Global warming increases the risk of pests and weeds before wintering, and decreases the grain yield of winter wheat. Therefore, the sowing date should be delayed properly. But the variety of winter wheat that can adapt to late sowing remains unclear. Here, we selected two winter wheat cultivars and evaluated four sowing date treatments on 1 October (early sowing), 8 (normal sowing), 15 (late sowing) and 22 (latest sowing) over two wheat-growing seasons at the experimental Station of Shandong Agricultural University (35°96′N, 117°06′E), Daiyue District, Taian, Shandong, China. We examined the effects of sowing date on a few traits, and found that, compared with normal sowing, though spike number decreased, grain yield was maintained above 9300 kg/ha under late and latest sowing. The main reason was that the more accumulated N from jointing to anthesis resulted in a higher grain number per spike. The higher net photosynthetic rate after anthesis, through optimizing N distribution in the canopy and increasing Rubisco content of flag leaves, improved dry matter accumulation rate and contribution ratio of vegetative organs, ultimately, ensured consistent grain weight. The grain yield of high-tillering winter wheat cultivars decreased from 9370 to 8346 kg/ha. The main reason was that spike number, accumulated N from jointing to anthesis and net photosynthetic rate decreased significantly, which reduced the dry matter accumulation rate, and only satisfied less grains to achieve consistent grain weight. Therefore, low-tillering winter wheat cultivars are more adaptable to late sowing, and can reduce the harm of global warming.

Single-leaf CO2 assimilation rate under saturating light (CA) varies as a function of leaf nitrogen content per unit leaf area (NL). Measured CA for many crop species also declines with developmental stage. Because these relationships may have strong implications for crop–weed competition, a field experiment was conducted to quantify corn and velvetleaf CA as influenced by leaf NL and stage of development. Crop and weed CA were measured on the most recent fully expanded leaves of plants grown in four nitrogen (N) application treatments. Both corn and velvetleaf CA increased with increasing NL, up to about 1.5 g N m−2. Corn and velvetleaf NL values were rarely less than 0.75 g N m−2, indicating that both species may restrict leaf growth in order to maintain constant NL under conditions of limited soil N. Corn CA declined to half its maximum by physiological maturity, whereas velvetleaf CA only declined 18%. Although velvetleaf CA was considerably lower than that of corn, the difference decreased as the growing season progressed. Because corn leaf production is complete by anthesis and velvetleaf continues vegetative production throughout its life cycle, velvetleaf will produce relatively greater quantities of biomass late in the season, which may increase competition for light.

Experiments were conducted to determine the effects of temperature on seed germination and growth of redroot pigweed, Palmer amaranth, and common waterhemp. At 15/10 C day and night temperature, respectively, no seed germination was observed in any species. Seed germination increased gradually as temperature increased. Germination peaked at 25/20 C in common waterhemp and at 35/30 C in redroot pigweed and Palmer amaranth. Seed germination of all three species declined when temperatures increased above 35/30 C. All three species produced less biomass at 15/10 C than at 25/20 C and 35/25 C. Redroot pigweed and common waterhemp biomass were similar at 15/10 C and higher than that of Palmer amaranth. However, Palmer amaranth produced more biomass than redroot pigweed and common waterhemp at 25/20 and 35/30 C. At 45/40 C, redroot pigweed, common waterhemp, and Palmer amaranth plants died 8, 9, and 25 d after initiation of heat treatment, respectively. The largest root volume among the three species was in Palmer amaranth grown at 35/30 C, whereas the smallest root volume was produced by Palmer amaranth grown at 15/10 C. Potential quantum efficiency (F v/F max) of Palmer amaranth was higher than that of redroot pigweed and common waterhemp at higher temperature. The greater growth of Palmer amaranth at higher temperatures may be attributed in part to its extensive root growth and greater thermostability of its photosynthetic apparatus.

The present study investigates the photobiont diversity of the boreal felt lichen, Erioderma pedicellatum. Previously sampled genetic data from Newfoundland were reanalyzed and new sequence data (16S rDNA, rbcLX) of the boreal felt lichen from Alaska (USA), Kamchatka (Russia), and North Trøndelag (Norway) were generated. The highest genetic diversity of the photobiont is found in Alaska and Kamchatka, indicating that these may be the primary sources of the species in the Northern Hemisphere. In Newfoundland, the photobiont of E. pedicellatum was screened on leaves of the symbiotic liverwort Frullania asagrayana and it was found to occur on trees where no other lichens were present, demonstrating that the geographical distribution, and possibly also the ecological requirement of the photobiont of E. pedicellatum, is wider than that of the lichen phenotype. Finally, a postulated association between the occurrence of the vegetatively reproducing Coccocarpia palmicola and the occurrence of the compatible photobiont of E. pedicellatum on the same tree could not be established.

