1 soil), which accompanied by unrealistic reducing conditions due to intense stimulation of microbial growth. The expression of effects only under extremely high application level of organic carbon (～?33 t C ha^?1) suggest that similar to aerobic conditions, extraplasmatic Fe-loading under transient hypoxia is probably of limited ecological significance for the iron nutrition of higher plants, at least in soils with a high redox buffer capacity as employed in the present study. Abbreviations: DHA – dehydrogenase activity; d.m. – dry matter; DOC – dissolved organic carbon; Eh – redox potential; PIXE – proton-induced X-ray emission; STIM – scanning transmission ion microscopy.     

Biogeochemistry of terrestrial soils as influenced by short-term flooding

Many terrestrial soils in the US Midwest are temporally flooded during the spring. The effects of short-term flooding on biogeochemical processes that occur in these soils are not fully understood and are the subject of this study. To evaluate these processes we investigated the redox-induced changes in the soil solution for three-cultivated and three-uncultivated/forest soils with different organic matter concentrations. The soils were flooded for 1, 3, 7, and 14-days under anoxic conditions in a biogeochemical reactor. Samples were analyzed for Eh; pH; NO₃ ^?; NH₄ ⁺; total dissolved Mn and Fe; soluble P; dissolved organic and inorganic carbon (DOC–DIC); and evolved CO₂. We found strongly contrasting responses of the terrestrial soils to flooding. Reducing conditions were established quickly in the uncultivated and more slowly in the cultivated soils. Concomitant changes in pH were higher for the uncultivated soils. The uncultivated soils showed a higher increase in the amount of NH₄ ⁺, P, Fe, Mn than the cultivated soils over the 14-day incubation. The total amount of carbon decomposed was much greater for the uncultivated soils with approximately 900 μg C (CO₂ + DOC + DIC) decomposed per gram of soil compared to a total decomposition of 240 μg C g _soil ^?1 for the cultivated soils indicating differences in the type of carbon decomposed. The rapid onset of reducing conditions for the uncultivated soils is attributed to a reactive carbon component that is either absent or occluded in the cultivated soils. This study demonstrates that the biogeochemically-induced changes in carbon dynamics in terrestrial soils are strongly influenced by short-term flooding and the history of soil management.     

Some physiological and morphological responses of Pyrus boissieriana to flooding

European pear is a flooding-sensitive species, and for its cultivation in lowland areas, it is necessary to carry out the grafting of scions of commercial pear varieties into rootstocks belonging to flooding-tolerant wild pear species. Flooding tolerance of Pyrus boissieriana—a type of wild pear—was studied as a promissory rootstock for commercial pear. For this purpose, 3-month-old plants of P. boissieriana were subjected for 30 days to control (C), well-irrigated treatment, short-term (15 days) flooding plus 15 days recovery (F + R) and long-term (30 days) continuous flooding (F). Physiological performance, plant morphological changes and biomass accumulation were assessed. Results showed that, although stomatal conductance, transpiration and photosynthesis were progressively decreased by flooding, when flooding was short term (i.e., 2 weeks, F + R treatment) plants were able to adequately recover their physiological activity (50–74 % with respect to controls). In contrast, when plants continued to be flooded (F treatment), the physiological activity became null and the plants died quickly after the water subsided. Adventitious rooting was the most conspicuous registered morphological response to flooding, despite that flooded plants had shorter shoots and roots than control plants. Leaf and root biomass were 63 and 89 % higher under short-term flooding (F + R) than under continuous flooding (F), condition in which plants did not survive. In conclusion, P. boissieriana appears to be a promising species for its use as rootstock of commercial pear in lowland areas prone to flooding of up to 2 weeks. However, if the flooding period is extended, plants of this species are at risk of perishing.     

Sulfur and molybdenum fractionation in marine and riverine alluvium paddy soils

Intermittently submergence and drainage status of paddy fields can cause alterations in morphological and chemical characteristics of soils. We conducted a sequential fractionation study to provide an insight into solubility of Sulfur (S) and Molybdenum (Mo) in flooded alluvial paddy soils. The samples (0–15 and 15–30 cm) were taken from marine and riverine alluvial soils in Kedah and Kelantan areas, respectively, and were sequentially extracted with NaHCO₃, NaOH, HCl, and HClO₄–HNO₃. Total S in upper and lower layers of Kedah and Kelantan ranged between 273 and 1121 mg kg^?1, and 177 to 1509 mg kg^?1, respectively. In upper layers and subsoil of Kedah, average total Mo were 0.34 and 0.27 mg kg^?1, respectively. Average total Mo in Kelantan were 0.25 mg kg^?1 (surface layer) and 0.28 mg kg^?1 (subsoil). Cation exchange capacity (CEC) was positively correlated with plant available amounts of Mo in upper layers of Kedah area. Also, total and medium-term plant-available S was correlated with total carbon (C) at lower layers of Kelantan soil series. But in surface layers of Kelantan soil series, CEC was strongly correlated with total and medium-term plant-available S. Our results indicates that the influence of flooding conditions on soil S and Mo contents in paddy fields may cause long-term changes in S and Mo chemical reactivities.     

