Impact of Atmospheric Sulfur Deposition on Sulfur Metabolism in Plants: H2S as Sulfur Source for Sulfur Deprived Brassica oleracea L.

Brassica oleracea L. was rather insensitive to atmospheric H₂S: growth was only negatively affected at ≥0.4 μl I^?1. Shoots formed a sink for H₂S and the uptake rate showed saturation kinetics with respect to the atmospheric concentration. The H₂S uptake rate was high in comparison with other species, which may reflect the high sulfur need of Brassica. The net uptake of sulfate by roots of hydroponically grown plants was substantially reduced after one week of exposure to 0.25 μl l^?1 H₂S, indicating that plants switched in part from sulfate to H₂S as sulfur source for plant growth. Plants were sulfur deficient after two weeks of sulfur deprivation, illustrated by reduced growth, which was more pronounced for shoots than for roots, and in enhanced shoot dry matter content. The latter could for the greater part be attributed to enhanced levels of soluble sugars and starch. Sulfur deficiency was further characterized by a low pigment content, extremely low levels of sulfate and water-soluble non-protein thiols, and by enhanced levels of nitrate and free amino acids, particularly in the shoots. Furthermore, sulfur deficient plants contained a lower total lipid content in shoots, whereas its content in roots was unaffected. The level of sulfolipids was decreased in both roots and shoots. When sulfur deprived plants were exposed to 0.25 μl I^?1 H₂S for one week, all sulfur deficiency symptoms were abolished and growth was restored. Furthermore, plants were able to grow with 0.4 μl I^?1 H₂S as the sole sulfur source. Water-soluble non-protein thiol content was enhanced in both shoots and roots of H₂S exposed plants, irrespective of the sulfate supply to the roots, whereas plants grown with H₂S as sole sulfur source contained very low sulfate levels. The interaction between atmospheric and pedospheric sulfur nutrition in plants is discussed.     

Diagnostic parameters for selecting against novel spruce (Picea abies) decline: I. Tree morphology and photosynthesis response to acute SO2 exposures

The dose- and time-response effects of single 4-h day-exposures to 0.50, 0.79, 1.28, 1.58, 2.38 or 3.35 μl l^?1 (ppm) SO₂ followed by single 3-h night-exposures of 0.60, 0.87, 1.54, 1.91, 2.91 or 3.98 μl l^?1 SO₂ on photosynthesis, transpiration and dark respiration were examined for nine East European (Carpatho-Ukrainian, ‘Rachovo’) half-sib families and for two populations, one from the FRG (‘Westerhof’) and one from the GDR (‘Schmiedefeld’) of Norway spruce [Picea abies (L.) Karst.], all in their 4th growing season. Even the lowest SO₂ concentration reduced photosynthesis and transpiration within 1 h. Photosynthesis of the different spruce types was affected significantly differently, the most sensitive spruce being suppressed 2.5 times more than the most tolerant spruce. ‘Westerhof’ was more resistant to SO₂ than the average ‘Rachovo’ half-sibs. Neither transpiration (stomatal reaction), which was affected alike by all SO₂ concentrations, nor SO₂ uptake, explained adequately the effects on photosynthesis. Night transpiration, but not dark respiratin, was stimulated by night SO₂ preceded by day SO₂ exposure. The gradient of different SO₂ sensitivities among young trees from the half-sib families demonstrated a significant negative correlation with the gradient of different sensitivities to novel decline symptoms of their parents growing in a rural seed orchard in Denmark, and with the gradients of four morphology parameters, (height, branching, branch density and the number of Lammas shoots) of the young trees, which in turn demonstrated a positive correlation with decline sensitivity in the seed orchard. The relative photosynthesis sensitivity and the morphology of half-sibs may serve as diagnostic parameters for laboratory selection of the most resistent trees to novel spruce decline in the field. There was a positive correlation between SO₂ induced scorching of Lammas shoots and the inhibition of photosynthesis, but not between the severity of SO₂ scorching and symptoms of novel spruce decline. The two visible types of symptoms looked very different.     

