Sagar V. Krupa

Plant responses to atmospheric CO2 enrichment with emphasis on roots and the rhizosphere

Empirical records provide incontestable evidence of global changes: foremost among these changes is the rising concentration of CO(2) in the earths atmosphere. Plant growth is nearly always stimulated by elevation of CO(2). Photosynthesis increases, more plant biomass accumulates per unit of water consumed, and economic yield is enhanced. The profitable use of supplemental CO(2) over years of greenhouse practice points to the value of CO(2) for plant production. Plant responses to CO(2) are known to interact with other environmental factors, e.g. light, temperature, soil water, and humidity. Important stresses including drought, temperature, salinity, and air pollution have been shown to be ameliorated when CO(2) levels are elevated. In the agricultural context, the growing season has been shortened for some crops with the application of more CO(2); less water use has generally, but not always, been observed and is under further study; experimental studies have shown that economic yield for most crops increases by about 33% for a doubling of ambient CO(2) concentration. However, there are some reports of negligible or negative effects. Plant species respond differently to CO(2) enrichment, therefore, clearly competitive shifts within natural communities could occur. Though of less importance in managed agro-ecosystems, competition between crops and weeds could also be altered. Tissue composition can vary as CO(2) increases (e.g. higher C: N ratios) leading to changes in herbivory, but tests of crop products (consumed by man) from elevated CO(2) experiments have generally not revealed significant differences in their quality. However, any CO(2)-induced change in plant chemical or structural make-up could lead to alterations in the plants interaction with any number of environmental factors-physicochemical or biological. Host-pathogen relationships, defense against physical stressors, and the capacity to overcome resource shortages could be impacted by rises in CO(2). Root biomass is known to increase but, with few exceptions, detailed studies of root growth and function are lacking. Potential enhancement of root growth could translate into greater rhizodeposition, which, in turn, could lead to shifts in the rhizosphere itself. Some of the direct effects of CO(2) on vegetation have been reasonably well-studied, but for others work has been inadequate. Among these neglected areas are plant roots and the rhizosphere. Therefore, experiments on root and rhizosphere response in plants grown in CO(2)-enriched atmospheres will be reviewed and, where possible, collectively integrated. To this will be added data which have recently been collected by us. Having looked at the available data base, we will offer a series of hypotheses which we consider as priority targets for future research.

Effects of atmospheric ammonia (NH3) on terrestrial vegetation: a review

At the global scale, among all N (nitrogen) species in the atmosphere and their deposition on to terrestrial vegetation and other receptors, NH3 (ammonia) is considered to be the foremost. The major sources for atmospheric NH3 are agricultural activities and animal feedlot operations, followed by biomass burning (including forest fires) and to a lesser extent fossil fuel combustion. Close to its sources, acute exposures to NH3 can result in visible foliar injury on vegetation. NH3 is deposited rapidly within the first 4-5 km from its source. However, NH3 is also converted in the atmosphere to fine particle NH4+ (ammonium) aerosols that are a regional scale problem. Much of our current knowledge of the effects of NH3 on higher plants is predominantly derived from studies conducted in Europe. Adverse effects on vegetation occur when the rate of foliar uptake of NH3 is greater than the rate and capacity for in vivo detoxification by the plants. Most to least sensitive plant species to NH3 are native vegetation > forests > agricultural crops. There are also a number of studies on N deposition and lichens, mosses and green algae. Direct cause and effect relationships in most of those cases (exceptions being those locations very close to point sources) are confounded by other environmental factors, particularly changes in the ambient SO2 (sulfur dioxide) concentrations. In addition to direct foliar injury, adverse effects of NH3 on higher plants include alterations in: growth and productivity, tissue content of nutrients and toxic elements, drought and frost tolerance, responses to insect pests and disease causing microorganisms (pathogens), development of beneficial root symbiotic or mycorrhizal associations and inter species competition or biodiversity. In all these cases, the joint effects of NH3 with other air pollutants such as all-pervasive O3 or increasing CO2 concentrations are poorly understood. While NH3 uptake in higher plants occurs through the shoots, NH4+ uptake occurs through the shoots, roots and through both pathways. However, NH4+ is immobile in the soil and is converted to NO3- (nitrate). In agricultural systems, additions of NO3- to the soil (initially as NH3 or NH4+) and the consequent increases in the emissions of N2O (nitrous oxide, a greenhouse gas) and leaching of NO3- into the ground and surface waters are of major environmental concern. At the ecosystem level NH3 deposition cannot be viewed alone, but in the context of total N deposition. There are a number of forest ecosystems in North America that have been subjected to N saturation and the consequent negative effects. There are also heathlands and other plant communities in Europe that have been subjected to N-induced alterations. Regulatory mitigative approaches to these problems include the use of N saturation data or the concept of critical loads. Current information suggests that a critical load of 5-10 kg ha(-1) year(-1) of total N deposition (both dry and wet deposition combined of all atmospheric N species) would protect the most vulnerable terrestrial ecosystems (heaths, bogs, cryptogams) and values of 10-20 kg ha(-1) year(-1) would protect forests, depending on soil conditions. However, to derive the best analysis, the critical load concept should be coupled to the results and consequences of N saturation.