River-wetland interaction and carbon cycling in a semi-arid riverine system: the Okavango Delta, Botswana

The Okavango River, in semi-arid northwestern Botswana, flows for over 400 km in a pristine wetland developed on a large (>22,000 km²) alluvial fan (Okavango Delta). An annual flood pulse inundates the floodplains of the wetlands and travels across the Delta in 4–6 months. In this study, we assess the effects of long hydraulic residence time, variable hydrologic interaction between river–floodplain–wetland and evapotranspiration on carbon cycling. We measured dissolved inorganic carbon (DIC) concentrations and stable carbon isotopes of DIC (δ¹³C_DIC) from river water when the Delta was not flooded (low water) and during flooding (high water). During low water, the average DIC concentration was 31 % higher and the δ¹³C_DIC 2.1 ‰ more enriched compared to high water. In the lower Delta with seasonally flooded wetlands, the average DIC concentration increased by 70 % during low water and by 331 % during high water compared to the Panhandle with permanently flooded wetlands. The increasing DIC concentration downriver is mostly due to evapoconcentration from transpiration and evaporation with increased transit time. The average δ¹³C_DIC between low and high water decreased by 3.7 ‰ in the permanently flooded reaches compared to an increase of 1.6 ‰ in the seasonally flooded reaches. The lower δ¹³C_DIC during high water in the permanently flooded reaches suggest that DIC influx from the floodplain-wetland affects river’s DIC cycling. In contrast, higher river channel elevations relative to the wetlands along seasonal flooded reaches limit hydrologic interaction and DIC cycling occurs mostly by water column processes and river-atmospheric exchange. We conclude that river-wetlands interaction and evapoconcentration are important factors controlling carbon cycling in the Okavango Delta.     

Oxygen input controls the spatial and temporal dynamics of arsenic at the surface of a flooded paddy soil and in the rhizosphere of lowland rice (Oryza sativa L.): a microcosm study

The impact of oxygen (O₂) input at the soil surface and in the rhizosphere of rice (Oryza sativa L.) on the spatial and temporal dynamics of arsenic (As) was investigated in a flooded paddy soil. A soil microcosm and root-mat technique were designed to mimic submerged conditions of paddy fields. Water-filled containers with (planted) or without (unplanted) 27-day-old rice seedlings were fitted for 20 days on top of microcosms containing an As-affected soil (Bangladesh). After the initial establishment of strongly reduced conditions (?230 mV) in both planted and unplanted soils, the redox potential gradually increased until the day 8 to reach?+?50 mV at 2 mm from the surface of unplanted soils only. This oxidation was associated with an accumulation of NH₄-oxalate extractable As (25.7 mg kg^?1) in the 0.5-mm top layer, i.e. at levels above the initial total content of As in the soil (14 mg kg^?1) and a subsequent depletion of As in soil solution at 2 mm from soil surface. Root O₂-leakage induced the formation of an iron (Fe) plaque in root apoplast, with no evidence of outer rhizosphere oxidation. Arsenic content reached 173 mg kg^?1 in the Fe plaque. This accumulation induced a depletion of As in soil solution over several millimetres in the rhizosphere. Arsenic contents in root symplast and shoots (112 and 2.3 mg kg^?1, respectively) were significantly lower than in Fe plaque. Despite a large As concentration in soil solution, Fe plaque appeared highly efficient to sequester As and to restrict As acquisition by rice. The oxidation-mediated accumulation of As in the Fe plaque and in the oxidised layer at the top of the soil mobilised 21 and 3% of the initial amount of As in the planted and unplanted soils, respectively. Soil solution As concentration steadily decreased during the last 16 days of the soil stage, likely indicating a decrease in the ability of the soil to re-supply As from the solid-phase to the solution. The driving force of As dynamic in soil was therefore attributed to the As diffusion from reduced to oxidised soil layers. These results suggest a large mobility of As in the soil during the flooded period, controlled by the setting of oxic/anoxic interfaces at the surface of soil in contact with flooding water and in the rhizosphere of rice.     

