Effects of ozone on growth, yield and leaf gas exchange rates of four Bangladeshi cultivars of rice (Oryza sativa L.)

To assess the effects of tropospheric O₃ on rice cultivated in Bangladesh, four Bangladeshi cultivars (BR11, BR14, BR28 and BR29) of rice (Oryza sativa L.) were exposed daily to charcoal-filtered air or O₃ at 60 and 100 nl l⁻¹ (10:00-17:00) from 1 July to 28 November 2008. The whole-plant dry mass and grain yield per plant of the four cultivars were significantly reduced by the exposure to O₃. The exposure to O₃ significantly reduced net photosynthetic rate of the 12th and flag leaves of the four cultivars. The sensitivity to O₃ of growth, yield and leaf gas exchange rates was not significantly different among the four cultivars. The present study suggests that the sensitivity to O₃ of yield of the four Bangladeshi rice cultivars is greater than that of American rice cultivars and is similar to that of Japanese rice cultivars and that O₃ may detrimentally affect rice production in Bangladesh.     

Establishing ozone flux–response relationships for winter wheat: Analysis of uncertainties based on data for UK and Polish genotypes

The work outlined in this paper had three objectives. The first was to explore the effects of ozone pollution on grain yield and quality of commercially-grown winter wheat cultivars. The second was to derive a stomatal ozone flux model for winter wheat and compare with those already developed for spring wheat. The third was to evaluate exposure- versus flux–response approaches from a risk assessment perspective, and explore the implications of genetic variation in modelled ozone flux.Fifteen winter wheat cultivars were grown in open-top chambers where they were exposed to four levels of ozone. During fumigation, stomatal conductance measurements were made over the lifespan of the flag leaf across a range of environmental conditions. Although significant intra-specific variation in ‘ozone sensitivity’ (in terms of impacts on yield) was identified, yield was inversely related (R² = 0.63, P < 0.001) to the accumulated hourly averaged ozone exposure above 40 ppb during daylight hours (AOT40) across the dataset. The adverse effect of ozone on yield was principally due to a decline in seed weight. Algorithms defining the influence of environmental variables on stomatal uptake were subtly different from those currently in use, based on data for spring wheat, to map ozone impacts on pan-European cereal yield. Considerable intra-specific variation in phenological effects was identified. This meant that an ‘average behaviour’ had to be derived which reduced the predictive capability of the derived stomatal flux model (R² = 0.49, P < 0.001, 15 cultivars included). Indeed, given the intra-specific variability encountered, the flux model that was derived from the full dataset was no better in predicting O₃ impacts on wheat yield than was the AOT40 index. The study highlights the need to use ozone risk assessment tools appropriate to specific vegetation types when modelling and mapping ozone impacts at the regional level.     

Differential protection of ethylenediurea (EDU) against ambient ozone for five cultivars of tropical wheat

The antiozonant EDU (ethylenediurea) was used to assess the impact of ambient O₃ under field conditions on five cultivars of tropical wheat (Triticum aestivum L.). EDU solution (0 ppm and 400 ppm) was applied as soil drench (100 ml plant^?1) 10 days after germination (DAG) at an interval of 12 days. EDU-treated plants showed significant increments in stomatal conductance, photosynthetic rate, variable fluorescence, total chlorophyll, ascorbic acid, proline and protein contents and protective enzymes (POX, SOD and APX) activities in HUW468, HUW510 and HUW234 cultivars, while, a reverse trend was observed for lipid peroxidation. EDU application restored grain yield significantly by maintaining higher levels of antioxidants, metabolites and enzymes in cultivars HUW468 and HUW510. Sonalika and PBW343 showed least response of measured parameters under EDU treatment suggesting their greater resistance to O₃. EDU, thus proved its usefulness in screening suitable wheat cultivars for areas experiencing elevated concentrations of O₃.     