Atmospheric ozone: Formation and effects on vegetation

Ozone (O(3)) is present both in the troposphere and the stratosphere. Troposphere O(3) is predominantly produced by photochemical reactions involving precursors generated by natural processes and to a much larger extent by mans activities. There is evidence for a trend towards increasing tropospheric O(3) concentrations. However, tropospheric O(3) is known to account for only 10% of the vertical O(3) column above the earths surface. The stratosphere accounts for an additional 90% of the O(3) column. There is evidence to suggest that there are losses in the stratospheric O(3) due to the updraft of O(3) destroying pollutants generated by both natural processes and by human activity. Such a loss in stratospheric O(3) can result in alterations of incidence in the ultraviolet (UV) radiation to the earths surface. Tropospheric O(3) is known to be highly phytotoxic. Appropriate exposures to O(3) can result in both acute (symptomatic) and chronic (changes in growth, yield or productivity and quality) effects. Chronic effects are of great concern in terms of both crops and forests. A number of experimental techniques are available to evaluate the chronic effects of O(3) on plants. There are limitations attached to the use of these techniques. However, results obtained, with such techniques are valuable if interpreted in the appropriate context. Among all field evaluation techniques, open-top chambers are the most frequently used method for evaluating the chronic effects of O(3) on crops. The National Crop Loss Assessment Program (NCLAN) of the United States is the largest such effort. However, given the limitations of the open-top chambers and the experimental aspects of NCLAN, its results must be interpreted with caution. On the other hand, acute effects can be evaluated with less complexity through the use of biological indicator plants. The numerical modelling of such effects are also far less complicated than establishing numerical cause and effects relationships for chronic effects. Confounding the acute or chronic responses of plants to O(3), is the presence of other kinds and forms of pollutants in the ambient atmosphere and the incidence of pathogens and pests. The resulting complex interactions and joint effects on plants are poorly understood. Future research must address these issues. In the final analysis we have re-emphasized the fact that plant health is the product of its interaction with the physical and chemical climatology and pathogens and pests. What we have described in this context is the importance of tropospheric O(3) within the chemical climatology of our environment and its effects on vegetation.

Passive sampling of ambient, gaseous air pollutants: an assessment from an ecological perspective

During some past two decades there has been a growing interest among air pollution-vegetation effects-scientists to use passive sampling systems for quantifying ambient, gaseous air pollutant concentrations, particularly in remote and wilderness areas. On the positive side, excluding the laboratory analysis costs, passive samplers are inexpensive, easy to use and do not require electricity to operate. Therefore, they are very attractive for use in regional-scale air quality assessments. Passive samplers allow the quantification of cumulative air pollutant exposures, as total or average pollutant concentrations over a sampling duration. Such systems function either by chemical absorption or by physical adsorption of the gaseous pollutant of interest onto the sampling medium. Selection of a passive sampler must be based on its known or tested characteristics of specificity and linearity of response to the chemical constituent being collected. In addition, the effects of wind velocity, radiation, temperature and relative humidity must be addressed in the context of absorbent/adsorbent performance and sampling rate. Because of all these considerations, passive samplers may provide under- or overestimations of the cumulative exposures, compared to the corresponding data from co-located continuous monitors or active samplers, although such statistical variance can be minimized by taking necessary precautions. On the negative side, cumulative exposures cannot identify short-term (<few hours) pollutant episodes or regulatory non-compliance, where appropriate. Equally important, cumulative exposures (e.g. with ozone, a non-accumulating pollutant in plants) cannot account for the stochasticity and the dynamics of air pollutant exposure and plant (particularly deciduous vegetation) response. Although still being tested, statistical methods appear to be available to reconstruct passive sampler data to mimic the dynamics of the co-located continuous monitoring data. Regional level air quality and ecological risk assessments (multi-point models) as a whole should consider this type of approach in the future, using co-located passive samplers, with continuous monitors at selected locations as calibration points for mapping the air quality distribution on temporal and spatial scales.

