Effect of SO2 on the reproduction of pea aphids, Acyrthosiphon pisum, and the impact of SO2 and aphids on the growth and yield of peas

Pea aphids feeding from birth to maturity on pea plants (Pisum sativum) exposed to SO(2) concentrations of 50 nl litre(-1) or 80 nl litre(-1) showed a significant 19% increase in the rate of nymph production during the reproductive period, compared to control aphids feeding on plants in charcoal-filtered air. The higher nymph production resulted in a mean 4.6% increase in the intrinsic rate of population increase (rm). In longer term glasshouse fumigation experiments pea aphid populations were, on average, 1.8 times greater on pea plants in ambient air plus 45 nl litre(-1) SO(2) than in ambient air alone. Aphid infestation in ambient air caused a 42% reduction in pea yield and affected most plant parameters adversely. Ambient air plus SO(2) had no direct effect on yield, but, in combination with aphid infestation, a further 10% reduction in yield was recorded.     

Air pollution and agricultural aphid pests. I: Fumigation experiments with SO2 and NO2

A wide range of aphid/host-crop systems was surveyed by means of fumigation experiments in closed chambers for sensitivity to SO(2) and NO(2) at a concentration of 100 nl litre(-1). Aphid performance was measured by the mean relative growth rate (MRGR) of individual aphids. In all cases, except for Acrythosiphon pisum (Harris) on Vicia fabaL., there were increases in the MRGR of the aphids feeding on fumigated plants as compared to clean air controls, both during and post fumigation. The increases in MRGR ranged from 6 to 75%, with the majority falling between 25 and 40%. A. pisum on V. faba showed a consistent negative response, with decreases in MRGR between -9 and -12%. The changes in aohid MRGR were not due to direct effects, as no significant differences in MRGR were observed between fumigated and clean air chambers when aphids were fed on artificial diet sachets during fumigation.     

Relationship between SO2 dose and growth of the pea aphid, Acyrthosiphon pisum, on peas

A series of experiments was carried out in controlled environment cabinets to investigate the effects of SO(2) dose on the mean relative growth rate (MRGR) of pea aphids, Acyrthosiphon pisum, feeding on the pea plant, Pisum sativum. There was a significant linear increase in the MRGR of aphids feeding on SO(2)-fumigated plants, relative to control aphids feeding on plants in charcoal-filtered air. The increase in MRGR reached a maximum of 11% at SO(2) concentrations between 90 and 110 nl litre(-1). MRGR declined at higher SO(2) concentrations until above 220 nl litre(-1) it was below that of the controls. The dose-response curve is discussed in relation to reported changes in the nitrogen metabolism of plants subject to air pollution, the response of aphids to host plant nitrogen and possible toxic effects of high concentrations of SO(2) on the aphid.     

An open-top chamber study with filtered and non-filtered air to evaluate the effects of air pollutants on crops

Four non-filtered and four charcoal-filtered open-top chambers were employed to determine the effects of ambient levels of gaseous air pollutants at Braunschweig, FRG, on growth and yield of potted plants of winter and spring barley. During the exposure period (November 1985-August 1986) monthly mean values of gaseous air pollutants (microg m(-3)) ranged between 34 and 127 for SO(2), 34 and 52 for NO(2) and 12 and 33 for O(3) in winter (November-March), and 16 to 26 for SO(2), 20 to 33 for NO(2) and 42 to 53 for O(3) in spring-summer (April-August). Monthly 2% percentile values for these gases reached (microg m (-3)) 561 for SO(2), 140 for NO(2) and 170 for O(3). The filtering efficiencies of the charcoal filters used averaged 60% for SO(2), 50% for NO(2) and 70% for O(3). All plants of winter barley from the unchambered plot were killed by severe frost periods in winter, 1986. Little frost damage occurred on plants grown in the chambers. Air filtration resulted in higher numbers of plants of winter barley per pot, i.e. a higher number of individuals per area, and a higher dry weight of whole plants and ears compared to the non-filtered atmosphere. In the experiments with spring barley, fresh and dry weight of whole plants were lower and dry weight of leaves were higher in the filtered open-top chambers. These effects could not be observed at all harvests which were carried out during the growing season. Grain yield and sulphur content of the leaves of both barley cultivars were not affected by the air filtration. Production of biomass of spring barley grown in ambient air was higher than of that grown in open-top chambers.     