Reduced ozone by air filtration consistently improved grain yield in wheat

This study considered effects of reduced [O₃] on wheat yield. Open-top chamber charcoal filtered air treatments were compared with non-filtered treatments for field-grown wheat. 30 experiments meeting requirements were found, representing nine countries in North America, Europe and Asia. 26 experiments reported improved yield and 4 experiments reduced yield by filtration, a significant positive effect. Average yield improvement was 9%. Average daytime [O₃] was reduced by filtration from 35 to 13 nmol mol⁻¹. Filtration efficiency was 63% for O₃ and 56% for SO₂. For NO_x it was observed that NO₂ was reduced and NO increased by filtration. Thus, filters convert NO₂ to NO. Most experiments reported low or very low [SO₂] and [NO_x]. Thus, O₃ can be concluded to be the main phytotoxic component in the experiments. Elevated [NO₂] was observed in one experiment. The conclusion is that current [O₃] over large parts of the world adversely affect wheat yield.     

Assessing the impacts of current and future concentrations of surface ozone on crop yield with meta-analysis

Meta-analysis was conducted to quantitatively assess the effects of rising ozone concentrations ([O₃]) on yield and yield components of major food crops: potato, barley, wheat, rice, bean and soybean in 406 experimental observations. Yield loss of the crops under current and future [O₃] was expressed relative to the yield under base [O₃] (≤26 ppb). With potato, current [O₃] (31–50 ppb) reduced the yield by 5.3%, and it reduced the yield of barley, wheat and rice by 8.9%, 9.7% and 17.5%, respectively. In bean and soybean, the yield losses were 19.0% and 7.7%, respectively. Compared with yield loss at current [O₃], future [O₃] (51–75 ppb) drove a further 10% loss in yield of soybean, wheat and rice, and 20% loss in bean. Mass of individual grain, seed, or tuber was often the major cause of the yield loss at current and future [O₃], whereas other yield components also contributed to the yield loss in some cases. No significant difference was found between the responses in crops grown in pots and those in the ground for any yield parameters. The ameliorating effect of elevated [CO₂] was significant in the yields of wheat and potato, and the individual grain weight in wheat exposed to future [O₃]. These findings confirm the rising [O₃] as a threat to food security for the growing global population in this century.     

Assessment of competitive ability of two Indian wheat cultivars under ambient O3 at different developmental stages

The concentrations of O₃ are increasing, which may have potential adverse effects on crop yield. This paper deals with assessing the intraspecific variability of two wheat cultivars (PBW 343 and M 533) at different growth stages using open top chambers. Mean O₃ concentrations were 50.2 and 53.2 ppb, and AOT40 values were 9 and 12.1 ppm h, respectively, in 2008–2009 and 2009–2010. Reproductive stage showed higher AOT40 values (6.9 and 9.2 ppm h) compared to vegetative (2.23 and 2.9 ppm h). Critical levels of a 3-month AOT 40 of 3 ppm h led to 6 % yield reduction in two wheat cultivars for two consecutive years. Variations in photosynthesis rate, stomatal conductance (g_s), Fv/Fm ratio, photosynthetic pigments, primary and secondary metabolites, morphological parameters, and yield attributes were measured at vegetative and reproductive stages. Reductions in number of leaves, leaf area, total biomass, root/shoot ratio, RGR, photosynthetic pigments, protein content, and Fv/Fm ratio in PBW 343 were more than M 533 at reproductive stage. Photosynthetic rate did not vary between the cultivars, but g_s was higher in PBW 343 compared to M 533 under ambient O₃. Higher total phenolics and peroxidase activity were recorded in M 533 at reproductive stage conferring higher resistance at latter age. Results of O₃ resistance showed that M 533 was sensitive compared to PBW 343 during vegetative stage but developed more resistance at reproductive stage. PBW 343 with larger leaf area and high g_s is more sensitive than M 533 with smaller leaf area and low g_s. The study suggests that the sensitivity varied with plant growth stage, and the plant showing higher sensitivity during vegetative period developed more resistance during reproductive period due to higher defense mechanism. Though the yield reductions were same in both cultivars under ambient O₃, the mechanism of acquiring the resistance is different between the cultivars.     