The long-term utility of DNA banks is predicated on the stability of DNA during storage. The quality and yield of DNA extracted from seeds from four garden species, which varied in age from 1 to 135 years old, was used to examine the early stages of DNA degradation. Seeds that were 70 years old yielded high molecular weight DNA, which permitted amplification of a 650 bp segment of the internal transcribed spacer (ITS) region of the nuclear ribosomal DNA and a 487 bp segment of the plastid-encoded large subunit of rubisco (rbcL). DNA extracted from seeds estimated to be about 135 years old was degraded, but still permitted amplification when routine extraction procedures were used. The rbcL sequences obtained from c. 135-year-old cucurbit seed DNA were consistent with species identifications based on seed morphological features; however, ITS sequences from the same samples were determined to be of fungal origin. A comparison of our results with studies of DNA stability in leaf specimens suggested that DNA degraded within both tissues, but perhaps more slowly within seeds. Evidence that high-quality DNA can be extracted from old, non-viable seeds expands the utility of seed banks in preserving genetic resources.

The present work aimed to differentiate between proteolytic activities of plants and micro-organisms during the incubation of grass in cattle rumens. Freshly cut ryegrass was placed in bags of varying permeability and incubated for 16 h in the rumens of dairy cows that had previously grazed a ryegrass sward, supplemented with 4 kg dairy concentrate daily. Woven polyester bags (50 μm pore size) permitted direct access of the micro-organisms and rumen fluid enzymes to the plant material. The polythene was impermeable even to small molecules such as NH3. Dialysis tubing excluded micro-organisms and rumen enzymes/metabolites larger than 10 kDa. DM loss was 46·3 % in polyester, 36·2 % in polythene and 38·1 % in dialysis treatments. It is possible that the DM loss within polythene bags occurred due to a solubilisation of plant constituents (e.g. water-soluble carbohydrates) rather than microbial attachment/degradation processes. The final protein content of the herbage residues was not significantly different between treatments. Regardless of bag permeability, over 97 % of the initial protein content was lost during incubations in situ. Electrophoretic separation showed that Rubisco was extensively degraded in herbage residues whereas the membrane-associated, light-harvesting protein remained relatively undegraded. Protease activity was detected in herbage residues and bathing liquids after all incubation in situ treatments. Although rumen fluid contains proteases (possibly of plant and microbial origin), our results suggest that, owing to cell compartmentation, their activity against the proteins of intact plant cells is limited, supporting the view that plant proteases are involved in the degradation of proteins in freshly ingested herbage.

Lobaria pulmonaria is an epiphytic lichen that, in south-eastern Canada, inhabits deciduous forests where it must acclimate to large seasonal changes in temperature and in light caused by closing and opening of the leaf canopy. On a seasonal timescale, this acclimation occurs via large shifts in the macromolecular complexes of the photosynthetic system, within a photobiont population that shows no seasonal change in cell numbers. In this study, samples of L. pulmonaria were harvested in February and in May from a natural population near Sackville, New Brunswick, and subjected to two simulated intense seasonal changes: (1) early spring warming, simulated by a shift from high light at 5°C to high light at 16°C (February shift), and (2) late spring canopy closure, simulated by a shift from high light at 16°C to low light at 16°C (May shift). Thallus samples were collected daily throughout each week-long shift. There were no significant changes in photobiont cell population size or in the fraction of cells dividing during either shift. During the first day of the February temperature shift, there were, however, large changes in the pools of chlorophyll, the major light capture molecule in the photobionts, the PsbA (D1) core protein of photosystem II whose turnover is highly responsive to changing light and temperature, and the RbcL major subunit of the carbon-fixing RUBISCO enzyme whose levels correlate strongly with achieved photosynthesis in lichens. A static population of photobionts was therefore able to perform large and rapid macromolecular reallocations to cope with rapid environmental change. No significant changes were seen in the chlorophyll, photosystem II or RUBISCO pools across the May light shift, although seasonal-scale macromolecular reallocation does occur in response to decreased light in the summer.