Growth and yield responses of spring wheat (Triticum aestivum L. CV. Turbo) grown in open-top chambers to ozone and water stress

Spring wheat (Triticum aestivum L.) cv. Turbo was exposed to different levels of ozone and water supply in open-top chambers in 1991. The plants were grown either in charcoal filtered air (CF), not filtered air (NF), in charcoal filtered air with proportional addition of ambient ozone (CF1), or in charcoal filtered air with twice proportional addition of ambient ozone (CF2). The mean seasonal ozone concentrations (24 h mean) were 2.3, 20.6, 17.3, and 24.5 nl litre(-1) for CF, NF, CF1, and CF2 treatments, respectively. Ozone enhanced senescence and reduced growth and yield of the wheat plants. At final harvest, dry weight reductions were mainly due to reductions in ear weight. Grain yield loss by ozone mainly resulted from depressions of 1000 grain weight, whereas numbers of ears per plant and of grains per ear remained unchanged. Pollutants other than ozone did not alter the response to ozone, as was obvious from comparisons between CF1 and NF responses. Water stress alone did not enhance senescence, but also reduced growth and yield. However, yield loss mainly resulted from reductions in the number of ears per plant; 1000 grain weight was not influenced by water stress. No water supply by ozone treatment interactions were detected for any of the estimated parameters.     

Root and shoot growth, assimilate partitioning and cell proliferation in roots of Sitka spruce (Picea sitchensis) grown in filtered and unfiltered chambers

Rooted cuttings of clonal Sitka spruce (Picea sitchensis (Bong.) Carr.) were grown from April to October in 1 m long tubes sunk into the ground inside open top chambers. The same experiment was repeated in each of two consecutive years using a different clone of Sitka spruce each year. Air was either passed directly into the chambers (ambient air) or passed over charcoal filters which removed the majority of gaseous pollutants before entering the chambers (filtered air). Ambient pollution did not appear to influence the growth of Sitka spruce at least over the experimental period used. No significant differences were found between plants exposed to ambient or filtered air in terms of shoot and root dry mass, needle dry mass, root length, carbohydrate content of roots and needles, and in the percentage of meristematic cells close to the apex in each phase or interphase or undergoing mitosis.     

Effects of ozone and soil water deficit on roots and shoots of field-grown soybeans

Water-stressed and well-watered soybean (Glycine max cvs. Williams and Corsoy) plants were exposed to increasing seasonal doses of ozone (O(3)) using open-top field chambers and ambient air plots. Chamber O(3) treatments included charcoal filtered (CF) air, non-filtered (NF) air, NF + 0.03, NF + 0.06 and NF + 0.09 microl litre(-1) O(3). Soil water potentials measured at 25 and 45 cm averaged -0.40 MPa and -0.05 MPa, respectively, for the plots in the water-stressed and well-watered series. Total root length/core, root length densities, and biomasses (dry weights) were determined. With Williams, a very popular cultivar in recent years, total root length for all O(3) treatments averaged 58% more under water-stress conditions than in well-watered plots, but the range was from 136% to 11% more for NF air and NF + 0.09 microl litre(-1) O(3), respectively. Increasing the O(3) exposure dose did not affect root lengths or weights in the well-watered series. With Corsoy, water stress did not significantly increase root development. In both soil moisture regimes, with both cultivars, there was a linear decrease in seed yield and top dry weight as the O(3) exposure dose increased.     