Physiological effects of a long term exposure to low concentrations of NH3, NO2 and SO2 on Douglas fir (Pseudotsuga menziesii)

The above-ground parts of two years old seedlings of Douglas fir (Pseudotsuga menziesii) were exposed to filtered air, NH₃, NO₂₊, SO₂ (66, 96 and 95 μg m^?3, respectively), to a mixture of NO₂+NH₃ (55 + 82 μg m^?3) or SO₂+NO₂ (128 + 129 μg m^?3), for 8 months in fumigation chambers. Both chlorophyll fluorescence and gas exchange measurements were carried out on shoots which had sprouted at the beginning of the exposure period. The chlorophyll fluorescence measurements were performed after 3 and 5 months of exposure (average shoot age 70 and 140 days, respectively). Light response curves of electron transport rate (J) were determined, in which J was deduced from chlorophyll fluorescence. In addition, light response curves of net CO₂ assimilation were determined after 5 months of exposure. After 3 months of exposure (average shoot age 70 days) all exposure treatments showed a lower maximum electron transport rate (J_max) as compared to the control shoots (filtered air). A large reduction (45%) was observed for shoots exposed to SO₂+NO₂. During the exposure period between 3 and 5 months (average shoot age 70 and 140 days, respectively) a decrease of J_max was observed for all treatments. J_max had further declined some time after termination of the exposure, when average shoot age was 310 days. Shoots exposed to SO₂ and SO₂+NO₂ also showed a reduction in maximum net CO₂ assimilation (P_max) as compared to the control shoots. However, shoots exposed to NO₂ showed no reduction and even a higher P_max was observed for shoots exposed to NH₃ or NO₂+NH₃. Needles of these treatments also showed a higher chlorophyll content which might explain the contradictory results obtained for these treatments: the increased amount of photosynthetic units counteracts the reduction in J_max and consequently no reduction in P_max is measured. Shoots exposed to SO₂ and SO₂+NO₂ also showed a reduction in maximum stomatal conductance (g_s). However, the stomatal opening was larger than could be expected on basis of their (maximum) CO₂ assimilation rate. Consequently, water use efficiency of these shoots was lower than that of the control shoots. Also shoots exposed to NO₂ had a lower water use efficiency due to a significantly higher maximum g_s. Shoots exposed to NH₃ showed a high transpiration rate in the dark, indicating imperfect stomatal closure.     

Acquisition and within-plant allocation of 13C and 15N in CO2-enriched Quercus robur plants

We assessed the effects of doubling atmospheric CO₂ concentration, [CO₂], on C and N allocation within pedunculate oak plants (Quercus robur L.) grown in containers under optimal water supply. A short-term dual ¹³CO₂ and ¹⁵NO₃^? labelling experiment was carried out when the plants had formed their third growing flush. The 22-week exposure to 700 μl l^?1 [CO₂] stimulated plant growth and biomass accumulation (+53% as compared with the 350 μl l^?1 [CO₂] treatment) but decreased the root/shoot biomass ratio (-23%) and specific leaf area (-18%). Moreover, there was an increase in net CO₂ assimilation rate (+37% on a leaf dry weight basis; +71% on a leaf area basis), and a decrease in both above- and below-ground CO₂ respiration rates (-32 and -26%, respectively, on a dry mass basis) under elevated [CO₂]. ¹³C acquisition, expressed on a plant mass basis or on a plant leaf area basis, was also markedly stimulated under elevated [CO₂] both after the 12-h ¹³CO₂ pulse phase and after the 60-h chase phase. Plant N content was increased under elevated CO₂ (+36%), but not enough to compensate for the increase in plant C content (+53%). Thus, the plant C/N ratio was increased (+13%) and plant N concentration was decreased (-11%). There was no effect of elevated [CO₂] on fine root-specific ¹⁵N uptake (amount of recently assimilated ¹⁵N per unit fine root dry mass), suggesting that modifications of plant N pools were merely linked to root size and not to root function. N concentration was decreased in the leaves of the first and second growing flushes and in the coarse roots, whereas it was unaffected by [CO₂] in the stem and in the actively growing organs (fine roots and leaves of the third growth flush). Furthermore, leaf N content per unit area was unaffected by [CO₂]. These results are consistent with the short-term optimization of N distribution within the plants with respect to growth and photosynthesis. Such an optimization might be achieved at the expense of the N pools in storage compartments (coarse roots, leaves of the first and second growth flushes). After the 60-h ¹³C chase phase, leaves of the first and second growth flushes were almost completely depleted in recent ¹³C under ambient [CO₂], whereas these leaves retained important amounts of recently assimilated ¹³C (carbohydrate reserves?) under elevated [CO₂].     

