Role of ethylene in alleviation of cadmium-induced photosynthetic capacity inhibition by sulphur in mustard

Sulphur (S) assimilation leads to the formation of glutathione (GSH) and alleviation of cadmium (Cd) stress. GSH is synthesized from its immediate metabolite cysteine, which also serves as a metabolite for ethylene formation through S‐adenosyl methionine. To assess the role of ethylene in S‐induced alleviation of Cd stress on photosynthesis, the effects of S or ethephon (ethylene source) on GSH and ethylene were examined in mustard (Brassica juncea L. cv. Varuna). Sufficient‐S at 100 mg S kg^?1 soil alleviated Cd‐induced photosynthetic inhibition more than excess‐S (200 mg S kg^?1 soil) via ethylene by increased GSH. Under Cd stress, plants were less sensitive to ethylene, despite high ethylene evolution, and showed photosynthetic inhibition. Ethylene sensitivity of plants increased with ethephon or sufficient‐S, triggering the induction of an antioxidant system, and leading to increased photosynthesis even under Cd stress. The effects of ethephon and S under Cd stress were similar. The effects of S were reversed by ethylene biosynthesis inhibitor, aminoethoxyvinylglycine (AVG), suggesting that ethylene plays an important role in S‐induced alleviation of Cd stress on photosynthesis.     

Variation in the carbon and oxygen isotope composition of plant biomass and its relationship to water‐use efficiency at the leaf‐ and ecosystem‐scales in a northern Great Plains grassland

Measurements of the carbon (δ¹³C_m) and oxygen (δ¹⁸O_m) isotope composition of C₃ plant tissue provide important insights into controls on water‐use efficiency. We investigated the causes of seasonal and inter‐annual variability in water‐use efficiency in a grassland near Lethbridge, Canada using stable isotope (leaf‐scale) and eddy covariance measurements (ecosystem‐scale). The positive relationship between δ¹³C_m and δ¹⁸O_m values for samples collected during 1998–2001 indicated that variation in stomatal conductance and water stress‐induced changes in the degree of stomatal limitation of net photosynthesis were the major controls on variation in δ¹³C_m and biomass production during this time. By comparison, the lack of a significant relationship between δ¹³C_m and δ¹⁸O_m values during 2002, 2003 and 2006 demonstrated that water stress was not a significant limitation on photosynthesis and biomass production in these years. Water‐use efficiency was higher in 2000 than 1999, consistent with expectations because of greater stomatal limitation of photosynthesis and lower leaf c_i/ca during the drier conditions of 2000. Calculated values of leaf‐scale water‐use efficiency were 2–3 times higher than ecosystem‐scale water‐use efficiency, a difference that was likely due to carbon lost in root respiration and water lost during soil evaporation that was not accounted for by the stable isotope measurements.     

Antioxidant enzyme activities and hormonal status in response to Cd stress in the wetland halophyte Kosteletzkya virginica under saline conditions

Salt marshes constitute major sinks for heavy metal accumulation but the precise impact of salinity on heavy metal toxicity for halophyte plant species remains largely unknown. Young seedlings of Kosteletzkya virginica were exposed during 3 weeks in nutrient solution to Cd 5 µM in the presence or absence of 50 mM NaCl. Cadmium (Cd) reduced growth and shoot water content and had major detrimental effect on maximum quantum efficiency (F_v/F_m), effective quantum yield of photosystem II (Y(II)) and electron transport rates (ETRs). Cd induced an oxidative stress in relation to an increase in O₂^?? and H₂O₂ concentration and lead to a decrease in endogenous glutathione (GSH) and α‐tocopherol in the leaves. Cd not only increased leaf zeatin and zeatin riboside concentration but also increased the senescing compounds 1‐aminocyclopropane‐1‐carboxylic acid (ACC) and abscisic acid (ABA). Salinity reduced Cd accumulation already after 1 week of stress but was unable to restore shoot growth and thus did not induce any dilution effect. Salinity delayed the Cd‐induced leaf senescence: NaCl reduced the deleterious impact of Cd on photosynthesis apparatus through an improvement of F_v/F_m, Y(II) and ETR. Salt reduced oxidative stress in Cd‐treated plants through an increase in GSH, α‐tocopherol and ascorbic acid synthesis and an increase in glutathione reductase (EC 1.6.4.2) activity. Additional salt reduced ACC and ABA accumulation in Cd+NaCl‐treated leaves comparing to Cd alone. It is concluded that salinity affords efficient protection against Cd to the halophyte species K. virginica, in relation to an improved management of oxidative stress and hormonal status.     