Chronic ozone exposure alters the growth of leaves, stems and roots of hybrid Populus

Rooted cuttings of hybrid Populus (DN34, Populus deltoides x nigra) were grown outdoors in pots in open-top chambers at Ithaca, NY (74.5 degrees W, 42.5 degrees N), during 1988 and 1989 (experiment 1) and during 1989 and 1990 (experiment 2). Ambient air was passed through charcoal filters to produce a 0.5 times ambient ozone treatment, and ozone generated from oxygen was added to produce one and two times ambient ozone treatments. In experiment 1, treatments were applied for 8-12 h each day for 112 days of the 1988 growing season, then the plants were grown outdoors with ambient ozone in 1989. In experiment 2, treatments were applied for 9 h each day for 98 days of the 1989 growing season, then the plants were grown outdoors with ambient ozone in 1990. Chronic exposure to ozone caused the following changes (statistically significant in one or both experiments at p<0.05): (1) earlier leaf abscission, (2) decreased stem basal diameter, (3) decreased stem mass, (4) decreased internode length, (5) decreased shoot height p=0.005, and (6) decreased leaf size in the growing season following ozone treatment. There was also strong evidence that ozone increased the number of leaves produced p=0.055. Finally, there was some evidence that ozone increased the ratio of shoot mass to root mass p=0.093.     

A cumulative ozone uptake-response relationship for the growth of Norway spruce saplings

Norway spruce saplings [Picea abies (L.) Karst.] were exposed during four growing seasons to different ozone treatments in open-top chambers: charcoal filtered air (CF), non-filtered air (NF) and non-filtered air with extra ozone (NF+, 1.4xambient concentrations). The CF and NF+ ozone treatments were combined with phosphorous deficiency and drought stress treatments. The total biomass of the trees was harvested at different intervals during the experimental period. The ozone uptake to current-year needles of the Norway spruce saplings was estimated using a multiplicative stomatal conductance simulation model. There was a highly significant correlation between the reduction of total biomass and the estimated cumulative ozone uptake, which did not vary when different thresholds were applied for the rate of ozone uptake. The reduction of the total biomass was estimated to 1% per 10 mmol m(-2) cumulated ozone uptake, on a projected needle area basis.     

The response of spruce shoot aphid Cinara pilicornis hartig to ambient and filtered air at two elevations and pollution climates

The effects of ambient air compared to filtered air on the reproduction of females and mean relative growth rate (MRGR) of nymphs of C. pilicornis on Norway spruce was determined in open-top chambers at Wengernalp in the Swiss Alps (1900 m a.s.l.) and at Sch?nenbuch near the city of Basle (400 m a.s.l.). The ambient concentration of O(3), the main pollutant at both sites, varied between 45 and 120 microg m(-3) (24-h mean) at both sites. A 5-8 day exposure of spruce saplings to ambient compared to filtered air enhanced the MRGR of nymphs of C. pilicornis of local and northern origins at Sch?nenbuch. The cumulative numbers of offspring of C. pilicornis were higher in ambient air chambers than in filtered air chambers at Sch?nenbuch, but not at Wengernalp. Filtration of ambient air did not affect significantly the levels of total free amino acids or reducing sugars in phloem sap or concentration of total phenolics in needles at the end of the growing season. The results suggest that ambient air with elevated O(3) and with high daily fluctuations, as it can be observed in Sch?nenbuch, affects aphid performance on conifers more than ambient air with also elevated, but rather constant levels of O(3), as it can be observed in mountain forest areas.     

The effects of ozone-exposed sugar maple seedlings on the biological performance and the feeding preference of the forest tent caterpillar (Malacosoma disstria Hbn.)

The effects of exposure of sugar maple (Acer saccharum Marsh.) to ozone on the entire larval stage of a native insect have not been previously investigated. This study reports the effects of sugar maple seedlings exposed to different ozone concentrations on the relative performance and the feeding preference of the forest tent caterpillar (Malacosoma disstria Hbn.). Three-year-old seedlings were set in nine open-top field chambers in the spring of 1992 and 1993. Three ozone concentrations were generated: charcoal-filtered ambient air (0x), ambient air (1x) and three times ambient air (3x). In 1992, female and male larval development time did not differ among ozone treatments. In 1993, female larvae reared on 3x developed faster than those on 0x and 1x, while male larvae were not affected. Ozone treatments did not influence pupal weights except for males in 1993 where pupae reared on 0x were heavier than 1x but did not differ from 3x. Larval and pupal survival rates were not affected by ozone in either year. Finally, 4th and 5th instar larvae showed a significant feeding preference for 3x foliage in 1993 but not in 1992. The response of the forest tent caterpillar to ozone exposed seedlings varied between years and could be more sensitive to annual climatic variations than ozone.     