Elevated CO2 enhances plant growth in droughted N2-fixing alfalfa without improving water status

The long-term interaction between elevated CO₂ and soil water deficit was analysed in N₂-fixing alfalfa plants in order to assess the possible drought tolerance effect of CO₂. Elevated CO₂ could delay the onset of drought stress by decreasing transpiration rates, but this effect was avoided by subjecting plants to the same soil water content. Nodulated alfalfa plants subjected to ambient (400 μmol mol^?1) or elevated (700 μmol mol^?1) CO₂ were either well watered or partially watered by restricting water to obtain 30% of the water content at field capacity (ampproximately 0.55 g water cm^?3). The negative effects of soil water deficit on plant growth were counterbalanced by elevated CO₂. In droughted plants, elevated CO₂ stimulated carbon fixation and, as a result, biomass production was even greater than in well-watered plants grown in ambient CO₂. Below-ground production was preferentially stimulated by elevated CO₂ in droughted plants, increasing nodule biomass production and the availability of photosynthates to the nodules. As a result, total nitrogen content in droughted plants was higher than in well-watered plants grown in ambient CO₂. The beneficial effect of elevated CO₂ was not correlated with a better plant water status. It is concluded that elevated CO₂ enhances growth of droughted plants by stimulating carbon fixation, preferentially increasing the availability of photosynthates to below-ground production (roots and nodules) without improving water status. This means that elevated CO₂ enhances the ability to produce more biomass in N₂-fixing alfalfa under given soil water stress, improving drought tolerance.     

Diagnostic parameters for selecting against novel spruce (Picea abies) decline: III. Response of photosynthesis and transpiration to O3 exposures

The dose- and time-response effects of single 4 h day-time exposures of 0.064, 0.166, 0.336, 0.452 or 0.693 μl l^?1 (ppm) O₃ followed by single 4 h night-time exposures of 0.078, 0.198, 0.378, 0.502 or 0.747 μl l^?1 O₃ on photosynthesis, transpiration and dark respiration were examined for nine Carpatho-Ukrainian (‘Rachovo’) half-sib families and for two populations. ‘Westerhof’ from the FRG and ‘Schmiedefeld’ from the GDR, of Norway spruce [Picea abies (L.) Karst.], all in their 4th growing season. Needles were scorched by 4 h exposures to 0.336 μl l^?1 O₃ and higher. The lag before photosynthesis and transpiration responded significantly to O₃ decline took from a few minutes at the highest concentration to several hours at the lower concentrations. Recovery of photosynthesis and transpiration was absent or extremely slow. Photosynthesis of the different spruce types was affected significantly differently, the most sensitive spruce having its photosynthesis suppressed 1.9 times and its transpiration 1.6 times more than the most tolerant spruce. The physiological responses of ‘Westerhof’ were less sensitive than the average ‘Rachovo’ half-sibs. Neither night transpiration nor dark respiration were affected by high doses of night O₃, preceded by day O₃ exposures. The gradients of different photosynthesis and transpiration sensitivities of the young half-sibs (and ‘Westerhof’) demonstrated a significant, positive, mutual correlation, and significant positive correlations with the gradient of novel decline symptoms of their parents growing in Danish forests. The relative photosynthesis and transpiration sensitivities may thus serve as diagnostic parameters in laboratory tests for selection against novel spruce decline.     

Diagnostic parameters for selecting against novel spruce (Picea abies) decline: II. Response of photosynthesis and transpiration to acute NOx exposures

The dose- and time-response effects of sequential 3 h+3 h NO→NO₂ day time exposures [0–9 μl l^?1 (ppm) NO, 0–7.5 μl l^?1 NO₂] followed by 3 h+3 h NO→NO₂ night-time exposures (0–9.5 μl l^?1 NO, 0–9 μl l^?1 NO₂) on photosynthesis, transpiration and dark respiration were examined for nine Carpatho-Ukrainian (‘Rachovo’) half-sib families and for two populations, one from the FRG (‘Westerhof’) and one from the GDR (‘Schmiedefeld’) of Norway spruce [Picea abies (L.) Karst.], all in their 4th growing season. In a second exposure series the exposure sequence was reversed. None of the treatments induced needle scorching. The higher NO_x (NO or NO₂) concentrations reduced photosynthesis and transpiration within 1 h. The physiology of the different spruce types was affected significantly differently, the most sensitive spruce having its photosynthesis suppressed 6.6 times and its transpiration 5.5 times more than the most tolerant. ‘Westerhof’ was more sensitive to NO₂ than the average ‘Rachovo’ half-sibs. The gradients of different photosynthesis and transpiration sensitivities among the half-sibs (and ‘Westerhof’) demonstrated a significant, positive, mutual correlation, but significant negative correlations with the gradient of novel decline symptoms among their parents growing in Danish forests. The relative photosynthesis and transpiration sensitivies may thus serve as diagnostic parameters for laboratory selection of the most resistant trees to novel spruce decline. The average NO₂ flux density was three times larger than the average NO flux density. Only for NO₂ and in light was stomatal NO_x uptake larger than the total NO_x uptake. Both night transpiration and dark respiration were stimulated by high concentrations of night NO_x, preceded by day NO_x exposures.     