An Air Quality Data Analysis System for Interrelating Effects,Standards, and Needed Source Reductions: Part 8. An Effective Mean O3 Crop Reduction Mathematical Model

A plant injury mathematical model, applied previously to acute and chronic leaf injury data, is used here to model National Crop Loss Assessment Network (NCLAN) data for 15 cultivars and to calculate species parameters from the cultivar analyses. Percent crop yield reduction is estimated as a function of a new parameter, the effective mean O₃ concentration: m_e = [(Σ c_h ^?1/v)/n]^?v, where c_h is the hourly average ambient O₃ concentration for each daytime hour (defined here as 9:00 A.M.–4:00 P.M., always standard time) of data available at an air sampling site for summer (defined here as June 1–August 31), n is the total number of such available hours, and v is an exposure time-concentration parameter, calculated here to be approximately –0.376. Crop yield reduction for soybean is calculated here as z = 0.478 In (tm_e ^2-66) – 0.42, where z is the Gaussian transform of percent crop reduction, t is the hours of exposure (525 h is used here; 7 h/day for 75 days), and In indicates that the natural logarithm is taken of the quantity within parentheses. Crop yield reductions for seven plant species are estimated with similar equations for each of the 1824 site-years of 1981–1983 hourly O₃ concentration data available in the National Aerometric Data Bank (NADB). County-average effective mean O₃ concentrations are indicated by shading on a U.S. map. State-average O₃ parameters and estimated percent crop yield reductions are tabulated. The National Ambient Air Quality Standard (NAAQS) for O₃ specifies that, on the average, the second highest daily maximum 1-h average O₃ concentration in a year shall not exceed 0.12 ppm. For years 1981-1983,71% of the NADB sites recorded annual second highest daily maximum 1-h average O3 concentrations below 0.125 ppm (for summer daytime hours). Ambient O₃ concentrations reduced the total U.S. crop yield an estimated 5% for years 1981–1983. (Summer, daytime, and all acronyms are always used herein as defined above.)     

A comparison of North American and Asian exposure–response data for ozone effects on crop yields

Modelling-based studies to assess the extent and magnitude of ozone (O₃) risk to agriculture in Asia suggest that yield losses of 5–20% for important crops may be common in areas experiencing elevated O₃ concentrations. These assessments have relied on European and North American dose–response relationships and hence assumed an equivalent Asian crop response to O₃ for local cultivars, pollutant conditions and climate. To test this assumption we collated comparable dose–response data derived from fumigation, filtration and EDU experiments conducted in Asia on wheat, rice and leguminous crop species. These data are pooled and compared with equivalent North American dose–response relationships. The Asian data show that at ambient O₃ concentrations found at the study sites (which vary between ～35–75 ppb 4–8 h growing season mean), yield losses for wheat, rice and legumes range between 5–48, 3–47 and 10–65%, respectively. The results indicate that Asian grown wheat and rice cultivars are more sensitive to O₃ than the North American dose–response relationships would suggest. For legumes the scatter in the data makes it difficult to reach any equivalent conclusion in relative sensitivities. As such, existing modelling-based risk assessments may have substantially underestimated the scale of the problem in Asia through use of North American derived dose–response relationships.     

Leaf size and surface characteristics of Betula papyrifera exposed to elevated CO2 and O3

Betula papyrifera trees were exposed to elevated concentrations of CO₂ (1.4 × ambient), O₃ (1.2 × ambient) or CO₂ + O₃ at the Aspen Free-air CO₂ Enrichment Experiment. The treatment effects on leaf surface characteristics were studied after nine years of tree exposure. CO₂ and O₃ increased epidermal cell size and reduced epidermal cell density but leaf size was not altered. Stomatal density remained unaffected, but stomatal index increased under elevated CO₂. Cuticular ridges and epicuticular wax crystallites were less evident under CO₂ and CO₂ + O₃. The increase in amorphous deposits, particularly under CO₂ + O_3, was associated with the appearance of elongated plate crystallites in stomatal chambers. Increased proportions of alkyl esters resulted from increased esterification of fatty acids and alcohols under elevated CO₂ + O₃. The combination of elevated CO₂ and O₃ resulted in different responses than expected under exposure to CO₂ or O₃ alone.     

Ozone exposure of field-grown winter wheat affects soil mesofauna in the rhizosphere

A 2-year open-top chamber experiment with field-grown winter wheat (Triticum aestivum L. cv. Astron) was conducted to examine the effects of ozone on plant growth and selected groups of soil mesofauna in the rhizosphere. From May through June in each year, plants were exposed to two levels of O₃: non-filtered (NF) ambient air or NF+ 40 ppb O₃ (NF+). During O₃ exposure, soil sampling was performed at two dates according to different plant growth stages. O₃ exposure reduced above- and below-ground plant biomass in the first year, but had little effect in the second year. The individual density of enchytraeids, collembolans and soil mites decreased significantly in the rhizosphere of plants exposed to NF+ in both years. Differences were highest around anthesis, i.e. when plants are physiologically most active. The results suggest that elevated O₃ concentrations may influence the dynamic of decomposition processes and the turnover of nutrients.     