Yield and grain quality of spring wheat (Triticum aestivum L., cv. Drabant) exposed to different concentrations of ozone in open-top chambers

Spring wheat (Triticum aestivum L., cv. Drabant) was exposed to different concentrations of ozone in open-top chambers for two growing seasons, 1987 and 1988, at a site located in south-west Sweden. The chambers were placed in a field of commercially grown spring wheat. The treatments were charcoal-filtered air (CF), non-filtered air (NF) and non-filtered air plus extra ozone (NF(+)). In 1988, one additional ozone concentration (NF(++)) was used. Grain yield was affected by the ozone concentration of the air. Air filtration resulted in an increase in grain yield of about 7% in both years, compared to NF. The addition of ozone (NF(+), NF(++)) reduced grain yield and increased the content of crude protein of the grain in both years. Filtration of the air had no significant effect on the content of crude protein, compared to NF. The results showed a strong positive chamber effect on grain yield in the cold and wet summer of 1987. In 1988, there was no net chamber effect on grain yield. The relative differences between the CF, NF and NF(+) treatments with respect to grain yield were of the same magnitude in the two years, despite the very different weather conditions.     

Influence of ozone and ethylenediurea (EDU) on growth and yield of bean (Phaseolus vulgaris L.) in open-top field chambers

Greenhouse and ambient air experiments have shown ethylene diurea (EDU) to be a strong and specific protective suppressant of ozone injury in plants. To examine how EDU affects plant responses to various ozone (O(3)) levels under controlled field conditions, Phaseolus vulgaris L. cv. Lit was treated with 150 ppm EDU every 14 days and exposed in open-top chambers to charcoal-filtered air (CF), nonfiltered air (NF) or two cf treatments with ozone added. The ozone treatments were proportional additions of one (CF1) and two (CF2) times ambient ozone levels. The mean ozone concentrations in the CF, NF, CF1 and CF2 treatments were 0.98, 14.1, 14.98 and 31.56 nl litre(-1). A two-way split plot ANOVA revealed that shoot dry weight was significantly reduced by ozone. EDU treatment was highly significant for leaf dry weight, root dry weight and shoot dry weight, but not for pod dry weight; leading to a higher biomass of EDU-treated plants. Ozone/EDU interactions were significant for root weight only, indicating that EDU reduced growth suppression by ozone. These results show that EDU action on plant biomass could be interpreted as a delay in senescence since EDU-treated plants showed a significant decreased biomass loss even in the CF treatment.     

Ozone-induced nitrate formation in needles and leaves of Picea abies, Fagus sylvatica and Quercus robur

Much attention has been paid to ozone as a major cause of novel forest decline in Europe. In combination with acidic mist, O(3) has been observed to increase ion leaching. Besides cations lake Mg(2+), Ca(2+), K(+), NH(4)(+), considerable amounts of nitrate were found to be leached by acidic mist from needles of Norway spruce. Controlled fumigation experiments, with 100, 300, and 600 microg O(3)m(-3) over 22 days continuously, have led to a nitrate accumulation of 94.1 +/- 14.8, 119.4 +/- 28.7 and 198.9 +/- 14.9 microg NO(3)(-1) g(-1) FW, respectively, in leaves of Quercus robur. Similar values were found in leaves of Fagus sylvatica and current and previous year needles of Picea abies. Nitrate levels of controls receiving charcoal filtered air were well below 40 microg NO(3)(-) g (-1) FW. Statistically significant elevated nitrate levels were observed after only 48 h of continuous fumigation with 600 microg O(3)m(-3), in all tree species tested, and after 144 h in the 100 microg O(3)m(-3) treatment. In another experiment, trees of Picea abies were kept in two charcoal (C) and two Purafil plus charcoal (P/C) ventilated chambers, and fumigated with O and 500 microg O(3)m(-3) in cabinets of each filter-type in order to eliminate NO(x) from chamber air. After 29 days of continuous ozone fumigation, NO(3)(-) accumulation in needles amounted to 102.0 +/- 37.7 and 137.4 +/- 40.5 microg g(-1) FW in P/C and C-filtered chambers, respectively. Nitrate contents of controls were below 30 microg NO(3)(-)g(-1) FW at the end of the experiment. No significant differences in NO(3)(-) accumulation between filter treatments were observed. Since NO(x) was reduced by more than 95% in the Purafil/charcoal versus the charcoal treatment, NO(3)(-) accumulation in needles can be attributed predominantly to the influence of ozone and not to direct NO(2) uptake of needles by the possible oxidation of NO to NO(2) in the presence of ozone.     