Effect of selenium on growth and antioxidant capacity of <Emphasis Type="Italic">Triticum aestivum</Emphasis> L. during development of lead-induced oxidative stress

Effects of two selenium concentrations—0.4 and 0.8 mg Se⁶⁺ per kilogram of soil (treatments Se0.4 and Se0.8)—on seedling growth, chlorophyll content (Chl (a + b)), the content of thiobarbituric acidreactive substances (TBARs) indicative of peroxidation rates, and the activities of antioxidant enzymes (ascorbate peroxidase, AsP; glutathione reductase, GR; and guaiacol peroxidase, GPX) were studied in roots and leaves of wheat (Triticum aestivum L., cv. Triso) plants that were exposed for 14 days to oxidative stress induced by 50 and 100 mg Pb²⁺ per kilogram of soil (treatments Pb50 and Pb100, respectively). The pollution of soil with Pb²⁺ inhibited growth, lowered Chl (a + b) content, and intensified peroxidation. The content of TBARs increased by 44 and 72% in leaves and by 25 and 45% in roots for treatments Pb50 and Pb100, respectively. Activities of the antioxidant enzymes GR and GPX were higher in Pb²⁺-treated than in untreated (control) plants. The introduction of Se⁶⁺ into Pb²⁺-free soil (treatment Se0.4) was found to promote growth, stimulate AsP activity by 40% in leaves, and enhance AsP, GR, and GPX activities in roots by 38, 33, and 74%, respectively. The content of TBARs was reduced in Se⁶⁺-treated plants. By contrast, the treatment Se0.8 suppressed growth, elevated TBARs content, and stimulated activities of antioxidant enzymes in roots and leaves. The addition of 0.4 mg Se⁶⁺/kg to Pb²⁺-contaminated soil alleviated the negative influence of lead on plant growth, whereas the addition of 0.8 mg Se⁶⁺/kg aggravated the effect of Pb²⁺ contamination, especially in treatment (Pb100+Se0.8). Thus, the effectiveness of exogenous Se⁶⁺ application on growth and adaptive potential of plants cultivated under optimal pollutant-free conditions and on soils contaminated with lead depended on the concentration of Se⁶⁺ supplemented to soil and on the content of the stressor agent.     

Growth and chlorophyll a fluorescence in Erythrina crista-galli L. plants under flooding conditions

The present study aimed to analyse the alterations in the growth and chlorophyll content of Erythrina crista-galli plants in response to flooding and to evaluate possible photosystem II (PSII) damage through chlorophyll a fluorescence transient analysis using the JIP test. Plants cultured from seeds were grown in 0.5-L pots in a greenhouse and transferred to 5-L pots. Two treatments were employed: plants were either flooded at the root and maintained with a sheet of water above the soil or not flooded (control). Evaluations were performed at 10, 20, 30 and 40 days after the treatment was initiated. The experiment was set up in a randomised block design, and the results were subjected to analysis of variance (ANOVA). The means were compared by Tukey’s test at a 5 % significance level. The flooding of the plants induced the formation of lenticels, adventitious roots and larger intercellular spaces in the root cortex and inhibited the growth of shoots at early time points compared to control plants. However, growth resumed by the end of the experimental period. The Erythrina crista-galli plants manifested a reduction in chlorophyll content as a function of the duration of exposure to flooding. However, there were changes in the chlorophyll fluorescence parameters, indicating impairment of the structure and function of PSII, and the performance indices were the most representative parameters to describe the effects of flooding. The results showed that the Erythrina crista-galli tolerated flooding by initially investing energy into the formation of morpho-anatomical structures to maintain oxygen diffusion and ATP production. The effect of flooding on the leaves was reflected in increased energy dissipation through chlorophyll a fluorescence, likely as a mechanism to protect the photosynthetic apparatus.     

Do changes in flood pulse duration disturb soil carbon dioxide emissions in semi-arid floodplains?