Differential Na2SO4 tolerance in tobacco plants regenerated from Na2SO4-grown callus

Abstract The regenerated shoots from sodium sulphate (Na₂SO₄) grown callus of tobacco (Nicotiana tabacum L. cv. Wisconsin 38) were evaluated for Na₂SO₄ tolerance based on shoot proliferation and rooting in vitro, and seed germination in vivo in response to Na₂SO₄. An increase in Na₂SO₄ concentration resulted in significantly decreasing shoot fresh weight, number of shoots, shoot length and leaf size, and increasing per cent shoot dry weight of both control and Na₂SO₄-grown cultures. In rooting, shoots of Na₂SO₄-grown cultures exhibited the highest per cent rooting (85%) in the presence of 1% w/v Na₂SO₄. However, per cent rooting, root number per rooted cutting and root fresh weight decreased significantly with increasing Na₂SO₄ concentration when shoots were transferred to the medium in the absence of Na₂SO₄ for 4-monthly passages. Following acclimatization of the rooted shoots of Na₂SO₄-grown cultures, phenotypic variation was observed during growth and development. There were 13.2% sterile plants. Fertile plants were sorted into normal (N), tolerant (T), and sensitive (S) categories and the respective percentages of plants were 31.6, 44.7 and 10.5, based on per cent germination, germination velocity index and seedling survival to Na₂SO₄. The response of N, T and S types to Na₂SO₄ in subsequent shoot proliferation was similar to that of seed germination.     

Response of wheat canopy CO2 and water gas-exchange to soil water content under ambient and elevated CO2

The nature of the interaction between drought and elevated CO₂ partial pressure (pC_a) is critically important for the effects of global change on crops. Some crop models assume that the relative responses of transpiration and photosynthesis to soil water deficit are unaltered by elevated pC_a, while others predict decreased sensitivity to drought at elevated pC_a. These assumptions were tested by measuring canopy photosynthesis and transpiration in spring wheat (cv. Minaret) stands grown in boxes with 100 L rooting volume. Plants were grown under controlled environments with constant light (300 µmol m^?2 s^?1) at ambient (36 Pa) or elevated (68 Pa) pC_a and were well watered throughout growth or had a controlled decline in soil water starting at ear emergence. Drought decreased final aboveground biomass (?15%) and grain yield (?19%) while elevated pC_a increased biomass (+24%) and grain yield (+29%) and there was no significant interaction. Elevated pC_a increased canopy photosynthesis by 15% on average for both water regimes and increased dark respiration per unit ground area in well‐watered plants, but not drought‐grown ones. Canopy transpiration and photosynthesis were decreased in drought‐grown plants relative to well‐watered plants after about 20–25 days from the start of the drought. Elevated pC_a decreased transpiration only slightly during drought, but canopy photosynthesis continued to be stimulated so that net growth per unit water transpired increased by 21%. The effect of drought on canopy photosynthesis was not the consequence of a loss of photosynthetic capacity initially, as photosynthesis continued to be stimulated proportionately by a fixed increase in irradiance. Drought began to decrease canopy transpiration below a relative plant‐available soil water content of 0.6 and canopy photosynthesis and growth below 0.4. The shape of these responses were unaffected by pC_a, supporting the simple assumption used in some models that they are independent of pC_a.     

Plant responses to H2S and SO2 fumigation. II. Differences in metabolism of H2S and SO2 in spinach

Fumigation of spinach (Spinacia oleracea L. cvs Estivato and Monosa) with H₂S or SO, for 1 to 6 days resulted in accumulation of sulfhydryl (SH) compounds in the shoots of both H₂S- and SO₂-exposed plants. The sulfate concentration in shoots of SO₂-exposed plants increased linearly with time. SH accumulation showed saturation kinetics as a function of time as well as H₂S concentration, ascribed to the internal H₂S concentration in the plant and the availability of substrates for glutathione synthesis, respectively. SH compounds accumulated more at lower exposure temperatures, whereas sulfate accumulation was more pronounced at higher temperatures. These results are discussed in relation to the possible foliar uptake of H₂S and SO₂, the temperature dependence of uptake and the water solubility of these gases. The possibility of SO₂-induced H₂S emission rather than sulfate accumulation as a source for SH accumulation is also discussed. Cessation of fumigation resulted in a decrease in SH compounds and sulfate content that could be accounted for by sulfur metabolism and growth, respectively.     