Effects of elevated CO2 concentration on the polyamine levels of field-grown soybean at three O3 regimes

Effects of increased ozone (O3) and carbon dioxide (CO2) on polyamine levels were determined in soybean (Glycine max L. Merr. cv. Clark) grown in open-top field chambers. The chamber treatments consisted of three O3 regimes equal to charcoal filtered (CF), non-filtered (NF), and non-filtered plus 40 nl litre(-1) O3 and CO2 treatments equal to 350, 400 and 500 microl litre(-1) for a total of nine treatments. Leaf samples were taken at three different times during the growing season. Examination of growth and physiological characteristics, such as photosynthesis, stomatal resistance, and shoot weight, revealed that increasing CO2 ameliorated the deleterious effects of increased O3. Results from the initial harvest, at the pre-flowering growth stage (23 days of treatment), showed that increasing O3 at ambient CO2 caused increases in putrescine (Put) and spermidine (Spd) of up to six-fold. These effects were lessened with increased CO2. Elevated CO2 increased polyamines in plants treated with CF air, but had no effect in the presence of ambient or enhanced O3 levels. Leaves harvested during peak flowering (37 days of treatment) showed O3-induced increases in Put and Spd at ambient CO2 concentrations. However, increased CO2 levels inhibited this response by blocking the O3-induced polyamine increase. Leaves harvested during the pod fill stage (57 days of treatment) showed no significant O3 or CO2 effects on polyamine levels. Our results demonstrate that current ambient O3 levels induce the accumulation of Put and Spd early in the growing season and that further increases in O3 could result in even greater polyamine increases. These results are consistent with a possible antiozonant function for polyamines. The ability of increased CO2 to protect soybeans from O3 damage, however, does not appear to involve polyamine accumulation.     

Effects of dietary and gaseous fluoride on the aphid Aphis fabae

A large proportion of leaf fluoride consists of surface deposits and a sucking herbivore would be expected to take in a smaller load of fluoride in its food than a chewing herbivore. In these experiments, fluoride was applied aerially, through the leaves, and systemically, via the roots, to compare uptake by aphids and effects on their fecundity. Fluoride applied via roots was taken up by both the plants and the aphids, but at high treatment rates the aphids had much lower concentrations than the foliage. When the plants and aphids were fumigated with HF the aphids had much greater loads than the plant shoots, which was due to deposition of F on the insect surfaces. There were no effects of the treatments on aphid reproduction or development time. The aphids obtained some fluoride through their diet which suggests that fluoride is present in the phloem sap, previously thought to be of minor importance.     

Indirect effects of air pollutants: Changes in plant/parasite interactions

The impact of air pollutants on plant/parasite-interactions has been investigated. It could be demonstrated that fumigation of Vicia faba L. with 0.15 ppm SO2 (400 microg m(-3)) or 0.2 ppm NO2 (400 microg m(-3)) during 7 days caused changes in plant metabolism which resulted in higher growth rates of the aphid Aphis fabae Scop. feeding on these plants. Fumigation of V. faba with 0.085 ppm O3 during 2 or 3 days, however, caused decreased aphid growth on fumigated plants. That result could be reversed by higher O3 concentrations or through the presence of NOx during O3 fumigation. Ambient air comprising a mixture of pollutant gases had a strong enhancing effect on aphid performance. Thus, the growth of A. fabae on field bean plants was significantly higher in ambient summertime. London air than in charcoal-filtered air. Similarly, the growth of Macrosiphon rosae L. on rose bushes (Rosa sp., cv. Nina Weibull) was improved in ambient summertime Munich air; the increase in growth rate averaged about 20%.     