In semi-arid floodplains the average times between floods have been cited to drive metabolic and biogeochemical responses during the subsequent flooding pulse. However, the interaction effects of flood pulse duration and the length of time between floods on the carbon budget are not well understood. Using field experiments, flood pulses—dry cycles were simulated (SF plots—short flood/dry cycles: 15 flood days + 7 dry + 15 flood and LF plots—long flood/dry cycles: 21 flood + 14 dry + 21 flood) in a semi-arid floodplain in Central Spain, in order to study the effects on soil CO₂ emissions. Differences on soil water content among SF, LF and control plots were statistically significant throughout the experiment (p < 0.01). Soil CO₂ emission rates during drying time were significantly related with the duration of previous flooding and inter-flooding intervals (R ² = 0.52–0.64, p = 0.03). During the first stage of desiccation, the high soil water content appears to limit aerobic metabolism. Soil respiration rates similar to those of control plots measurements occurred 1–2 weeks later. Then, soil respiration increased to a maximum rate which was delayed 5–8 weeks, as high soil water content limited microbial activity. While more than 7 days of inundation promoted denitrification, organic nutrients supplied by flood water increased 1% soil respiration during drying. Differences between SF and LF plots in soil CO₂ emissions only appeared after floodplain soil had been subjected to two consecutive flood-dry cycles; 70 days after the second inundation ended, CO₂ fluxes achieved similar values in all treatments. Daily soil CO₂ emission rates during the entire study period (117 days) were comparable, independently of the flood duration and the time between floods (75.76 ± 1.59 and 77.94 ± 0.45 mmol CO₂ m^?2 day^?1, in SF and LF, respectively). Flood disturbance affects site-specific microbial processes, but only during very short time periods. The mechanism by which soil microbial communities cope or adapt to new conditions needs to be reassessed in future research in order to determine the long-term effects of hydrological changes in the soil carbon balance of semi-arid floodplains.     

Antioxidative responses and morpho-anatomical adaptations to waterlogging in Sesbania virgata

Sesbania virgata (Leguminosae) is tolerant of long periods of soil inundation. However, its morphological adaptations to anoxia and its response to possible damage from oxidative stress are still unknown. Here, we provide new information that helps to explain the ability of S. virgata plants to grow in flooded environments. Plants containing six expanded leaves were placed in masonry tanks and were subjected to the following conditions: control (well watered), soil waterlogging (water to the setup level of 1 cm above the soil surface—roots and parts of the stems flooded), and complete submergence (whole plant flooded). Plants exposed to flooding (soil waterlogging and complete submergence) significantly increased their production of hydrogen peroxide (H₂O₂), indicating the extent of oxidative injury posed by stress conditions. We demonstrate that plants exposed to flooding develop an efficient scavenger of ROS (generated during stress) in the roots through the coordinated action of nonenzymatic ascorbic acid (Asc) and dehydroascorbate (DHA) as well as the enzymatic antioxidants superoxide dismutase (SOD), ascorbate peroxidase (APX), dehydroascorbate reductase (DHAR), and glutathione reductase (GR) that are present in the tissues. Moreover, we observed the development of morpho-anatomical structures such as adventitious roots, lenticels, and cracks in the stem of plants under soil waterlogging. The secondary root of plants under soil waterlogging showed a thinner cortex and larger number of elements of small diameter vessels. Numerous aerenchymas were observed in the newly formed in the adventitious roots. We conclude that these antioxidative responses and morpho-anatomical adaptations in the roots are part of a suite of adaptations that allow S. virgata plants to survive long periods of flooding, notably under waterlogged conditions.     

Dissolved organic matter and nutrient dynamics of a coastal freshwater forested wetland in Winyah Bay, South Carolina

Seasonally flooded, freshwater cypress-tupelo wetlands, dominated by baldcypress (Taxodium distictum), water tupelo (Nyssa aquatica), and swamp tupelo (Nyssa sylvatica var. biflora) are commonly found in coastal regions of the southeastern United States. These wetlands are threatened due to climate change, sea level rise, and coastal urban development. Understanding the natural biogeochemical cycles of nutrients in these forested wetlands as ecosystems services such as carbon sequestration and nitrogen processing can provide important benchmarks to guide conservation plans and restoration goals. In this study, surface water and soil pore water samples were collected weekly from a cypress-tupelo wetland near Winyah Bay, South Carolina and analyzed for dissolved organic carbon (DOC), dissolved organic nitrogen (DON), inorganic nitrogen, and phosphate during its flooding period between October 2010 and May 2011. DOC was further characterized by specific ultra-violet absorbance at 254 nm, spectral slope ratio (S_R) (ratio of two spectral slopes between 275–295 nm and 350–400 nm), E2/E3 ratio (ratio between A254 and A365), and fluorescence excitation-emission matrix. In addition, litterfall was collected on a monthly basis for a year while the biomass of the detritus layer (i.e., decomposed duff lying on the wetland floor) was determined before and after the flooding period. Results of the field study showed that concentrations of DOC, DON, NH₄ ⁺–N, and (NO₂ ^? + NO₃ ^?)–N in the surface water were generally higher during the fall, or peak litterfall season (October to December), than in the spring season (March to May). Highest concentrations of 54.8, 1.48, 0.270, and 0.0205 mg L^?1, for DOC, DON, NH₄ ⁺–N, and (NO₂ ^? + NO₃ ^?)–N respectively, in surface waters were recorded during October. Lower SUVA, but higher S_R and E2/E3 ratios of DOC, were observed at the end of the flooding season comparing to the initial flooding, suggesting the wetland system converts high aromatic and large DOC molecules into smaller and hydrophilic fractions possibly through photochemical oxidation. A similar trend was observed in soil pore water, but the pore water generally had greater and relatively stable concentrations of dissolved nutrients than surface water. No obvious temporal trend in phosphate concentration and total nitrogen to total phosphorus ratio (N:P) were found. Results of the laboratory extraction and mass balance calculation suggested fresh litter was a major source of DOC whereas decomposed duff was the source of dissolved nitrogen in surface water. In summary, the biogeochemistry of this isolated cypress-tupelo wetland is not only driven by the vegetation within the wetland system but also by hydrology and weather conditions such as groundwater table position, precipitation, and temperature.     