The impact of UV-B radiation and ozone on terrestrial vegetation

Although terrestrial vegetation has been exposed to UV-B radiation and ozone over the course of evolutionary history, it is essential to view the effects on vegetation of changing levels of these factors in the context of other features of climate change, such as increasing CO(2) levels and changes in temperature and precipitation patterns. Much of our understanding of the impacts of increased UV-B and ozone levels has come from studies of the effects of each individual factor. While such information may be relevant to a wider understanding of the roles that these factors may play in climate change, experience has shown that the interactions of environmental stresses on vegetation are rarely predictable. A further limitation on the applicability of such information results from the methodologies used for exposing plants to either factor. Much of our information comes from growth chamber, greenhouse or field studies using experimental protocols that made little or no provision for the stochastic nature of the changes in UV-B and ozone levels at the earths surface, and hence excluded the roles of repair mechanisms. As a result, our knowledge of dose-response relationships under true field conditions is both limited and fragmentary, given the wide range of sensitivities among species and cultivars. Adverse effects of increased levels of either factor on vegetation are qualitatively well established, but the quantitative relationships are far from clear. In both cases, sensitivity varies with stage of plant development. At the population and community levels, differential responses of species to either factor has been shown to result in changes in competitiveness and community structure. At the mechanistic level, ozone generally inhibits photosynthetic gas exchange under both controlled and field conditions, and although UV-B is also inhibitory in some species under controlled conditions, others appear to be indifferent, particularly in the field. Both factors affect metabolism; a common response is increased secondary metabolism leading to the accumulation of phenolic compounds that, in the case of UV-B, offer the leaf cell some protection from radiation. Virtually no information is available about the effects of simultaneous or sequential exposures. Since both increased surface UV-B and ozone exposures have spatial and temporal components, it is important to evaluate the different scenarios that may occur, bearing in mind that elevated daytime ozone levels will attenuate the UV-B reaching the surface to some extent. The experimentation needed to acquire unequivocal effects data that are relevant to field situations must therefore be carried out using technologies and protocols that focus on quantification of the interactions of UV-B and ozone themselves and their interactions with other environmental factors.

Photochemical oxidants: state of the science.

Atmospheric photochemical processes resulting in the production of tropospheric ozone (O(3)) and other oxidants are described. The spatial and temporal variabilities in the occurrence of surface level oxidants and their relationships to air pollution meteorology are discussed. Models of photooxidant formation are reviewed in the context of control strategies and comparisons are provided of the air concentrations of O(3) at select geographic locations around the world. This overall oxidant (O(3)) climatology is coupled to human health and ecological effects. The discussion of the effects includes both acute and chronic responses, mechanisms of action, human epidemiological and plant population studies and briefly, efforts to establish cause-effect relationships through numerical modeling. A short synopsis is provided of the interactive effects of O(3) with other abiotic and biotic factors. The overall emphasis of the paper is on identifying the current uncertainties and gaps in our understanding of the state of the science and some suggestions as to how they may be addressed.

Ambient ozone (O3) and adverse crop response : a unified view of cause and effect

This paper presents a cohesive view of the dynamics of ambient O(3) exposure and adverse crop response relationships, coupling the properties of photochemical O(3) production, flux of O(3) from the atmosphere into crop canopies and the crop response per se. The results from two independent approaches ((a) statistical and (b) micrometeorological) were analyzed for understanding cause-effect relationships of the foliar injury responses of tobacco cv Bel-W3 to the exposure dynamics of ambient O(3) concentrations. Similarly, other results from two independent approaches were analyzed in: (1) establishing a micrometeorological relationship between hourly ambient O(3) concentrations and their vertical flux from the air into a natural grassland canopy; and (2) establishing a statistical relationship between hourly ambient O(3) concentrations in long-term, chronic exposures and crop yield reductions. Independent of the approach used, atmospheric conditions appeared to be most conducive and the crop response appeared to be best explained statistically by the cumulative frequency of hourly ambient O(3) concentrations between 50 ppb and 90 ppb (100 and 180 microg m(-3)). In general, this concentration range represents intermediate or moderately enhanced hourly O(3) values in a polluted environment. Further, the diurnal occurrence of this concentration range (often approximately between 0900 and 1600 h in a polluted, agricultural environment) coincided with the optimal CO(2) flux from the atmosphere into the crop canopy, thus high uptake. The frequency of occurrence of hourly O(3) concentrations > 90 ppb (180 microg m(-3)) appeared to be of little importance and such concentrations in general appeared to occur during atmospheric conditions which did not facilitate optimal vertical flux into the crop canopy, thus low uptake. Alternatively, when > 90 ppb (180 microg m(-3)) O(3) concentrations occurred during the 0900-1600 h window, their frequency of occurrence was low in comparison to the 50-90 ppb (100-180 microg m(-3)) range. Based on the overall results, we conclude that if the cumulative frequency of hourly ambient O(3) concentrations between 50-62 ppb (100-124 microg m(-3)) occurred during 53% of the growing season and the corresponding cumulative frequency of hourly O(3) concentrations between 50-74 ppb (100-148 microg m(-3)) occurred during 71% of the growing season, then yield reductions in sensitive crops could be expected, if other factors supporting growth, such as adequate soil moisture are not limiting.