Physiological effects of long term exposure to low concentrations of SO2 and NH3 on poplar leaves

Shoots of poplar (Populus euramericana L. cv. Flevo) were exposed to filtered air, SO₂, NH₃ or a mixture of SO₂ and NH₃ for 7 weeks in fumigation chambers. After this exposure gas exchange measurements were carried out using a leaf chamber. As compared to leaves exposed to filtered air, leaves pretreated with 112 μg m^?3 SO₂ showed a small reduction in maximum CO₂ assimilation rate (P_max) and stomatal conductance (g_s). They also showed a slightly higher quantum yield and dark respiration. In addition, the fluorescence measurements indicated that the Calvin cycle of the leaves pretreated with 112 μg m^?3 SO₂ was more rapidly activated after transition from dark to light. An exposure to 64 μg m^?3 NH₃ had a positive effect on P_max, stomatal conductance and NH₃ uptake of the leaves. This positive effect was counteracted by an SO₂ concentration of 45 μg m^?3. The exposure treatments appeared to have no effect on the relationship between net CO₂-assimilation and g_s. Also, no injury of the leaf cuticle or of epidermal cells was observed. Resistance analysis showed that NH₃ transfer into the leaf can be estimated from data on the boundary layer and stomatal resistance for H₂O transfer and NH₃ concentration at the leaf surface, irrespective of whether the leaves are exposed for a short or long time to NH₃ or to a mixture of NH₃ and SO₂. In contrast SO₂ uptake into the leaves was only partly correlated to the stomatal resistance. The results suggest a large additional uptake of this gas by the leaves. The possibility of a difference in path length between SO₂ and H₂O molecules is proposed.     

Comparative ecophysiology of CAM and C3 bromeliads. III. Environmental influences on CO2 assimilation and transpiration

Abstract Field measurements of the gas exchange of epiphytic bromeliads were made during the dry season in Trinidad in order to compare carbon assimilation with water use in CAM and C₃ photosynthesis. The expression of CAM was found to be directly influenced by habitat and microclimate. The timing of nocturnal CO₂ uptake was restricted to the end of the dark period in plants found at drier habitats, and stomatal conductance in two CAM species was found to respond directly to humidity or temperature. Total night-time CO₂ uptake, when compared with malic-acid formation (measured as the dawn-dusk difference in acidity, ΔH⁺), could only account for 10–40% of the total ΔH⁺ accumulated. The remaining malic acid must have been derived from the refixation of respired CO₂ (recycling). Within the genus Aechmea (12 samples from four species), recycling was significantly correlated with night temperature at the six sample sites. Recycling was lowest in A. fendleri (54% of ΔH⁺ derived from respired CO₂), a CAM bromeliad with little water-storage parenchyma that is restricted to wetter, cooler regions of Trinidad. Gas-exchange rates of C₃ bromeliads were found to be similar to those of the CAM bromeliads, with CO₂ uptake from 1 to 3 μmol m^?2 s^?1 and stomatal conductances generally up to 100 mmol m^?2 s^?1. The midday depression of photosynthesis occurred in exposed habitats, although photosynthetically active radiation (PAR) limited photosynthesis in shaded habitats. CO₂ uptake of the C₃ bromeliad Guzmania lingulata was saturated at around 500 μmol m^?2 s^?1 PAR, suggesting that epiphytic plants found in the shaded forest understorey are shade-tolerant rather than shade-demanding. Transpiration ratios (TR) during CO₂ fixation in CAM (Phase I and IV) and C₃ bromeliads were compared at different sites in order to assess the efficiency of water utilization. For the epiphytes displaying marked uptake of CO₂, TR were found to be lower than many previously published values. In addition, the average TR values were very similar for dark CO₂ uptake in CAM (42 ± 41, n= 12), Phase IV of CAM (69 ± 36, n= 3) and for C₃ photosynthesis (99 ± 73, n= 4) in these plants. It appears that recycling of respired CO₂ by CAM bromeliads and efficient use of water in all phases of CO₂ uptake are physiological adaptations of bromeliads to arid microclimates in the humid tropics.     