Methane oxidation and methane driven redox process during sequential reduction of a flooded soil ecosystem

A laboratory incubation study conducted to assess the temporal variation of CH₄ oxidation during soil reduction processes in a flooded soil ecosystem. A classical sequence of microbial terminal electron accepting process observed following NO₃ ^? reduction, Fe³⁺ reduction, SO₄ ^2? reduction and CH₄ production in flooded soil incubated under initial aerobic and helium-flushed anaerobic conditions. CH₄ oxidation in the slurries was influenced by microbial redox process during slurry reduction. Under aerobic headspace condition, CH₄ oxidation rate (k) was stimulated by 29 % during 5 days (NO₃ ^? reduction) and 32 % during both 10 days (Fe³⁺) and 20 days (early SO₄ ^2? reduction) over unreduced slurry. CH₄ oxidation was inhibited at the later methanogenic period. Contrastingly, CH₄ oxidation activity in anaerobic incubated slurries was characterized with prolonged lag phase and lower CH₄ oxidation. Higher CH₄ oxidation rate in aerobically incubated flooded soil was related to high abundance of methanotrophs (r?=?0.994, p?<?0.01) and ammonium oxidizers population (r?=?0.184, p?<?0.05). Effect of electron donors NH₄ ⁺, Fe^2+, S^2? on CH₄ oxidation assayed to define the interaction between reduced inorganic species and methane oxidation. The electron donors stimulated CH₄ oxidation as well as increased the abundance of methanotrophic microbial population except S^2? which inhibited the methanotrophic activity by affecting methane oxidizing bacterial population. Our result confirmed the complex interaction between methane-oxidizing microbial groups and redox species during sequential reduction processes of a flooded soil ecosystem.     

Induced anti-oxidant activity in soybean alleviates oxidative stress under moderate boron toxicity

The effect of B toxicity on antioxidant responses of soybean (Glycine max) cv. Athow was investigated by growing plants for 43 days at 0.2 (control), 2 and 12 mg B kg^?1. At the end of the treatment period, shoot growth, lipid peroxidation level, the activities of antioxidant enzymes including superoxide dismutase (SOD), peroxidase (POX), catalase (CAT), ascorbate peroxidase (APX) and glutathione reductase (GR), and their isoenzymes in leaves were measured. Boron concentration in leaves was significantly increased by the increasing levels of B treatment from 43 to 522 mg kg^?1, and shoot dry matter was depressed at 12 mg B kg^?1. Significant increases in SOD, CAT, and APX activities were determined in leaves under 12 mg B kg^?1; however, GR activities were decreased while POX activity was unchanged. Increased enzymic antioxidant activity arose from a combination of newly formed isoenzymes and activation of existing isoenzymes. By contrast, SOD and GR activities were decreased by 2 mg B kg^?1 concentration as compared to the control groups while POX activity was increased and the activity of CAT did not change. Malondialdehyde content increased under 2 mg B kg^?1 but decreased under 12 mg B kg^?1. These results suggest that higher antioxidant activity observed under 12 than at 2 mg B kg^?1 provided higher free radical-scavenging capacity, and thus a lower level of lipid peroxidation in Athow. While the induction of increased antioxidant activity was related to internal boron levels, the signaling and coordination of responses remain unclear.     