Responses to ozone pollution of alfalfa exposed to increasing salinity levels

Stomatal closure and biosynthesis of antioxidant molecules are two fundamental components of the physiological machinery that lead to stress adaptation during plant's exposure to salinity. Since high stomatal resistance may also contribute in counteracting O₃ damages, we hypothesized that soil salinization may increase O₃ tolerance of crops. An experiment was performed with alfalfa grown in filtered (AOT40 = 0 in both years) and non-filtered (AOT40 = 9.7 in 2005 and 6.9 ppm h in 2006) open-top chambers. Alfalfa yield was reduced by O₃ (−33%) only in plants irrigated with salt-free water, while the increasing levels of soil salinity until 1.06 dS m⁻¹ reduced both stomatal conductance and plant O₃ uptake, thus linearly reducing O₃ effects on yield. Therefore a reliable flux-based model for assessing the effects of O₃ on crop yield should take into account soil salinity.     

Ozone, nitric acid, and ammonia air pollution is unhealthy for people and ecosystems in southern Sierra Nevada, California

Two-week average concentrations of ozone (O₃), nitric acid vapor (HNO₃) and ammonia (NH₃) were measured with passive samplers during the 2002 summer season across the central Sierra Nevada Mountains, California, along the San Joaquin River drainage. Elevated concentrations of the pollutants were determined with seasonal means for individual sites ranging between 62 and 88 ppb for O₃, 1.0-3.8 μg m⁻³ for HNO₃, and 2.6-5.2 μg m⁻³ for NH₃. Calculated O₃ exposure indices were very high, reaching SUM00-191 ppm h, SUM60-151 ppm h, and W126-124 ppm h. Calculated nitrogen (N) dry deposition ranged from 1.4 to 15 kg N ha⁻¹ for maximum values, and 0.4-8 kg N ha⁻¹ for minimum values; potentially exceeding Critical Loads (CL) for nutritional N. The U.S., California, and European 8 h O₃ human health standards were exceeded during 104, 108, and 114 days respectively, indicating high risk to humans from ambient O₃.     

Contrasting effects of ozone under different water supplies in two Mediterranean tree species

The effects of ozone (O₃) exposure under different water availabilities were studied in two Mediterranean tree species: Quercus ilex and Ceratonia siliqua. Plants were exposed to different O₃ concentrations in open top chambers (charcoal-filtered air (CF), non-filtered air (NF)) and non-filtered air plus 40 ppb_v of O₃ ((7:00–17:00 solar time) (NF+)) during 2 years, and to different water regimes (IR, sample irrigation, and WS, reduced water dose to 50%) through the last of those 2 years. AOT40 in the NF+ treatment was 59265 ppb_v h (from March 1999 to August 1999) while in the NF treatment, the AOT40 was 6727 ppb_v h for the same period. AOT40 was always 0 in the CF treatment. WS plants presented lower stomatal conductances and net photosynthetic rates, and higher foliar N concentrations than IR plants in both species. The irrigation treatment did not change the response trends to ozone in Q. ilex, the most sensitive species to O₃ ambient concentrations, but it changed those of C. siliqua, the least sensitive species, since its ozone-fumigated WS plants did not decrease their net photosynthetic rates nor their biomass accumulation as it happened to its ozone-fumigated IR plants. These results show interspecific variations in O₃ sensitivity under different water availabilities.     