After more than a decade of soil moisture deficit,tropical rainforest trees maintain photosynthetic capacity,despite increased leaf respiration

Determining climate change feedbacks from tropical rainforests requires an understanding of how carbon gain through photosynthesis and loss through respiration will be altered. One of the key changes that tropical rainforests may experience under future climate change scenarios is reduced soil moisture availability. In this study we examine if and how both leaf photosynthesis and leaf dark respiration acclimate following more than 12 years of experimental soil moisture deficit, via a through‐fall exclusion experiment (TFE) in an eastern Amazonian rainforest. We find that experimentally drought‐stressed trees and taxa maintain the same maximum leaf photosynthetic capacity as trees in corresponding control forest, independent of their susceptibility to drought‐induced mortality. We hypothesize that photosynthetic capacity is maintained across all treatments and taxa to take advantage of short‐lived periods of high moisture availability, when stomatal conductance (g_s) and photosynthesis can increase rapidly, potentially compensating for reduced assimilate supply at other times. Average leaf dark respiration (R_d) was elevated in the TFE‐treated forest trees relative to the control by 28.2 ± 2.8% (mean ± one standard error). This mean R_d value was dominated by a 48.5 ± 3.6% increase in the R_d of drought‐sensitive taxa, and likely reflects the need for additional metabolic support required for stress‐related repair, and hydraulic or osmotic maintenance processes. Following soil moisture deficit that is maintained for several years, our data suggest that changes in respiration drive greater shifts in the canopy carbon balance, than changes in photosynthetic capacity.     

Proteomic study of Carissa spinarum in response to combined heat and drought stress

Carissa spinarum is one of the secondary advantage plants grown in dry‐hot valleys in China, which can survive under stress conditions of high temperature and extreme low humidity. Here, we studied the physiological and proteomic changes of C. spinarum in response to 42°C heat stress treatment in combination with drought stress. Dynamic changes in the leaf proteome were analyzed at four time points during the stress treatment and recovery stages. Approximately, 650 protein spots were reproducibly detected in each gel. Forty‐nine spots changed their expression levels upon heat and drought treatment, and 30 proteins were identified by MS and 2‐D Western blot. These proteins were classified into several categories including HSP, photosynthesis‐related protein, RNA‐processing protein and proteins involved in metabolism and energy production. The potential roles of these stress‐responsive proteins are discussed.     

<Emphasis Type="Italic">Cannabis sativa</Emphasis> L. growing on heavy metal contaminated soil: growth,cadmium uptake and photosynthesis

The effects of different cadmium concentrations [17 mg(Cd) kg⁻¹(soil) and 72 mg(Cd) kg^{− 1}(soil)] on Cannabis sativa L. growth and photosynthesis were examined. Hemp roots showed a high tolerance to Cd, i.e. more than 800 mg(Cd) kg⁻¹(d.m.) in roots had no major effect on hemp growth, whereas in leaves and stems concentrations of 50 – 100 mg(Cd) kg⁻¹(d.m.) had a strong effect on plant viability and vitality. For control of heavy metal uptake and xylem loading in hemp roots, the soil pH plays a central role. Photosynthetic performance and regulation of light energy consumption were analysed using chlorophyll fluorescence analysis. Seasonal changes in photosynthetic performance were visible in control plants and plants growing on soil with 17 mg(Cd) kg⁻¹(soil). Energy distribution in photosystem 2 is regulated in low and high energy phases that allow optimal use of light and protect photosystem 2 from overexcitation, respectively. Photosynthesis and energy dissipation were negatively influenced by 72 mg(Cd) kg⁻¹(soil). Cd had detrimental effects on chlorophyll synthesis, water splitting apparatus, reaction centre, antenna and energy distribution of PS 2. Under moderate cadmium concentrations, i.e. 17 mg(Cd) kg⁻¹(soil), hemp could preserve growth as well as the photosynthesis apparatus, and long-term acclimation to chronically Cd stress occurred.     