Effects of air pollution on rice yield in the Pakistan Punjab

A study using open-top chambers ventilated with ambient or charcoal filtered air in the vicinity of Lahore, Pakistan demonstrated reductions of 42% and 37% in the grain yield of two cultivars of rice (Oryza sativa L.). This yield reduction was primarily due to the reduction in mean panicle number per plant, although significant effects of filtration on 1000 grain weight and the number of filled grains per panicle were also detected. The 6-h daily mean O(3) concentrations were only 10-20 nl litre(-1) during the monsoon season, but increased to 30-55 nl litre(-1) later in the growing season, while the mean NO(2) concentration during the experiment was 12 nl litre(-1). On the basis of experience in North America and Japan, the reductions in yield in the present study are substantially greater than might be predicted. The reasons for this discrepancy are discussed, together with the implications for effects on rice yield in other regions of south and south-east Asia.     

Ambient air pollution induced changes in amino acid pattern of phloem sap in host plants-relevance to aphid infestation

In pollution exclusion experiments on the verge of a widely-used motorway, it has been shown that roadway emissions increase the abundance of the aphid Aphis fabae Scop. on two different host plants, Viburnum opulus L. and Phaseolus vulgaris L. Analysis of phloem sap of excised leaves revealed on increase in almost all of the detected amino acids due to ambient air pollution. To investigate the significance of these changes to aphid development, the patterns analyzed were translated into artificial diets, representing the amino acid composition either of filtered air or ambient air host plants. On 'ambient air' diets, the larvae were significantly larger at birth compared to the progeny on 'filtered air' diets. Also the mean relative growth rates (MRGR) of larvae reared on 'ambient air' diets were significantly increased. At the same time, a conspicuous shorter development time of the larvae could be observed on 'ambient air' diets.     

Effects of open-top chamber fumigations with ozone on three fungal leaf diseases of wheat and the mycoflora of the phyllosphere

Spring wheat (Triticum aestivum L. cv. Turbo) was exposed in open-top chambers to six different ozone levels (8-h daily means from 12.4 to 122 microg m(-3)), to non-filtered air and to chamberless field conditions for 31 days from seedling stage through ear emergence. Powdery mildew (Erysiphe graminis DC. f.sp. tritici Marchal) which developed during the exposure period was significantly enhanced from 0.3/0.6% (two chamber replicates), 1.2/2.1%, 0.9/2.2% in charcoal-filtered air (CF) to 1.5/1.6%, 3.7/4.3%, 4.4/4.6% at the highest level of ozone, on the flag leaf, second and third leaf position, respectively. Post-exposure inoculation with Septoria nodorum Berk. led to increases of disease severity on the flag leaf from 40.9/43.6% in CF to 66.3/70.6% at the highest ozone concentration and on the ears from 15.7/16.5% to 26.3/26.6%. In the same comparison, severity of spot blotch following inoculation with Bipolaris sorokiniana (Sacc.) Shoem. (syn. Helminthosporium sativum Pamm., King et Bakke) was increased on the flag leaf from 3.6/8.9% to 12.3/23.4%. The three diseases examined correlated significantly with the ozone treatments in fumigated chambers. Disease severity was enhanced even on undamaged plant tissue (flag leaves). Infections of the two facultative pathogens on lower leaf positions started only in part from visible ozone lesions, mildew did not start from such lesions. No significant effects of ozone in the chambers on the saprobial colonization of the phyllosphere were detected, whereas there were marked differences in this respect between plants from the field and the chambers. At the highest ozone treatment, contents of chlorophyll a and carotenoids on the second leaf position declined significantly, which was associated with symptoms of premature senescence. Senescing effects of ozone are therefore assumed to be one major factor in predisposing wheat for necrotrophic leaf pathogens. Surprisingly, injurious and predisposing effects of ozone were completely absent in chambers supplied with non-filtered air containing ambient ozone at doses equivalent to those in CF + ozone chambers. Evidently, biological effects of ozone in pure air and in ambient air may differ markedly.