Estimated crop yield losses due to surface ozone exposure and economic damage in India

In this study, we estimate yield losses and economic damage of two major crops (winter wheat and rabi rice) due to surface ozone (O₃) exposure using hourly O₃ concentrations for the period 2002–2007 in India. This study estimates crop yield losses according to two indices of O₃ exposure: 7-h seasonal daytime (0900–1600 hours) mean measured O₃ concentration (M7) and AOT40 (accumulation exposure of O₃ concentration over a threshold of 40 parts per billion by volume during daylight hours (0700–1800 hours), established by field studies. Our results indicate that relative yield loss from 5 to 11 % (6–30 %) for winter wheat and 3–6 % (9–16 %) for rabi rice using M7 (AOT40) index of the mean total winter wheat 81 million metric tons (Mt) and rabi rice 12 Mt production per year for the period 2002–2007. The estimated mean crop production loss (CPL) for winter wheat are from 9 to 29 Mt, account for economic cost loss was from 1,222 to 4,091 million US$ annually. Similarly, the mean CPL for rabi rice are from 0.64 to 2.1 Mt, worth 86–276 million US$. Our calculated winter wheat and rabi rice losses agree well with previous results, providing the further evidence that large crop yield losses occurring in India due to current O₃ concentration and further elevated O₃ concentration in future may pose threat to food security.     

Ozone risk assessment for agricultural crops in Europe: Further development of stomatal flux and flux–response relationships for European wheat and potato

Applications of a parameterised Jarvis-type multiplicative stomatal conductance model with data collated from open-top chamber experiments on field grown wheat and potato were used to derive relationships between relative yield and stomatal ozone uptake. The relationships were based on thirteen experiments from four European countries for wheat and seven experiments from four European countries for potato. The parameterisation of the conductance model was based both on an extensive literature review and primary data. Application of the stomatal conductance models to the open-top chamber experiments resulted in improved linear regressions between relative yield and ozone uptake compared to earlier stomatal conductance models, both for wheat (r²=0.83) and potato (r²=0.76). The improvement was largest for potato. The relationships with the highest correlation were obtained using a stomatal ozone flux threshold. For both wheat and potato the best performing exposure index was AF_st6 (accumulated stomatal flux of ozone above a flux rate threshold of 6 nmol ozone m⁻² projected sunlit leaf area, based on hourly values of ozone flux). The results demonstrate that flux-based models are now sufficiently well calibrated to be used with confidence to predict the effects of ozone on yield loss of major arable crops across Europe. Further studies, using innovations in stomatal conductance modelling and plant exposure experimentation, are needed if these models are to be further improved.     

Leaf age affects the responses of foliar injury and gas exchange to tropospheric ozone in Prunus serotina seedlings

We investigated the effect of leaf age on the response of net photosynthesis (A), stomatal conductance (g_wv), foliar injury, and leaf nitrogen concentration (N_L) to tropospheric ozone (O₃) on Prunus serotina seedlings grown in open-plots (AA) and open-top chambers, supplied with either carbon-filtered or non-filtered air. We found significant variation in A, g_wv, foliar injury, and N_L (P < 0.05) among O₃ treatments. Seedlings in AA showed the highest A and g_wv due to relatively low vapor pressure deficit (VPD). Older leaves showed significantly lower A, g_wv, N_L, and higher foliar injury (P < 0.001) than younger leaves. Leaf age affected the response of A, g_wv, and foliar injury to O₃. Both VPD and N_L had a strong influence on leaf gas exchange. Foliar O₃-induced injury appeared when cumulative O₃ uptake reached 8-12 mmol m⁻², depending on soil water availability. The mechanistic assessment of O₃-induced injury is a valuable approach for a biologically relevant O₃ risk assessment for forest trees.     

Ozone sensitivity of the Mediterranean terophyte Trifolium striatum is modulated by soil nitrogen content

The influence of nitrogen (N) fertilization on the sensitivity to ozone (O₃) has been studied using the O₃ sensitive species Trifolium striatum, an annual species frequently found in therophitic grasslands of the dehesas in the central Iberian Peninsula. The experiment was carried out in open-top chambers using three different O₃ exposure treatments: charcoal-filtered air (CFA), non-filtered air (NFA) and non-filtered air plus 40 nl l⁻¹ added over ambient concentrations (NFA+). After 30 days of exposure to the different treatments, plants were placed under ambient conditions until seed ripeness was completed. Three N fertilization treatments were established receiving final doses of 10 (low), 30 (medium) or 60 (high) kg N ha⁻¹ throughout the growing period divided into four proportional doses at every 15 days. Ozone affected phenology and reduced total aerial and flower biomass and seed production in NFA and NFA+ treatments. Carry-over effects of O₃ on the reproductive ability of this species were found resulting in reductions in flower biomass and seed production detected 1 month after O₃ exposure. By contrast, N fertilization increased aerial and flower biomass. Significant O₃×N interactive effects were detected since N fertilization counterbalanced O₃-induced effects only when plants were exposed to moderate O₃ levels (NFA) but not under high O₃ concentrations (NFA+). The observed O₃ effects on reproductive ability of T. striatum might determine changes in species viability and future diversity of the ecosystem.     