Effect of soil flooding on photosynthesis,carbohydrate partitioning and nutrient uptake in the invasive exotic Lepidium latifolium

Lepidium latifolium L. is an invasive exotic crucifer that has spread explosively in wetlands and riparian areas of the western United States. To understand the ecophysiological characteristics of L. latifolium that affect its ability to invade riparian areas and wetlands, we examined photosynthesis, chlorophyll concentration, carbohydrate partitioning and nutrient uptake in L. latifolium in response to soil flooding. Photosynthesis of flooded plants was about 60–70% of the rate of unflooded controls. Chlorophyll concentrations of flooded plants were about 60–70% of the unflooded plants during 15–50 days of flooding. Flooding resulted in an increase in leaf starch concentration, but root starch concentration was not significantly affected. However, concentrations of soluble sugar were significantly higher in both leaves and roots of flooded plants than unflooded controls. On day 50 after initial flooding, the concentrations of N, P, K and Zn in leaves of flooded plants were lower than in control plants. The concentrations of Mn and Fe in leaves of flooded plants were eight and two times those of control plants, respectively. In contrast, N, P, K and Zn concentrations of roots of flooded plants were slightly higher than in unflooded plants. The concentrations of Fe and Mn in roots of flooded plants were 15 and 150 times those of the control plants, respectively. The transport of P, K, and Zn to shoots decreased and that of Mn increased under flooding. The accumulation of N, K and Zn in roots decreased and that of Mn increased in response to flooding. The results suggested that the maintenance of relatively high photosynthesis and the accumulation of soluble sugar in roots of flooded plants are important adaptations for this species in flooded environments. Despite a reduction in photosynthesis and disruption in nutrient and photosynthate allocation in response to flooding, L. latifolium was able to survive 50 days of flooding stress. Overall, L. latifolium performed like a facultative hydrophyte species under flooding.     

Flooding-induced membrane damage, lipid oxidation and activated oxygen generation in corn leaves

Flooding effects on membrane permeability, lipid peroxidation and activated oxygen metabolism in corn (Zea mays L.) leaves were investigated to determine if activated oxygens are involved in corn flooding-injury. Potted corn plants were flooded at the 4-leaf stage in a controlled environment. A 7-day flooding treatment resulted in a significant increase in chlorophyll breakdown, lipid peroxidation (malondialdehye content), membrane permeability, and the production of superoxide (O ₂ ^- ) and hydrogen peroxide (H₂O₂) in corn leaves. The effects were much greater in older leaves than in younger ones. Spraying leaves with 8-hydroxyquinoline (an O ₂ ^- scavenger) and sodium benzoate (an .OH scavenger) reduced the oxidative damage and enhanced superoxide dismutase (SOD) activity. A short duration flooding treatment elevated the activities of SOD, catalase, ascorbate peroxidase (AP), and glutathione reductase (GR), while further flooding significantly reduced the enzyme activities but enhanced the concentrations of ascorbic acid and reduced form glutathione (GSH). It was noted that the decline in SOD activity was greater than that in H₂O₂ scavengers (AP and GR). The results suggested that O ₂ ^- induced lipid peroxidation and membrane damage, and that excessive accumulation of O ₂ ^- is due to the reduced activity of SOD under flooding stress.     

Interactions among phosphorus, pH and Eh in reforested mangroves, Vietnam: a three-dimensional spatial analysis

Sediment reduction is frequently proposed to increase available phosphorus (P) but several studies found also decreases. Another important factor for P liberation is the pH. We investigated the relative importance of Eh and pH on P cycling in reforested mangroves. Sediment P compounds, pH and Eh were analysed over depth along five transects of two areas in the Saigon River Delta and compared with leaf P levels. A three-dimensional spatial approach was used to investigate pH and Eh effects on P compound distribution at different sediment depth and locate layers of predominant P uptake. Along an inundation gradient, submergence durations of 254 to 2 days per year caused a large Eh gradient within the top 20 cm, whereas Eh response was small within 20–45 cm depth. At individual layers, a correlation between Eh and Al/Fe–P was only found in the upper depth interval (0–20 cm). No significant effect of Eh or Al/Fe–P on Morgan-P (available P) was detected. Minor effects on P composition changes by the Eh were caused through generally strongly reduced sediment at deeper layers (>20 cm). In contrast, pH variations produced extreme differences in both, Ca–P and available P content at layers of apparently predominant P uptake (>20 cm). As available P was correlated with Ca–P (p < 0.001), leaf P (p < 0.001) and pH (non linear correlation) it is likely that the pH sensitive Ca–P fraction is a more effective source for mangrove tree growth than Al/Fe–P. The predominant pH effect on P uptake within these reforested mangroves differs from a proposed reduction-governed P cycling in wetlands.     