The use of transgenic crops as feeds for ruminant animals has prompted study of the possible uptake of transgene fragments by ruminal micro-organisms and/or intestinal absorption of fragments surviving passage through the rumen. The persistence in buffered ruminal contents of seven different recombinant DNA fragments from GM rapeseed expressing the 5-enolpyruvylshikimate-3-phosphate synthase (EPSPS) transgene was tracked using PCR. Parental and transgenic (i.e. glyphosphate-tolerant; Roundup Ready®, Monsanto Company, St Louis, MO, USA) rapeseed were incubated for 0, 2, 4, 8, 12, 24 and 48 h as whole seeds, cracked seeds, rapeseed meal, and as pelleted, barley-based diets containing 65 g rapeseed meal/kg. The seven transgene fragments ranged from 179 to 527 bp and spanned the entire 1363 bp EPSPS transgene. A 180 bp ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) small subunit fragment and a 466 bp 16S rDNA fragment were used as controls for endogenous rapeseed DNA and bacterial DNA respectively. The limit of detection of the PCR assay, established using negative controls spiked with known quantities of DNA, was 12·5 pg. Production of gas and NH3 was monitored throughout the incubation and confirmed active in vitro fermentation. Bacterial DNA was detected in all sample types at all time points. Persistence patterns of endogenous (Rubisco) and recombinant (EPSPS) rapeseed DNA were inversely related to substrate digestibility (amplifiable for 48, 8 and 4 h in whole or cracked seeds, meal and diets respectively), but did not differ between parental and GM rapeseed, nor among fragments. Detection of fragments was representative of persistence of the whole transgene. No EPSPS fragments were amplifiable in microbial DNA, suggesting that transformation had not occurred during the 48 h incubation. Uptake of transgenic DNA fragments by ruminal bacteria is probably precluded or time-limited by rapid degradation of plant DNA upon plant cell lysis.

An elachistacean epiphyte, Neoleptonema yongpilii E.-Y. Lee & I.K. Lee, gen. et sp. nov., is reported from Korean coasts. The plants are distinguished by having unbranched assimilatory filaments with intercalary plurilocular sporangia as well as lateral plurilocular sporangia from the cortex. The new genus differs from the genera Leptonematella P. Silva and Halothrix Reinke by having pod-shaped plurilocular sporangia on the medulla, and from Elachista Duby and Proselachista Y.P. Lee & Garbary by having intercalary plurilocular sporangia and a poorly developed medulla. The phylogenetic relationships of Neoleptonema yongpilii were inferred from the spacer sequences between the genes coding for the large and small subunits of the RuBisCO gene. The new genus is a member of a poorly resolved clade consisting of several genera within the Elachistaceae, within which it is more closely related to Halothrix and Elachista nipponica than to Leptonematella.