Cadmium-induced subcellular accumulation of O2·− and H2O2 in pea leaves

Cadmium is a toxic metal that produces disturbances in plant antioxidant defences giving rise to oxidative stress. The effect of this metal on H₂O₂ and O₂^·? production was studied in leaves from pea plants growth for 2 weeks with 50 µm Cd, by histochemistry with diaminobenzidine (DAB) and nitroblue tetrazolium (NBT), respectively. The subcellular localization of these reactive oxygen species (ROS) was studied by cytochemistry with CeCl₃ and Mn/DAB staining for H₂O₂ and O₂^·?, respectively, followed by electron microscopy observation. In leaves from pea plants grown with 50 µm CdCl₂ a rise of six times in the H₂O₂ content took place in comparison with control plants, and the accumulation of H₂O₂ was observed mainly in the plasma membrane of transfer, mesophyll and epidermal cells, as well as in the tonoplast of bundle sheath cells. In mesophyll cells a small accumulation of H₂O₂ was observed in mitochondria and peroxisomes. Experiments with inhibitors suggested that the main source of H₂O₂ could be a NADPH oxidase. The subcellular localization of O₂^·? production was demonstrated in the tonoplast of bundle sheath cells, and plasma membrane from mesophyll cells. The Cd‐induced production of the ROS, H₂O₂ and O₂^·?, could be attributed to the phytotoxic effect of Cd, but lower levels of ROS could function as signal molecules in the induction of defence genes against Cd toxicity. Treatment of leaves from Cd‐grown plants with different effectors and inhibitors showed that ROS production was regulated by different processes involving protein phosphatases, Ca²⁺ channels, and cGMP.     

Growth and Major Nutrient Concentrations in Brassica campestris Supplied with Different NH4^＋/NO3 Ratios

In the present study, we investigated whether growth and main nutrient ion concentrations of cabbage （Brassica campestris L.） could be increased when plants were subjected to different NH4^＋/NO3- ratios. Cabbage seedlings were grown in a greenhouse in nutrient solutions with five NH4^＋/NO3- ratios （1：0; 0.75：0.25; 0.5：0.5; 0.25：0.75; and 0：1）. The results showed that cabbage growth was reduced by 87% when the proportion of NH4^＋-N in the nutrient solution was more than 75% compared with a ratio NH4^＋/NO3- of 0.5：0.5 35 d after transplanting, suggesting a possible toxicity due to the accumulation of a large amount of free ammonia in the leaves. When the NH4＋/NO3- ratio was 0.5：0.5, fresh seedling weight, root length, and H2PO4- （P）, K^＋, Ca^2＋, and Mg^2＋ concentrations were all higher than those in plants grown under other NH4^＋/NO3- ratios. The nitrate concentration in the leaves was the lowest in plants grown at 0.5： 0.5 NH4^＋/NO3-. The present results indicate that an appropriate NH4^＋/NO3- ratio improves the absorption of other nutrients and maintains a suitable proportion of N assimilation and storage that should benefit plant growth and the quality of cabbage as a vegetable.     

Can plants exposed to SO2 excrete sulfuric acid through the roots?

Hydroponically grown pea plants (Pisum sativum L., cv. Kleine Rheinländerin) and barley seedlings (Hordeum vulgare L., cv. Gerbel) were fumigated for several days with 1 or 2 μl l^?1 SO₂. Both species accumulated sulfate during fumigation, although the nutrient medium lacked sulfate. In pea, SO₂-dependent sulfate accumulation in different plant parts accounted for 60 percent of the SO₂ sulfur which, as calculated from a determination of boundary and stomatal flux resistances had entered the leaves. Up to 55% of the air-borne sulfate was translocated from pea leaves to roots during the period of fumigation, but no or only little sulfate was excreted into the nutrient solution. In contrast, barley retained sulfate in the leaves, and sulfate translocation from shoot to the root system could not be observed. In both species, protons were excreted by the roots. In fumigated plants, proton loss was higher than in untreated controls in pea, but not in barley. In pea, SO₂-dependent proton loss into the medium accounted for up to 50% of the sulfuric acid formed from SO₂. Proton excretion was strongly dependent on potassium availability in the nutrient medium. Cation uptake by the plants during fumigation was sufficient to compensate for proton loss, suggesting proton/cation exchange at the interface between root and medium. We conclude that by oxidation to sulfuric acid, plants are capable of detoxifying SO₂ taken up by the leaves. Depending on plant species, either both protons and sulfate anions can be exported from the leaves, or the proton load on leaf cells can be relieved by proton/cation exchange at the plasmalemma. Finally, the problem of airborne plant acidification may be solved by proton/cation exchange at the level of roots. The burden of acidification is then shifted from the plant to the nutrient medium. Appreciable amounts of sulfate can be excreted neither by pea nor by barley plants.     