Measurements of S(IV) and organic anions in Minnesota rain

Abstract In 167 fractions from 84 rain events collected at two sites in Minnesota during the summer of 1982, S(IV) accounted 1.12% of the measured anions and thus usually contributed very little of the free acidity. A few events were observed, however, in which the S(IV) contribution was large. Organic acids were observed in nearly every fraction, with acetate being the most abundant. The samples were not preserved to prevent the loss of organic acids, which made an accurate assessment of the contribution of the organics to the acidity impossible and may have accounted for the low amounts of formate observed. The pH of samples in storage increased with time, a phenomenon which appeared to be related to gas exchange.

Forest responses to tropospheric ozone and global climate change: An analysis

In this paper an analysis is provided on: what we know, what we need to know, and what we need to do, to further our understanding of the relationships between tropospheric ozone (O(3)), global climate change and forest responses. The relationships between global geographic distributions of forest ecosystems and potential geographic regions of high photochemical smog by the year 2025 AD are described. While the emphasis is on the effects of tropospheric O(3) on forest ecosystems, discussion is presented to understand such effects in the context of global climate change. One particular strong point of this paper is the audit of published surface O(3) data by photochemical smog region that reveals important forest/woodland geographic regions where little or no O(3) data exist even though the potential threat to forests in those regions appears to be large. The concepts and considerations relevant to the examination of ecosystem responses as a whole, rather than simply tree stands alone are reviewed. A brief argument is provided to stimulate the modification of the concept of simple cause and effect relationships in viewing total ecosystems. Our knowledge of O(3) exposure and its effects on the energy, nutrient and hydrological flow within the ecosystem are described. Modeling strategies for such systems are reviewed. A discussion of responses of forests to potential multiple climatic changes is provided. An important concept in this paper is that changes in water exchange processes throughout the hydrological cycle can be used as early warning indicators of forest responses to O(3). Another strength of this paper is the integration of information on structural and functional processes of ecosystems and their responses to O(3). An admitted weakness of this analysis is that the information on integrated ecosystem responses is based overwhelmingly on the San Bernardino Forest ecosystem research program of the 1970s because of a lack of similar studies. In the final analysis, it is recommended that systems ecology be applied in examining the joint effects of O(3), carbon dioxide and ultraviolet-B radiation on forest ecosystems.

Foliar sulphur species in pine: a new indicator of a forest ecosystem under air pollution stress.

An eleven-year foliar sulphur (S) monitoring program was carried out from 1976 to 1986 near a sulphur recovery-gas plant in west-central Alberta, Canada, as part of a case study designed to determine the effects of chronic, low concentration sulphur gas emissions on the forest ecosystem surrounding the gas plant. Measurements of both foliar total sulphur (ST) and foliar inorganic sulphur (SO4-S) concentration in lodgepole x jack pine trees at the end of each of the 11 growing seasons were taken to provide an indication of S loading of the forest from industrial sulphur emissions. To measure the state of the forest ecosystem, foliar ST was separated into foliar accumulated sulphur (inorganic sulphur or SO4-S) and foliar assimilated sulphur (organic sulphur or S0) and the ratio of SO4-S/S0 taken. Foliar S0 was calculated as the difference between foliar ST and foliar SO4-S. The median SO4-S/S0 ratio, with all three years of needles considered, varied from 0.29 at a reference location (AV) to 0.88 at the location with the highest stress (AI). The corresponding mean values ranged from 0.3 at the reference location to 2.2 at the location of highest stress. The mean seasonal photosynthetic rate of current years foliage of the pine trees and soil pH were reduced at a stressed location (AI) compared to the reference location (AV), between 1976 and 1981. Over this same time period the mean foliar SO4-S/S0 ratio increased from 0.4 +/- 0.1 to 1.0 +/- 0.3 at the stressed location (AI) and remained nearly the same at the reference location (AV) at 0.3 +/- 0.1. This research suggests that the foliar SO4-S/S0 ratio is a useful indicator of the state of forest ecosystems under S air pollution stress. It is concluded that foliar S separated into various fractions has potential as an early warning environmental management tool.