Summary 11

I. INTRODUCTION 12

II. LICHEN BIONT CHARACTERISTICS 12

1. The mycobiont 12

2. The photobiont 13

III. LICHEN GROWTH 13

1. Allocation of resources 13

2. Growth rates and environmental limitations 14

3. Maintaining an optimal energy use efficiency 15

IV. CARBON ACQUISITION 16

1. Water relations 16

(a) Desiccation tolerance 16

(b) Activation upon re-hydration 17

(c) Diffusion of water and CO2 18

2. Photobiont CO2 fixation 20

(a) CO2 acquisition modes 20

(b) Significance of the CCM 22

3. Light and nitrogen relations 22

(a) The light-response curve 23

(b) Photosynthetic capacity 25

(c) Coping with high light 27

V. CARBON SINKS AND EXPENDITURES 28

1. Carbon translocation 28

2. Carbon sinks 29

3. Respiration 30

VI. CONCLUDING REMARKS 30

Acknowledgements 31

References 31

Lichens are nutritionally specialized fungi (the mycobiont component) that derive carbon and in some cases nitrogen from algal or cyanobacterial photobionts. The mycobiont and photobiont live together in an integrated thallus, but they lack specific tissue for the transport of metabolites and resources between them. Carbon is acquired through photosynthesis in the photobiont, which is active when the lichen is wet and exposed to light. Lichen photosynthesis is limited primarily by water, light and nitrogen, but can also be constrained by slow diffusion of CO2 within the wet thallus. The assimilated carbon is exported from photobiont to mycobiont, which also predominates in terms of biomass, and apparently regulates the size of the photobiont population. It has therefore generally been assumed that most of the carbon is used for growth and maintenance of the fungal hyphae. However, the extent of photobiont respiration in relation to mycobiont respiration has seldom been quantified; neither do we know the pool sizes of various carbon sinks within lichens. Owing to this lack of fundamental data we do not know whether, or how, carbohydrate resources are regulated to maintain an optimal balance between energy input and expenditures in these symbiotic organisms. This review summarizes data on growth, carbon gain and carbon expenditures in lichens, with particular emphasis on factors determining the photosynthetic capacity of their photobionts. An attempt is made to introduce an economic analysis of lichen growth processes, a view that has often been adopted in studies of higher plants. Areas in which more data are needed for the construction of a model on ‘lichen resource allocation patterns’ are discussed.

The effects of elevated concentrations of atmospheric carbon dioxide and ozone on diurnal patterns of photosynthesis have been investigated in field-grown spring wheat (Triticum aestivum). Plants cultivated under realistic agronomic conditions, in open-top chambers, were exposed from emergence to harvest to reciprocal combinations of two carbon dioxide and two ozone treatments: [CO2] at ambient (380 μmol mol−1, seasonal mean) or elevated (692 μmol mol−1) levels, [O3] at ambient (27 nmol mol−1, 7 hr seasonal mean) or elevated (61 nmol mol−1) levels. After anthesis, diurnal measurements were made of flag-leaf gas-exchange and in vitro Rubisco activity and content. Elevated [CO2] resulted in an increase in photoassimilation rate and a loss of excess Rubisco activity. Elevated [O3] caused a loss of Rubisco and a decline in photoassimilation rate late in flag-leaf development. Elevated [CO2] ameliorated O3 damage. The mechanisms of amelioration included a protective stomatal restriction of O3 flux to the mesophyll, and a compensatory effect of increased substrate on photoassimilation and photosynthetic control. However, the degree of protection and compensation appeared to be affected by the natural seasonal and diurnal variations in light, temperature and water status.

Photosynthetic acclimation and the interactions between carbon (C) and nitrogen (N) metabolism have been studied in the red macroalga Gracilaria sp. from Cádiz, Spain, cultured under different inorganic C and N levels. The use of chemostats and buffered medium allowed continuous restoration of the alkaline reserve and constancy of pH during the experiments. The N∶C ratios and phycobiliprotein, chlorophyll a and soluble protein contents decreased when Gracilaria sp. was grown at low N levels. Algae grown in a high inorganic C concentration (5% CO2) displayed a higher soluble carbohydrate concentration and maximum photosynthesis rates but a lower photosynthesic affinity for inorganic C, and lower phycobiliprotein and Rubisco contents, than those cultured at low inorganic C levels (air CO2). The inorganic C enrichment also affected the N uptake and assimilation in Gracilaria sp., causing a decrease in the N uptake rate even under conditions of N sufficiency. These results reflect the significant influence of the inorganic C growth regime on N assimilation in Gracilaria sp.

The study reported was conducted to establish the impacts of photochemical oxidants (primarily ambient ozone, O3) on the yield of watermelon (Citrullus lanatus) at a site on the east coast of Spain. Fruit yield and quality were monitored in plots established in a commercial watermelon field exposed, in open-top chambers (OTCs), to non-filtered air (NFA; near-ambient levels of ozone) or charcoal-filtered air (decreased levels of photochemical oxidants including O3; CFA), or to ambient air (AA), during the 1988 and 1989 growing seasons. Ambient levels of O3 were found to exceed present UN-ECE (United Nations Economic Commission for Europe; Convention on Long-Range Transboundary Air Pollution) critical level guidelines for the protection of crop yield by approx. twofold in 1988 and by approx. fivefold in 1989. Plants exposed to NFA and AA developed visible O3 injury on the upper surface of sun-exposed older leaves, and fruit yield (annual marketable fruit weight and number) was found to be depressed in OTCs ventilated with NFA in comparison with those receiving CFA. Consistent with inter-annual variations in O3 exposure, greater yield losses were experienced in 1989 (39%) than in 1988 (19%), an effect mediated predominantly by a decline in fruit number rather than average fruit weight. Exposure to ambient levels of ozone also slightly decreased fruit quality (4–8% decline in soluble solids content). Leaf gas exchange measurements made in the field in 1988 revealed effects of O3 on fruit yield and quality to be associated with a decline in the net CO2 assimilation rate per unit leaf area under light saturation (Asat) and stomatal conductance to water vapour (gs), and enhanced rates of dark respiration. A/ci curves (where A is the net CO2 assimilation rate per unit leaf area and ci is the mole fraction of CO2 in the leaf intercellular air space) constructed for plants grown in laboratory-based closed chambers, and exposed to an accumulated O3 exposure similar to that experienced by plants in the field, suggested that the likely cause of the decline in photosynthetic capacity was (1) a decrease in the amount and/or activity of Rubisco and (2) an impaired capacity for regeneration of ribulose 1,5-bisphosphate, which was not mediated through changes in the photochemical efficiency of photosystem II (Fv/Fm, where Fv is variable chlorophyll a fluorescence and Fm is maximum chlorophyll a fluorescence). No shift in the relative stomatal limitation to photosynthesis was observed under the influence of O3, suggesting that the decline in gs induced by the pollutant in both field and laboratory was the result, and not the cause, of the decrease in Asat. Ozone exposure also caused a decrease in C isotope discrimination (approx. 0.5‰), a shift that revealed a departure from predicted theory based on supporting leaf gas exchange measurements. The study demonstrates that ambient levels of photochemical oxidants on the Spanish Mediterranean coast are high enough to adversely influence the yield and physiology of an economically important crop grown in the region, and the magnitude of the effects was greater than would be predicted from exposure–yield-loss relationships for grain and fodder crops in central and northern Europe. Implications for the derivation of UN-ECE ozone standards are discussed, along with the mechanistic basis of the observed yield decreases.