Calcium acetate‐induced reduction of cadmium accumulation in Oryza sativa: Expression of auto‐inhibited calcium‐ATPase and cadmium transporters

• Calcium (Ca) signalling has an essential role in regulating plant responses to various abiotic stresses.
• This study applied Ca in various forms (Ca acetate and CaCl₂) and concentrations to reduce cadmium (Cd) concentration in rice and propose a possible mechanism through which Ca acts to control the Cd concentration in rice.
• The results showed that supplementation of Cd‐contaminated soil with Ca acetate reduced the Cd concentration in rice after exposure for 7 days in both hydroponic and soil conditions. The possible involvement of the auto‐inhibited Ca²⁺‐ATPase gene (ACA) might act to control the primary signal of the Cd stress response. The messages from ACA3 and ACA13 tended to up‐regulate the low‐affinity cation transporter (OsLCT1) and down‐regulate Cd uptake and the Cd translocation transporter, including the genes, natural resistance‐associated macrophage protein 5 (Nramp5) and Zn/Cd‐transporting ATPase 2 (HMA2), which resulted in a reduction in the Cd concentration in rice. After cultivation for 120 days, the application of Ca acetate into Cd‐contaminated soil inhibited Cd uptake of rice.
• Increasing the Ca acetate concentration in the soil lowered the Cd concentration in rice shoots and grains. Moreover, Ca acetate maintained rice productivity and quality whereas both aspects decreased under Cd stress.

     

Photosynthetic Responses of a Temperate Liana to Xylella fastidiosa Infection and Water Stress

Xylella fastidiosa is a xylem‐limited bacterial plant pathogen that causes bacterial leaf scorch in its hosts. Our previous work showed that water stress enhances leaf scorch symptom severity and progression along the stem of a liana, Parthenocissus quinquefolia, infected by X. fastidiosa. This paper explores the photosynthetic gas exchange responses of P. quinquefolia, with the aim to elucidate mechanisms behind disease expression and its interaction with water stress. We used a 2 × 2‐complete factorial design, repeated over two growing seasons, with high and low soil moisture levels and infected and non‐infected plants. In both years, low soil moisture levels reduced leaf water potentials, net photosynthesis and stomatal conductance at all leaf positions, while X. fastidiosa‐infection reduced these parameters at basally located leaves only. Intercellular CO₂ concentrations were reduced in apical leaves, but increased at the most basal leaf location, implicating a non‐stomatal reduction of photosynthesis in leaves showing the greatest disease development. This result was supported by measured reductions in photosynthetic rates of basal leaves at high CO₂ concentrations, where stomatal limitation was eliminated. Repeated measurements over the summer of 2000 showed that the effects of water stress and infection were progressive over time, reaching their greatest extent in September. By reducing stomatal conductances at moderate levels of water stress, P. quinquefolia maintained relatively high leaf water potentials and delayed the onset of photosynthetic damage due to pathogen and drought‐induced water stress. In addition, chlorophyll fluorescence measurements showed that P. quinquefolia has an efficient means of dissipating excess light energy that protects the photosynthetic machinery of leaves from irreversible photoinhibitory damage that may occur during stress‐induced stomatal limitation of photosynthesis. However, severe stress induced by disease and drought eventually led to non‐stomatal decreases in photosynthesis associated with leaf senescence.     

Three‐dimensional microscale modelling of CO2 transport and light propagation in tomato leaves enlightens photosynthesis

We present a combined three‐dimensional (3‐D) model of light propagation, CO₂ diffusion and photosynthesis in tomato (Solanum lycopersicum L.) leaves. The model incorporates a geometrical representation of the actual leaf microstructure that we obtained with synchrotron radiation X‐ray laminography, and was evaluated using measurements of gas exchange and leaf optical properties. The combination of the 3‐D microstructure of leaf tissue and chloroplast movement induced by changes in light intensity affects the simulated CO₂ transport within the leaf. The model predicts extensive reassimilation of CO₂ produced by respiration and photorespiration. Simulations also suggest that carbonic anhydrase could enhance photosynthesis at low CO₂ levels but had little impact on photosynthesis at high CO₂ levels. The model confirms that scaling of photosynthetic capacity with absorbed light would improve efficiency of CO₂ fixation in the leaf, especially at low light intensity.     

