Effects of source-sink relations on photosynthetic acclimation to elevated CO2

Abstract. While photosynthesis of C₃ plants is stimulated by an increase in the atmospheric CO₂ concentration, photosynthetic capacity is often reduced after long-term exposure to elevated CO₂. This reduction appears to be brought about by end product inhibition, resulting from an imbalance in the supply and demand of carbohydrates. A review of the literature revealed that the reduction of photosynthetic capacity in elevated CO₂ was most pronounced when the increased supply of carbohydrates was combined with small sink size. The volume of pots in which plants were grown affected the sink size by restricting root growth. While plants grown in small pots had a reduced photosynthetic capacity, plants grown in the field showed no reduction or an increase in this capacity. Pot volume also determined the effect of elevated CO₂ on the root/shoot ratio: the root/shoot ratio increased when root growth was not restricted and decreased in plants grown in small pots. The data presented in this paper suggest that plants growing in the field will maintain a high photosynthetic capacity as the atmospheric CO₂ level continues to rise.     

Free Air Temperature Increase (FATI): a new tool to study global warming effects on plants in the field

A new technique, called Free Air Temperature Increase (FATI), was developed to artificially induce increased canopy temperature in field conditions without the use of enclosures. This acronym was chosen in analogy with FACE (Free Air CO₂ Enrichment), a technique which produces elevated CO₂ concentrations [CO₂] in open field conditions. The FATI system simulates global warming in small ecosystems of limited height, using infrared heaters from which all radiation below 800 nm is removed by selective cut-off filters to avoid undesirable photomorpho-genetic effects. An electronic control circuit tracks the ambient canopy temperature in an unheated reference plot with thermocouples, and modulates the radiant energy from the lamps to produce a 2.5°C increment in the canopy temperature of an associated heated plot (continuously day and night). This pre-set target differential is relatively-constant over time due to the fast response of the lamps and the use of a proportional action controller (the standard deviation of this increment was <1°C in a 3 week field study with 1007 measurements). Furthermore, the increase in leaf temperature does not depend on the vertical position within the canopy or on the height of the stand. Possible applications and alternative designs are discussed.     

Physiological and growth responses of two African species, Acacia karroo and Themeda triandra, to combined increases in CO2 and UV-B radiation

The interactive effects of increased carbon dioxide (CO₂) concentration and ultraviolet-B (UV-B, 280–320 nm) radiation on Acacia karroo Hayne, a C₃ tree, and Themeda triandra Forsk., a C₄ grass, were investigated. We tested the hypothesis that A. karroo would show greater CO₂-induced growth stimulation than T. triandra, which would partially explain current encroachment of A. karroo into C₄ grasslands, but that increased UV-B could mitigate this advantage. Seedlings were grown in open-top chambers in a greenhouse in ambient (360 μmol mol^-1) and elevated (650 μmol mol^-1) CO₂, combined with ambient (1.56 to 8.66 kJ m^-2 day^-1) or increased (2.22 to 11.93 kJ m^-2 day^-1) biologically effective (weighted) UV-B irradiances. After 30 weeks, elevated CO₂ had no effect on biomass of A. karroo, despite increased net CO₂ assimilation rates. Interaction between UV-B and CO₂ on stomatal conductance was found, with conductances decreasing only where elevated CO₂ and UV-B were supplied separately. Increases in water use efficiencies, foliar starch concentrations, root nodule numbers and total nodule mass were measured in elevated CO₂. Elevated UV-B caused only an increase in foliar carbon concentrations. In T. triandra, net CO₂ assimilation rates were unaffected in elevated CO₂, but stomatal conductances and foliar nitrogen concentrations decreased, and water use efficiencies increased. Biomass of all vegetative fractions, particularly leaf sheaths, was increased in elevated CO₂. and was accompanied by increased leaf blade lengths and individual leaf and leaf sheath masses. However, tiller numbers were reduced in elevated CO₂. Significantly moderating effects of elevated UV-B were apparent only in individual masses of leaf blades and sheaths, and in total sheath and shoot biomass. The direct CO₂-induced growth responses of the species therefore do not support the hypothesis of CO₂-driven woody encroachment of C₄ grasslands. Rather, differential changes in resource use efficiency between grass and woody species, or morphological responses of grass species, could alter the competitive balance. Increased UV-B radiation is unlikely to substantially alter the CO₂ response of these species.     