The function of a leaf is photosynthesis, which requires the interception of light and access to atmospheric CO2 while controlling water loss. This paper examines the influence of leaf anatomy on both light capture and CO2 diffusion. As photosynthetic metabolism is spread between many chloroplasts, a leaf faces the challenge of matching light capture by a given chloroplast with the metabolic capacity of that chloroplast. Chloroplasts nearest the leaf surface receive the greatest irradiance and therefore absorb more light per unit chlorophyll than chloroplasts in the centre of a leaf. Electron transport and carbon fixation capacities per unit of chlorophyll decline with increasing depth in the leaf, to compensate for the decline in light absorbed per unit chlorophyll. Many key photosynthetic protein complexes in chloroplasts have nuclear encoded genetic information. Consequently, all chloroplasts within a given cell have a similar metabolic complement, which limits the potential gradient of photosynthetic capacity per unit chlorophyll across the leaf. A simple model couples light absorption through the leaf (based on the Beer–Lambert law) with the profile of chlorophyll through a leaf and the gradient in photosynthetic capacity. It is validated by comparison with 14CO2 fixation profiles through spinach leaves obtained in various studies. The model can account for published 14C fixation profiles obtained with blue, red and green light of different irradiances and white light applied in different combinations to the adaxial and abaxial surfaces of spinach leaves. The model confirms that spongy mesophyll increases the apparent extinction coefficient of chlorophyll compared to palisade tissue. The palisade tissue nearest the surface which receives light facilitates the penetration of light to a greater depth, while spongy mesophyll promotes scattering to enhance light absorption, thus reducing the gradient in light absorbed per unit chlorophyll through a leaf. CO2 fixation faces a diffusional limitation, which necessitates Rubisco to be spread evenly across the cell walls exposed to intercellular airspace. Mesophyll cell structure reflects the need to have a large cell surface per unit volume exposed to airspaces. The regular array of columnar cells in palisade tissue, or cell lobing in monocot leaves, results in greater exposed surface per unit tissue volume than spongy mesophyll. The exposed surface area per unit leaf area scales with photosynthetic capacity such that the difference in CO2 partial pressure between substomatal cavities and the sites of carboxylation within chloroplasts is, on average, independent of photosynthetic capacity of the leaf. However, Rubisco specific activity declines as the Rubisco content per unit leaf area increases due to greater internal diffusional limitations.