Sulphur protects mustard (<Emphasis Type="Italic">Brassica campestris</Emphasis> L.) from cadmium toxicity by improving leaf ascorbate and glutathione

In a pot-soil culture ameliorative effect of sulphur (S) (0 or 40 mg S kg⁻¹ soil) on cadmium (Cd) (0, 25, 50 and 100 mg Cd kg⁻¹ soil)-induced growth inhibition and oxidative stress in mustard (Brassica campestris L.) cultivar Pusa Gold was studied. Cadmium at 100 mg kg⁻¹ soil caused maximum increase in the contents of Cd and thiobarbituric acid reactive substances (TBARS) in leaves. Maximum reductions in growth (plant dry mass, leaf area), chlorophyll content, net photosynthetic rate (P_N) and the contents of ascorbate (AsA), glutathione (GSH) were observed with 100 mg Cd kg⁻¹ soil compared to control. The application of S helped in reducing Cd toxicity, which was greater for 25 and 50 mg Cd kg⁻¹ soil) compared to 100 mg Cd kg⁻¹ soil. Addition of S to Cd-treated plants showed decrease in Cd and TBARS content in leaves and restoration of growth and photosynthesis through increase in the contents of AsA and GSH. Net photosynthetic rate and plant dry mass were strongly and positively correlated with the contents of AsA and GSH. It is suggested that S may ameliorate Cd toxicity and protects growth and photosynthesis of mustard involving AsA and GSH.     

干旱胁迫下镉处理对互叶醉鱼草幼苗生长、镉积累及光合生理的影响

为探究干旱条件下,互叶醉鱼草(Buddleja alternifolia Maxim.)幼苗对重金属镉胁迫的生长及光合生理响应机制,以两年生互叶醉鱼草幼苗为试验材料,设置对照与干旱两个水分处理组(土壤相对含水率分别为:65%—60%,35%—30%),每个水分处理条件下再分别设置3个镉处理浓度(0.28、(0.6+0.28)、(1.2+0.28)mg/kg),共6个处理。测定不同水分及镉处理对互叶醉鱼草生长、生物量、光合参数及体内重金属含量的影响。结果表明:干旱与镉复合胁迫下植物的存活率为100%。镉胁迫、干旱与镉复合胁迫均不同程度抑制了互叶醉鱼草幼苗生长、生物量积累、植株的光合作用及叶绿素含量,且其光合和叶绿素含量的降幅明显大于单一镉胁迫。镉胁迫下,互叶醉鱼草幼苗单株最高镉富集量为69.33 mg/kg,而复合胁迫下单株最高镉富集量为50.68 mg/kg。以上结果表明:干旱胁迫能够加重镉胁迫对植物的影响,使复合胁迫下互叶醉鱼草生长、光合生理及镉富集能力下降。但单一镉胁迫下,互叶醉鱼草对镉具有更强的耐受性,并有较高的生物富集能力,且干旱与Cd复合胁迫下互叶醉鱼草幼苗仍有一定的镉积累量。因此在干旱半干旱区园林绿化以及Cd污染地区的生态建设中,互叶醉鱼草是一种具有巨大应用潜力和前景的灌木树种。     

Soil cadmium toxicity and nitrogen deposition differently affect growth and physiology in <Emphasis Type="Italic">Toxicodendron vernicifluum</Emphasis> seedlings

To ultimately determine whether different levels of soil nitrogen (N) deposition can modify the detrimental effects of cadmium (Cd), the seedlings of Toxicodendron vernicifluum (Strokes) F. A. Barkley were exposed to soil Cd stress (0, 5 and 15 mg kg^?1 dry soil), N deposition (0, 13 and 40 mg kg^?1 dry soil) and their combinations. Soil Cd stress caused damage in plant growth, photosynthesis and other physiological indexes, and in the ultrastructure of mesophyll cells. The effects of N deposition on growth, lipid peroxidation and enzyme activities depended on the relative amounts of N supplied. The combination of low N deposition and Cd stress was positive to plant growth, photosynthesis and enzyme activities, and it caused lower levels of Cd accumulation and lipid peroxidation compared with the effect of Cd stress alone. The combination of high N deposition and Cd stress led to a higher Cd accumulation and lipid peroxidation, and to lower enzyme activities, as compared with the effect of Cd stress alone. T. vernicifluum was found to be sensitive to soil Cd stress. Soil Cd had detrimental effects on T. vernicifluum seedlings, but the tolerance of T. vernicifluum to Cd increased under low N deposition.     