Morphological and photosynthetic responses of riparian plant Distylium chinense seedlings to simulated Autumn and Winter flooding in Three Gorges Reservoir Region of the Yangtze River, China

To evaluate the tolerance of riparian plant Distylium chinense in Three Gorges Reservoir Region to anti-season flooding, a simulation flooding experiment was conducted during Autumn and Winter, and morphology and photosynthesis of D. chinense seedlings and their recovery growth after soil drainage were analyzed in different duration of flooding and flooding depth. The seedlings were submitted to four treatments: (1) 40 seedlings unflooded and watered daily as control (Unflooded, CK); (2) 120 seedlings flooded at 1 cm above the ground level (F-1 cm); (3) 120 seedlings flooded at 12 cm above the ground level (F-12 cm) and (4) 120 seedlings completely submerged with 2 m water depth (F-2 m, top of plants at 2 m below water surface). The flooding survival, plant height, stem diameter, adventitious roots, stem lenticels, epicormic shoots, chlorophyll content and photosynthesis parameters were determined at 0, 15, 30, 90 days in flooding stress and 15, 60 days after soil drainage. The results showed that the survival of the seedlings subjected to flooding was 100% for all repeated measurements in all treatments. Adventitious roots, hypertrophied lenticels and stem hypertrophy were observed in the seedlings flooded for more than 15 d, and increased with the prolonged flooding duration, while disappeared after the soil was drained. Flooding duration and flooding depth showed significant individual and interactive effects on leaf chlorophyll a (Chl a), chlorophyll b (Chl b), and their ratio, chlorophyll (a + b), the net photosynthesis rate (P_n), transpiration rate (T_r), stomatal conductance (C_s), and inter-cellular CO₂ concentration (C_i) of D. chinense seedlings (P < 0.01). After 15 days of flooding, there was no significant decrease in P_n of the flooded seedlings as compared with that of the control seedlings. P_n of the flooded seedlings was significantly lower than that of the control seedlings after 30 days of flooding (P < 0.05), whereas P_n showed no significant difference among seedlings from three flooding depths. After 90 days of flooding, P_n of the F-2 m flooded seedlings was significantly lower than that of the controls, F-1 cm and F-12 cm flooded seedlings (P < 0.05), but still maintained high photosynthetic capacity. P_n of the F-1 cm and F-12 cm flooded seedlings rose gradually after soil drainage, while, it was significantly lower than that of the control seedlings after 15 days of recovery (P < 0.05). After 60 days of recovery, P_n of all seedlings flooded with different depths showed no significant difference as compared with that of the control seedlings and new leaves grew out in the F-2 m flooded seedlings. The effect of all flooding treatments on Gs, T_r, Chl a, Chl b, Chl a/Chl b and chl (a + b) was basically the same as their effect on P_n, while the effect of all flooding treatments on C_i was quite the contrary. Correlation analysis showed that P_n was positively relative with Gs, T_r, Chl a, Chl b and chl (a + b) (P < 0.05) and significantly negative with C_i (P < 0.05). Therefore, the present study demonstrates that D. chinense has high survival and good recovery growth after long-term flooding in anti-season flooding and could be taken as an excellent candidate species in the re-vegetation of water-level-fluctuation areas in Three Gorges Reservoir Region.     

Morphological and photosynthetic responses of riparian plant Distylium chinense seedlings to simulated Autumn and Winter flooding in Three Gorges Reservoir Region of the Yangtze River, China

To evaluate the tolerance of riparian plant Distylium chinense in Three Gorges Reservoir Region to anti-season flooding, a simulation flooding experiment was conducted during Autumn and Winter, and morphology and photosynthesis of D. chinense seedlings and their recovery growth after soil drainage were analyzed in different duration of flooding and flooding depth. The seedlings were submitted to four treatments: (1) 40 seedlings unflooded and watered daily as control (Unflooded, CK); (2) 120 seedlings flooded at 1 cm above the ground level (F-1 cm); (3) 120 seedlings flooded at 12 cm above the ground level (F-12 cm) and (4) 120 seedlings completely submerged with 2 m water depth (F-2 m, top of plants at 2 m below water surface). The flooding survival, plant height, stem diameter, adventitious roots, stem lenticels, epicormic shoots, chlorophyll content and photosynthesis parameters were determined at 0, 15, 30, 90 days in flooding stress and 15, 60 days after soil drainage. The results showed that the survival of the seedlings subjected to flooding was 100% for all repeated measurements in all treatments. Adventitious roots, hypertrophied lenticels and stem hypertrophy were observed in the seedlings flooded for more than 15 d, and increased with the prolonged flooding duration, while disappeared after the soil was drained. Flooding duration and flooding depth showed significant individual and interactive effects on leaf chlorophyll a (Chl a), chlorophyll b (Chl b), and their ratio, chlorophyll (a + b), the net photosynthesis rate (P_n), transpiration rate (T_r), stomatal conductance (C_s), and inter-cellular CO₂ concentration (C_i) of D. chinense seedlings (P < 0.01). After 15 days of flooding, there was no significant decrease in P_n of the flooded seedlings as compared with that of the control seedlings. P_n of the flooded seedlings was significantly lower than that of the control seedlings after 30 days of flooding (P < 0.05), whereas P_n showed no significant difference among seedlings from three flooding depths. After 90 days of flooding, P_n of the F-2 m flooded seedlings was significantly lower than that of the controls, F-1 cm and F-12 cm flooded seedlings (P < 0.05), but still maintained high photosynthetic capacity. P_n of the F-1 cm and F-12 cm flooded seedlings rose gradually after soil drainage, while, it was significantly lower than that of the control seedlings after 15 days of recovery (P < 0.05). After 60 days of recovery, P_n of all seedlings flooded with different depths showed no significant difference as compared with that of the control seedlings and new leaves grew out in the F-2 m flooded seedlings. The effect of all flooding treatments on Gs, T_r, Chl a, Chl b, Chl a/Chl b and chl (a + b) was basically the same as their effect on P_n, while the effect of all flooding treatments on C_i was quite the contrary. Correlation analysis showed that P_n was positively relative with Gs, T_r, Chl a, Chl b and chl (a + b) (P < 0.05) and significantly negative with C_i (P < 0.05). Therefore, the present study demonstrates that D. chinense has high survival and good recovery growth after long-term flooding in anti-season flooding and could be taken as an excellent candidate species in the re-vegetation of water-level-fluctuation areas in Three Gorges Reservoir Region.     