Cherry seedlings (Prunus avium) were grown from seed for two growing seasons in three ambient [CO2] (∼350 μmol mol−1) and three elevated [CO2] (ambient+∼350 μmol mol−1) open-top chambers, and in three outside blocks. A drying cycle was imposed in both the growing seasons to half the seedlings: days 69–115 in the first growing season, and in the second growing season days 212–251 on the same seedlings which had already experienced drought. Stomatal conductance was significantly reduced in elevated [CO2]-grown, unstressed seedlings in both the first and second growing seasons, but was not caused by a decrease in stomatal density. Droughted seedlings showed little or no reduction in stomatal conductance in response to elevated [CO2]. However, stomatal conductance was highly correlated with soil water status. Photosynthetic rate increased significantly in response to elevated [CO2] in both water regimes, leading to improvement in instantaneous transpiration efficiency over the whole duration of the experiment, but there was no relationship between instantaneous transpiration efficiency and long-term water use efficiency. The Amax was strongly reduced in the second growing season, but unaffected by [CO2] treatment. Although photosynthetic rate was not down-regulated, Rubisco activity was decreased by elevated [CO2], possibly because of the increased leaf carbon: nitrogen ratio which had occurred by the ends of the two growing seasons. Elevated [CO2] did not improve plant water relations (for example, bulk leaf – water potential, osmotic potentials at full and zero turgor, relative water content at zero turgor, bulk modulus of elasticity of the cell) and thus did not increase water-stress tolerance of cherry seedlings.

Nitrogen (N) and phosphorus (P) limitation affect the photosynthetic apparatus of Dunaliella tertiolecta in markedly different ways. When grown at 0·25 d−1 (18 % of the resource-saturated maximum rate, μmax=1·39 d−1) in chemostat cultures, N- and P-limited cells were chlorotic relative to nutrient-replete controls. The lutein-to-chlorophyll a ratio increased under both N and P limitation, whereas the neoxanthin-to-chlorophyll a ratio increased only under P limitation. The ratio of accessory photoprotective pigments (α- and β-carotene) to chlorophyll a increased under N-limited conditions. Despite differences in accessory pigment complement, chlorophyll a-specific light absorption coefficients of N- and P-limited cultures did not differ significantly, and were greater than in nutrient-replete conditions. In contrast, the initial slope of the photosynthesis–irradiance (PE) response curve (αChl) declined under nutrient-limiting conditions. There were slight reductions in the maximum quantum efficiency of photosynthesis (ϕm) in N- and P-limited cells. Reductions in ϕm were accompanied by reductions in the ratio of variable to maximum fluorescence (Fv/Fm), and the ratio of the photosystem II reaction centre protein D1 to the large subunit of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco). Differences in light-saturated gross photosynthesis rate, as measured by light/dark oxygen exchange, could be accounted for by changes in the abundance of the carboxylating enzyme Rubisco. Oxygen exchange and 14CO2 assimilation appeared to measure different processes in P-limited and N- limited cultures. At light-saturation, 14C-bicarbonate assimilation approximated gross photosynthesis (as measured by light/dark oxygen exchange) in P-limited cultures. In contrast, 14C-bicarbonate assimilation approximated net photosynthesis in N-limited cultures. When all culture conditions were compared, there was linear covariation of the rates of reductant supply via light absorption and photochemical charge separation with the rates of reductant demand for CO2 fixation and NO3− reduction.

Cloned cuttings of Betula pendula Roth were grown in field fumigation chambers at Birmensdorf throughout one growing season in filtered air with either <3 (control) or 90/40 nl l−1 O3 (day/night; ozone generated from pure oxygen). Each ozone regime was split into high and low soil nutrient regimes by watering plants with either a 0·05% or a 0·005% solution of a fertilizer which contained macronutrients and micronutrients.

Fertilization had a strong effect on plant growth, enzyme activities and the expression of ozone-induced effects at the biochemical level. The activities of PEPC and Rubisco were enhanced about threefold in the plants with high fertilization (HF). Significant effects of ozone were in most cases found only in the older leaves of the plants with low fertilization (LF). There, sucrose, glucose and fructose levels were enhanced. In both fertilization treatments, the number of starch granules along the minor veins was increased. These ozone effects point to a decreased or inhibited phloem loading. The increased PEPC activity and the enhanced malate levels in the ozone-exposed plants might be the result of a redirection of carbon flow from sucrose synthesis and translocation towards anapleurotic processes, which can feed detoxification and repair of ozone injury as indicated by enhanced respiration. These findings agree well with the observed effects of ozone in lowering the root[ratio ]shoot biomass ratio. Although there was a marked reduction in the O3/LF plants, O3/HF plants showed no significant response. Inositol was decreased under ozone exposure in both fertilizer treatments, contrasting with the pattern for carbohydrates.

These results demonstrate the role of fertilization as an important modifier of ozone-induced effects at the plant biochemical level. Well fertilized plants appear to cope better with the impact of ozone on metabolism.