Application of sulfur fertilizer reduces cadmium accumulation and toxicity in tobacco seedlings (<Emphasis Type="Italic">Nicotiana tabacum</Emphasis>)

Greenhouse pot experiments were carried out to illuminate the mitigation effects of sulfur (S) on cadmium (Cd) uptake and toxicity in tobacco using two levels of exogenous sulfur of S1 and S2 containing 47 and 38% total S. Results showed that Cd1 and Cd2 treatments of 1 and 5 mg Cd kg^?1 soil increased leaf Cd concentration and accumulation but reduced plant height, net photosynthetic rate (Pn) and biomass, with a much severe response in Cd2 treatment. Application of S2 fertilizer alleviates Cd toxicity, markedly decreased Cd concentration and improved photosynthesis, compared with Cd1 and Cd2 alone treatment; but S1 only alleviated Cd toxicity when tobacco was subjected to Cd1 stress. Our results suggest that S2 fertilizer was more effective in reducing Cd accumulation and toxicity in Tobacco than S1, and highlights a promising approach of S fertilizer application to lower leaf Cd accumulation in order to ensure product safety of tobacco grown in Cd polluted soils.     

Effects of soil flooding and changes in light intensity on photosynthesis of <Emphasis Type="Italic">Eugenia uniflora</Emphasis> L. seedlings

The increased frequency of heavy rains as a result of global climate change can lead to flooding and changes in light availability caused by the presence of thick clouds. To test the hypothesis that reduction in light availability can alleviate the harmful effects of soil flooding on photosynthesis, the authors studied the effects of soil flooding and acclimation from high to low light on the photosynthetic performance of Eugenia uniflora. Seedlings acclimated to full sunlight (about 35 mol m⁻² d⁻¹) for 5 months were transferred to partial sunlight (about 10 mol m⁻² d⁻¹) and were either subjected to soil flooding or not flooded. Chlorophyll fluorescence was measured throughout the flooding period and leaf gas exchange was measured 16 days after flooding was initiated. Minimal fluorescence yield (Fo) was significantly higher and the quantum efficiency of open PSII centres (Fv/Fm) was significantly lower in flooded than in non-flooded plants in full sunlight. Sixteen days after flooding was initiated, stomatal conductance (gs_sat) and net photosyntheses expressed on a leaf area (A_sat-area), weight (A_sat-wt) and chlorophyll (A_sat-Chl) basis decreased in response to soil flooding. Flooding decreased stomatal conductance by similar amounts in full and partial sunlight, but A_sat-area in partial and full sunlight was 3.4 and 16.8 times lower, respectively, in flooded than in non-flooded plants. These results indicate that changes from full to partial sunlight during soil flooding can alleviate the effects of flooding stress on photosynthesis in E. uniflora seedlings acclimated to full sunlight. The responses of photosynthesis in trees to flooding stress may be dependent on changes in light environment during heavy rains.     

Morphophysiological responses and programmed cell death induced by cadmium in <Emphasis Type="Italic">Genipa americana</Emphasis> L. (Rubiaceae)