Effect of increased CO2 concentration and temperature on the phyllosphere mycoflora of winter wheat flag leaves during ripening

The impact of elevated carbon dioxide (CO₂, 600/700 μmol mol^-1) and temperature (+ 4°C) on phyllosphere fungi colonising flag leaves of mini crops of winter wheat cv. Mercia between anthesis and harvest was determined in a computer-controlled environment facility in 1993 and 1994. In both years the total fungal populations (cm² leaf) were found to have increased due to exposure to either elevated CO₂ and elevated CO₂+ temperature treatments. This was mainly due to significant increases in populations of Cladosporium spp. (C. cladosporioides and C. herbarum) on the flag leaves during ripening. Other phyllosphere component species such as white and pink yeasts were not markedly affected by treatments. The range of fungal species found in such controlled environment chambers was narrower than that commonly found on flag leaves of field grown crops. Common and important colonisers of leaves and ripening ears such as Aureobasidium pullulans, Epicoccum nigrum and Fusarium spp. were seldom isolated.     

Effects of elevated CO2 and/or O3 on growth, development and physiology of wheat (Triticum aestivum L.)

Two cultivars of spring wheat (Triticum aestivum L. cvs. Alexandria and Hanno) and three cultivars of winter wheat (cvs. Riband, Mercia and Haven) were grown at two concentrations of CO₂ [ambient (355 pmol mol^?1) and elevated (708 μmol mol^?1)] under two O₃ regimes [clean air (< 5 nmol mol^?1 O₃) and polluted air (15 nmol mol^?1 O₃ at night rising to a midday maximum of 75 nmol mol^?1)] in a phytotron at the University of Newcastle-upon-Tyne. Between the two-leaf stage and anthesis, measurements of leaf gas-exchange, non-structural carbohydrate content, visible O₃ damage, growth, dry matter partitioning, yield components and root development were made in order to examine responses to elevated CO₂ and/or O₃. Growth at elevated CO₂ resulted in a sustained increase in the rate of CO₂ assimilation, but after roughly 6 weeks' exposure there was evidence of a slight decline in the photosynthetic rate (c.-15%) measured under growth conditions which was most pronounced in the winter cultivars. Enhanced rates of CO₂ assimilation were accompanied by a decrease in stomatal conductance which improved the instantaneous water use efficiency of individual leaves. CO₂ enrichment stimulated shoot and root growth to an equivalent extent, and increased tillering and yield components, however, non-structural carbohydrates still accumulated in source leaves. In contrast, long-term exposure to O₃ resulted in a decreased CO₂ assimilation rate (c. -13%), partial stomatal closure, and the accumulation of fructan and starch in leaves in the light. These effects were manifested in decreased rates of shoot and root growth, with root growth more severely affected than shoot growth. In the combined treatment growth of O₃-treated plants was enhanced by elevated CO₂, but there was little evidence that CO₂ enrichment afforded additional protection against O₃ damage. The reduction in growth induced by O₃ at elevated CO₂ was similar to that induced by O₃ at ambient CO₂ despite additive effects of the individual gases on stomatal conductance that would be expected to reduce the O₃ flux by 20%, and also CO₂-induced increases in the provision of substrates for detoxification and repair processes. These observations suggest that CO₂ enrichment may render plants more susceptible to O₃ damage at the cellular level. Possible mechanisms are discussed.     