Effects of elevated CO2, drought and temperature on the water relations and gas exchange of groundnut (Arachis hypogaea) stands grown in controlled environment glasshouses

Stands of groundnut (Arachis hypogaea L. cv. Kadiri‐3) were grown in controlled environment glasshouses at mean atmospheric CO₂ concentrations of 375 or 700 μmol mol^?1 and daily mean air temperatures of 28 or 32°C on irrigated or drying soil profiles. Leaf water (Ψ_l) and solute potential (Ψ_s), relative water content (RWC), stomatal conductance (g_l) and net photosynthesis (P_n) were measured at midday for the youngest mature leaf throughout the growing season. Elevated CO₂ and temperature had no detectable effect on the water relations of irrigated plants, but higher values of RWC, Ψ_l and Ψ_s were maintained for longer under elevated CO₂ during progressive drought. Turgor potential (Ψ_p) reached zero when Ψ_l declined to ?1.6 to ?1.8 MPa in all treatments; turgor was lost sooner when droughted plants were grown under ambient CO₂. A 4°C increase in mean air temperature had no effect on Ψ_s in droughted plants, but elicited a small increase in Ψ_l; midday g_l values were lower under elevated than under ambient CO₂, and Ψ_l and g_l declined below ?1.5 MPa and 0.25 cm s^?1, respectively, as the soil dried. Despite the low g_l values recorded for droughted plants late in the season, P_n was maintained under elevated CO₂, but declined to zero 3 weeks before final harvest under ambient CO₂. Concurrent reductions in g_l and increases in water use efficiency under elevated CO₂ prolonged photosynthetic activity during drought and increased pod yields relative to plants grown under ambient CO₂. The implications of future increases in atmospheric CO₂ for the productivity of indeterminate C₃ crops grown in rainfed subsistence agricultural systems in the semi‐arid tropics are discussed.     

The relationship between absorption of sulfur dioxide (SO2) and inhibition of photosynthesis in several plants

Photosynthetic rate, transpiration rate and SO₂ absorption rate were simultaneously measured under exposure to SO₂ (0.1–1.0 μl l ^?1) for 5 or 8 hr in six species belonging to C₄ or C₃ plants (Zea mays, Sorghum vulgare, Amaranthus tricolor, Oryza sativa, Avena sativa andHelianthus annuus). Distinct interspecific differences were found as to the extent of inhibition of photosynthetic rate. Calculation of diffusive resistance to H₂O(r) and SO₂(r′) showed that the ratio of r′/r was 1.9 irrespective of species and coincided well with the theoretical value based on molecular diffusion. Thus it was made clear that the absorption of SO₂ was dependent upon the gas exchange capacity of leaf blade. Using the ratio of r′/r the rate of SO₂ absorption could be calculated from transpiration rate and was compared with the inhibition rate of photosynthesis. In three C₄ species, the inhibition of photosynthesis increased linearly with the amount of SO₂ absorbed during a 5-hour period. The pattern of inhibition of photosynthesis inA. sativa andH. annuus among C₃ species was similar to that of C₄ species until the amount of SO₂ absorbed reached 60 mg-SO₂ m^?2 above which the inhibition abruptly increased. The inhibition of photosynthesis inO. sativa was exceptionally severe even with only a small amount of SO₂ absorbed.     

Gross primary productivity and transpiration flux of the Australian vegetation from 1788 to 1988 AD: effects of CO2 and land use change

We present a novel approach to estimating the transpiration flux and gross primary productivity (GPP) from Normalized Difference Vegetation Index, leaf functional types, and readily available climate data. We use this approach to explore the impact of variations in the concentration of carbon dioxide in the atmosphere ([CO₂]) and consequent predicted changes in vegetation cover, on the transpiration flux and GPP. There was a near 1 : 1 relationship between GPP estimated with this transpiration flux approach and that estimated using a radiation‐use efficiency (RUE) approach. Model estimates are presented for the Australian continent under three vegetation–[CO₂] scenarios: the present vegetation and hypothetical ‘natural’ vegetation cover with atmospheric CO₂ concentration ([CO₂]) of 350 μmol mol^?1 (pveg₃₅₀ and nveg₃₅₀), and for the ‘natural’ vegetation with [CO₂] 280 μmol mol^?1 (nveg₂₈₀). Estimated continental GPP is 6.5, 6.3 and 4.3 Gt C yr^?1 for pveg₃₅₀, nveg₃₅₀ and nveg₂₈₀, respectively_. The corresponding transpiration fluxes are 232, 224 and 190 mm H₂O yr^?1. The contribution of the raingreen and evergreen components of the canopy to these fluxes are also estimated.     