Cadmium (Cd) originating from atmospheric deposits, from industrial residues and from the application of phosphate fertilizers may accumulate in high concentrations in soil, water and food, thus becoming highly toxic to plants, animals and human beings. Once accumulated in an organism, Cd discharges and sets off a sequence of biochemical reactions and morphophysiological changes which may cause cell death in several tissues and organs. In order to test the hypothesis that Cd interferes in the metabolism of G. americana, a greenhouse experiment was conducted to measure eventual morphophysiological responses and cell death induced by Cd in this species. The plants were exposed to Cd concentrations ranging from 0 to 16 mg l⁻¹, in a nutritive solution. In TUNEL reaction, it was shown that Cd caused morphological changes in the cell nucleus of root tip and leaf tissues, which are typical for apoptosis. Cadmium induced anatomical changes in roots and leaves, such as the lignification of cell walls in root tissues and leaf main vein. In addition, the leaf mesophyll showed increase of the intercellular spaces. On the other hand, Cd caused reductions in the net photosynthetic rate, stomatal conductance and leaf transpiration, while the maximum potential quantum efficiency of PS2 (Fv/Fm) was unchanged. Cadmium accumulated in the root system in high concentrations, with low translocation for the shoot, and promoted an increase of Ca and Zn levels in the roots and a decrease of K level in the leaves. High concentrations of Cd promoted morphophysiological changes and caused cell death in roots and leaves tissues of G. americana.     

Does salt stress constrain spatial distribution of dune building grasses Ammophila arenaria and Elytrichia juncea on the beach?

Rising sea levels threaten coastal safety by increasing the risk of flooding. Coastal dunes provide a natural form of coastal protection. Understanding drivers that constrain early development of dunes is necessary to assess whether dune development may keep pace with sea‐level rise. In this study, we explored to what extent salt stress experienced by dune building plant species constrains their spatial distribution at the Dutch sandy coast. We conducted a field transplantation experiment and a glasshouse experiment with two dune building grasses Ammophila arenaria and Elytrigia juncea. In the field, we measured salinity and monitored growth of transplanted grasses in four vegetation zones: (I) nonvegetated beach, (II) E. juncea occurring, (III) both species co‐occurring, and (IV) A. arenaria dominant. In the glasshouse, we subjected the two species to six soil salinity treatments, with and without salt spray. We monitored biomass, photosynthesis, leaf sodium, and nutrient concentrations over a growing season. The vegetation zones were weakly associated with summer soil salinity; zone I and II were significantly more saline than zones III and IV. Ammophila arenaria performed equally (zone II) or better (zones III, IV) than E. juncea, suggesting soil salinity did not limit species performance. Both species showed severe winter mortality. In the glasshouse, A. arenaria biomass decreased linearly with soil salinity, presumably as a result of osmotic stress. Elytrigia juncea showed a nonlinear response to soil salinity with an optimum at 0.75% soil salinity. Our findings suggest that soil salinity stress either takes place in winter, or that development of vegetated dunes is less sensitive to soil salinity than hitherto expected.     

Melatonin mitigates cadmium and aluminium toxicity through modulation of antioxidant potential in Brassica napus L

• Melatonin has emerged as an essential molecule in plants, due to its role in defence against metal toxicity. Aluminium (Al) and cadmium (Cd) toxicity inhibit rapeseed seedling growth.
• In this study, we applied different doses of melatonin (50 and 100 µm ) to alleviate Al (25 µm ) and Cd (25 µm ) stress in rapeseed seedlings. Results show that Al and Cd caused toxicity in rapeseed seedling, as evidenced by a decrease in height, biomass and antioxidant enzyme activity.
• Melatonin increased the expression of melatonin biosynthesis‐related Brassica napus genes for caffeic acid O‐methyl transferase (BnCOMT) under Al and Cd stress. The genes BnCOMT‐1, BnCOMT‐5 and BnCOMT‐8 showed up‐regulated expression, while BnCOMT‐4 and BnCOMT‐6 were down‐regulated during incubation in water. Melatonin application increased the germination rate, shoot length, root length, fresh and dry weight of seedlings. Melatonin supplementation under Al and Cd stress increased superoxide dismutase, catalase, peroxidase, ascorbate peroxidase, proline, chlorophyll and anthocyanin content, as well as photosynthesis rate. Both Cd and Al treatments significantly increased hydrogen peroxide and malondialdehyde levels in rapeseed seedlings, which were strictly counterbalanced by melatonin. Analysis of Cd and Al in different subcellular compartments showed that melatonin enhanced cell wall and soluble fractions, but reduced the vacuolar and organelle fractions in Al‐ and Cd‐treated seedlings.
• These results suggest that melatonin‐induced improvements in antioxidant potential, biomass, photosynthesis rate and successive Cd and Al sequestration play a pivotal role in plant tolerance to Al and Cd stress. This mechanism may have potential implications in safe food production.