Apoplastic ascorbate contributes to the differential ozone sensitivity in two varieties of winter wheat under fully open-air field conditions

We studied leaf apoplastic ascorbates in relation to ozone (O₃) sensitivity in two winter wheat (Triticum aestivum L.) varieties: Yangfumai 2 (Y2) and Yangmai 16 (Y16). The plants were exposed to elevated O₃ concentration 27% higher than the ambient O₃ concentration in a fully open-air field from tillering stage until final maturity. The less sensitive variety Y16 had higher concentration of reduced ascorbate in the apoplast and leaf tissue by 33.5% and 12.0%, respectively, than those in the more sensitive variety Y2, whereas no varietal difference was detected in the decline of reduced ascorbate concentration in response to elevated O₃. No effects of O₃ or variety were detected in either oxidized ascorbate or the redox state of ascorbate in the apoplast and leaf tissue. The lower ascorbate concentrations in both apoplast and leaf tissue should have contributed to the higher O₃ sensitivity in variety Y2.     

A flux-based assessment of the effects of ozone on foliar injury, photosynthesis, and yield of bean (Phaseolus vulgaris L. cv. Borlotto Nano Lingua di Fuoco) in open-top chambers

Stomatal ozone uptake, determined with the Jarvis' approach, was related to photosynthetic efficiency assessed by chlorophyll fluorescence and reflectance measurements in open-top chamber experiments on Phaseolus vulgaris. The effects of O₃ exposure were also evaluated in terms of visible and microscopical leaf injury and plant productivity. Results showed that microscopical leaf symptoms, assessed as cell death and H₂O₂ accumulation, preceded by 3-4 days the appearance of visible symptoms. An effective dose of ozone stomatal flux for visible leaf damages was found around 1.33 mmol O₃ m⁻². Significant linear dose-response relationships were obtained between accumulated fluxes and optical indices (PRI, NDI, ΔF/F_m^′). The negative effects on photosynthesis reduced plant productivity, affecting the number of pods and seeds, but not seed weight. These results, besides contributing to the development of a flux-based ozone risk assessment for crops in Europe, highlight the potentiality of reflectance measurements for the early detection of ozone stress.     

Threat to food security under current levels of ground level ozone: A case study for Indian cultivars of rice

A higher ozone concentration in rural agricultural region poses threat to food production in developing countries. The present study was conducted to evaluate the growth, biomass accumulation and allocation pattern, quantitative and qualitative characteristics of grains for two tropical rice cultivars (Oryza sativa L. cv NDR 97 and Saurabh 950) at ambient O₃ concentrations at a rural site in the Indo Gangetic plains of India.Percent inhibition in number of leaves was higher for NDR 97, but in leaf area for Saurabh 950 grown in non filtered chambers (NFCs) compared to filtered chambers (FCs). Higher inhibition in root biomass was recorded in Saurabh 950 and in leaf and standing dead biomass for NDR 97. During vegetative phase, relative growth rate showed more percent inhibition in Saurabh 950, but at reproductive phase in NDR 97. Net assimilation rate showed higher values for Saurabh 950 than NDR 97 in NFCs but percent inhibition in leaf area ratio was higher for former than latter cultivar in NFCs. The ozone resistance was higher in NDR 97 during vegetative phase, but in Saurabh 950 at reproductive phase. Number of grains was higher in NDR 97 than Saurabh 950, but test weight and weight of grains m^?2 showed reverse trends. Concentrations of starch, protein, P, N, Ca, Mg and K decreased, while reducing and total soluble sugar increased in grains of both the cultivars in NFCs compared to FCs. The study concluded that under ambient condition of O₃ exposure, the two cultivars responded differently. Saurabh 950 favoured biomass translocation priority towards ear in reproductive phase and hence showed higher resistivity due to maintenance of higher test weight. NDR 97, however, showed better growth during vegetative period, but could not allocate efficiently to developing ears, hence higher number of unfilled grains in NFCs led lower test weight.