The impact of elevated carbon dioxide on the growth and gas exchange of three C4 species differing in CO2 leak rates

Recent work has suggested that the photosynthetic rate of certain C₄ species can be stimulated by increasing CO₂ concentration, [CO₂], even under optimal water and nutrients. To determine the basis for the observed photosynthetic stimulation, we tested the hypothesis that the CO₂ leak rate from the bundle sheath would be directly related to any observed stimulation in single leaf photosynthesis at double the current [CO₂]. Three C₄ species that differed in the reported degree of bundle sheath leakiness to CO₂, Flaveria trinervia, Panicum miliaceum, and Panicum maximum, were grown for 31–48 days after sowing at a [CO₂] of 350 μl l^?1 (ambient) or 700 μl l^?1 (elevated). Assimilation as a function of increasing [CO₂] at high photosynthetic photon flux density (PPFD, 1 600 μmol m^?2 s^?1) indicated that leaf photosynthesis was not saturated under current ambient [CO₂] for any of the three C₄ species. Assimilation as a function of increasing PPFD also indicated that the response of leaf photosynthesis to elevated [CO₂] was light dependent for all three C₄ species. The stimulation of leaf photosynthesis at elevated [CO₂] was not associated with previously published values of CO₂ leak rates from the bundle sheath, changes in the ratio of activities of PEP-carboxylase to RuBP carboxylase/oxgenase, or any improvement in daytime leaf water potential for the species tested in this experiment. In spite of the simulation of leaf photosynthesis, a significant increase in growth at elevated [CO₂] was only observed for one species, F. trinervia. Results from this study indicate that leaf photosynthetic rates of certain C₄ species can respond directly to increased [CO₂] under optimal growth conditions, but that the stimulation of whole plant growth at elevated carbon dioxide cannot be predicted solely on the response of individual leaves.     

Contrasting stream water NO3− and Ca2+ in two nearly adjacent catchments: the role of soil Ca and forest vegetation

Two nearly adjacent subcatchments, located in the Adirondack Mountains of New York State, US, with similar atmospheric inputs of N (0.6 kmol ha^?1 yr^?1), but markedly different stream water solute concentrations, provided a unique opportunity to evaluate the mechanisms causing this variation. Subcatchment 14 (S14) had much greater stream water Ca²⁺ and NO₃^? concentrations (851 and 73 μmol_c L^?1, respectively) than Subcatchment 15 (S15) (427 and 26 μmol_c L^?1, respectively). To elucidate factors affecting the variability in stream water concentrations, soil and forest floor samples from each subcatchment were analyzed for total elemental cations and extractable N species. Mineral soil samples were also analyzed for exchangeable cations. Tree species composition was characterized in each subcatchment and potential differences in land use history and hydrology were also assessed. Compared with S15, soils in S14 had significantly higher total elemental Ca²⁺ in the forest floor (380 vs. 84 μmol g^?1), Bs horizon (e.g. 1361 vs. 576 μmol g^?1) and C horizon (1340 vs. 717 μmol g^?1). Exchangeable Ca²⁺ was also significantly higher in the mineral soil (64 μmol g^?1 in S14 vs. 8 μmol g^?1 in S15). Extractable NO₃^? was higher in S14 compared with S15 in both the forest floor (0.1 vs. 0.01 μmol g^?1) and Bs horizon (0.2 vs. 0.07 μmol g^?1) while extractable NH₄⁺ was higher in S14 vs. S15 in the forest floor (7 vs. 5 μmol g^?1). The total basal area of ‘base‐rich indicator’ tree species (e.g. sugar maple, American basswood, eastern hophornbeam) was significantly greater in S14 compared with S15, which had species characteristic of sites with lower base concentrations (e.g. American beech and eastern white pine). The disparity in stream water Ca²⁺ and NO₃^?, concentrations and fluxes between S14 and S15 were explained by differences in tree species composition and soil properties rather than differences in land use or hydrology. The marked difference in soil Ca²⁺ concentrations in S14 vs. S15 corresponded to the higher stream water Ca²⁺ and the larger contribution of base‐rich tree species to the overstory biomass in S14. Soil under such species is associated with higher net mineralization and nitrification and likely contributed to the higher NO₃^? concentrations in the drainage waters of S14 vs. S15. Studies investigating differences in spatial and temporal patterns of the effects of chronic N deposition on surface water chemistry need to account for changes in tree species composition and how vegetation composition is influenced by soil properties, as well as climatic and biotic changes.