Plant growth and reproduction along CO2 gradients: non-linear responses and implications for community change

The effects of rising atmospheric CO₂ concentrations on natural plant communities will depend upon the cumulative responses of plant growth and reproduction to gradual, incremental changes in climatic conditions. We analysed published studies of plant responses to elevated CO₂ to address whether reproductive and total biomass exhibit similar enhancement to elevated vs. ambient CO₂ concentrations, and to assess the patterns of plant response along gradients of CO₂ concentrations. In six annual plant species, mean enhancement at double ambient vs. ambient CO₂ was 1.13 for total biomass and 1.30 for reproductive biomass. The two measures were significantly correlated, but there was considerable scatter in the relationship, indicating that reproductive responses cannot be consistently predicted from enhancement of total biomass. Along experimental CO₂ gradients utilizing three concentrations, there was a great diversity of response patterns, including positive, negative, non-monotonic and non-significant (flat) responses. The distribution of response patterns differed for plants grown in stands compared to those grown individually. Positive responses were less frequent in competitive environments, and non-monotonic responses were more frequent. These results emphasize that interpolation of plant response based on enhancement ratios measured at elevated vs. ambient CO₂ concentrations is not sufficient to predict community responses to incremental changes in atmospheric conditions. The consequences of differential response patterns were assessed in a simulation of community dynamics for four species of annual plants. The model illustrates that the final community composition at a future point in time depends critically on both the magnitude and the rate of increase of atmospheric CO₂.     

The effects on Arbutus unedo L. of long-term exposure to elevated CO2

Arbutus unedo is a sclerophyllous evergreen, characteristic of Mediterranean coastal scrub vegetation. In Italy, trees of A. unedo have been found close to natural CO₂ vents where the mean atmospheric carbon dioxide concentration is about 2200 μmol mol^?1. Comparisons were made between trees growing in elevated and ambient CO₂ concentrations to test for evidence of adaptation to long-term exposure to elevated CO₂. Leaves formed at elevated CO₂ have a lower stomatal density and stomatal index and higher specific leaf area than those formed at ambient CO₂, but there was no change in carbon to nitrogen ratios of the leaf tissue. Stomatal conductance was lower at elevated CO₂ during rapid growth in the spring. In mid-summer, under drought stress, stomatal closure of all leaves occurred and in the autumn, when stress was relieved, the conductance of leaves at both elevated and ambient CO₂ increased. In the spring, the stomatal conductance of the new flush of leaves at ambient CO₂ was higher than the leaves at elevated CO₂, increasing instantaneous water use efficiency at elevated CO₂. Chlorophyll fluorescence measurements suggested that elevated CO₂ provided some protection against photoinhibition in mid-summer. Analysis of A/C_i curves showed that there was no evidence of either upward or downward regulation of photosynthesis at elevated CO₂. It is therefore anticipated that A. unedo will have higher growth rates as the ambient CO₂ concentrations increase.     

Insects and fungi on a C3 sedge and a C4 grass exposed to elevated atmospheric CO2 concentrations in open-top chambers in the field

The effects of elevated atmospheric CO₂ concentration on plant-fungi and plant-insect interactions were studied in an emergent marsh in the Chesapeake Bay. Stands of the C₃ sedge Scirpus olneyi Grey, and the C₄ grass Spartina patens (Ait.) Muhl. have been exposed to elevated atmospheric CO₂ concentrations during each growing season since 1987. In August 1991 the severities of fungal infections and insect infestations were quantified. Shoot nitrogen concentration ([N]) and water content (WC) were determined. In elevated concentrations of atmospheric CO₂, 32% fewer S. olneyi plants were infested by insects, and there was a 37% reduction in the severity of a pathogenic fungal infection, compared with plants grown in ambient CO₂ concentrations. S. olneyi also had reduced [N], which correlated positively with the severities of fungal infections and insect infestations. Conversely, S. patens had increased WC but unchanged [N] in elevated concentrations of atmospheric CO₂ and the severity of fungal infection increased. Elevated atmospheric CO₂ concentration increased or decreased the severity of fungal infection depending on at least two interacting factors, [N] and WC; but it did not change the number of plants that were infected with fungi. In contrast, the major results for insects were that the number of plants infected with insects decreased, and that the amount of tissue that each insect ate also decreased.