The system modulating ROS content in germinating seeds of two Brazilian savanna tree species exposed to As and Zn

The effects of increasing arsenic (0, 10, 50, 100 mg L^?1) and zinc (0, 50, 80, 120, 200 mg L^?1) doses on germination and oxidative stress markers (H₂O₂, MDA, SOD, CAT, APX, and GR) were examined in two Brazilian savanna tree species (Anadenanthera peregrina and Myracrodruon urundeuva) commonly used to remediate contaminated soils. The deleterious effects of As and Zn on seed germination were due to As- and Zn-induced H₂O₂ accumulation and inhibition of APX and GR activities, which lead to oxidative damage by lipid peroxidation. SOD and CAT did not show any As- and Zn-induced inhibition of their activities as was seen with APX and GR. We investigated the close relationships between seed germination success under As and Zn stress in terms of GR and, especially, APX activities. Increased germination of A. peregrina seeds exposed to 50 mg L^?1 of Zn was related to increased APX activity, and germination in the presence of As (10 mg L^?1) was observed only in M. urundeuva seeds that demonstrated increased APX activity. All the treatments for both species in which germination decreased or was inhibited showed decreases in APX activity. A. peregrina seeds showed higher Zn-tolerance than M. urundeuva, while the reverse was observed with arsenic up to exposures of 10 mg L^?1.     

Involvement of antioxidant defense system in chill hardening-induced chilling tolerance in Jatropha curcas seedlings

Jatropha curcas L. is a sustainable energy plant with great potential for biodiesel production, and low temperature is an important limiting factor for its distribution and production. In this present work, chill hardening-induced chilling tolerance and involvement of antioxidant defense system were investigated in J. curcas seedlings. The results showed that chill hardening at 10 or 12 °C for 1 and 2 days greatly lowered death rate and alleviated electrolyte leakage as well as accumulation of the lipid peroxidation product malondialdehyde (MDA) of J. curcas seedlings under severe chilling stress at 1 °C for 1–7 days, indicating that the chill hardening significantly improved chilling tolerance of J. curcas seedlings. Measurement of activities of the antioxidant enzymes superoxide dismutase (SOD), ascorbate peroxidase (APX), catalase (CAT), peroxidase (POD), and glutathione reductase (GR), and the levels of the antioxidants ascorbic acid (AsA) and glutathione (GSH) showed the chill hardening at 12 °C for 2 days could obviously increase the activities of these antioxidant enzymes and AsA and GSH contents in the hardened seedlings. When the hardened and non-hardening (control) seedlings were subjected to severe chilling stress at 1 °C for 1–7 days, the chill-hardened seedlings generally maintained significantly higher activities of the antioxidant enzymes SOD, APX, CAT, POD, and GR, and content of the antioxidants AsA and GSH as well as ratio of the reduced antioxidants to total antioxidants [AsA/(AsA + DHA) and GSH/(GSH + GSSG)], when compared with the control without chill hardening. All above-mentioned results indicated that the chill hardening could enhance the chilling tolerance, and the antioxidant defense system plays an important role in the chill hardening-induced chilling tolerance in J. curcas seedlings.