Effects of manganese toxicity on photosynthesis of white birch (Betula platyphylla var. japonica) seedlings

The effects of manganese (Mn) toxicity on photosynthesis in white birch ( Betula platyphylla var. japonica ) leaves were examined by the measurement of gas exchange and chlorophyll fluorescence in hydroponically cultured plants. The net photosynthetic rate at saturating light and ambient CO₂ (C_a) of 35 Pa decreased with increasing leaf Mn concentrations. The carboxylation efficiency, derived from the difference in CO₂ assimilation rate at intercellular CO₂ pressures attained at C_a of 13 Pa and O Pa, decreased with greater leaf Mn accumulation. Net photosynthetic rate at saturating light and saturating CO₂ (5%) also declined with leaf Mn accumulation while the maximum quantum yield of O₂ evolution at saturating CO₂ was not affected. The maximum efficiency of PSII photochemistry (F_v/F_m) was little affected by Mn accumulation in white birch leaves over a wide range of leaf Mn concentrations (2–17 mg g₋₁ dry weight). When measured in the steady state of photosynthesis under ambient air at 430 μmol quanta m₋₂ s₋₁, the levels of photochemical quenching (qP) and the excitation capture efficiency of open PSII (F'^v/F'^m) declined with Mn accumulation in leaves. The present results suggest that excess Mn in leaves affects the activities of the CO² reduction cycle rather than the potential efficiency of photochemistry, leading to increases in Q_A reduction state and thermal energy dissipation, and a decrease in quantum yield of PSII in the steady state.     

Photoinhibition of photosynthesis in intact willow leaves in response to moderate changes in light and temperature

When willow leaves were transferred from 270 to 650 μmol m^-2 s^-1 photosynthetic photon flux density (PPFD), partial photoinhibition developed over the next hours. This was manifested as roughly parallel inhibitions of the ratio of variable over maximal chlorophyll fluorescence (F_v/F_M), and of the maximal quantum yield and the capacity of photosynthesis. This occurred even though photosynthesis was operating well below its capacity and only about one fourth of the reaction centres of photosystem (PS) II were in the closed state. When the air temperature was lowered from 25 to 15°C (18°C leaf temperature) photoinhibition was markedly accelerated. This temperature effect is suggested to be mediated largely by a decrease in the rate of energy dissipation through photosynthesis and indicated by a 50% increase in the number of closed PSII reaction centres. The pool size of the carotcnoid zeaxanthin and the extent of inhibition of the F_v/F_M ratio were positively correlated during the treatment. However, the relaxation following imposition of darkness was much faster for zeaxanthin than for the F_v/F_M ratio, ruling out the possibility of a direct causal relationship. The energy distribution between PSII and PSI was unaltered upon photoinhibition. However, the functioning of the PSII reaction centres was altered, as indicated by a rise in the minimal fluorescence, Fa.     

CO2 assimilation, respiration and chlorophyll fluorescence in peach leaves infected by Taphrina deformans

Photosynthesis, respiration and chlorophyll fluorescence parameters were determined in peach ( Prunus persica L. cv. Dixired) leaves naturally infected by Taphrina deformans (Berk.) Tul. and in healthy leaves (controls), in two successive springs. A drastic decrease in net photosynthesis and an evident increase in respiration in curled leaves were noted. The instantaneous PSII fluorescence yield, with no (F₀) and with (F₀) quenching component, and steady state fluorescence yield (under actinic light, F_s) were essentially unchanged. Maximal fluorescence in dark-adapted (F_m) and illuminated (F'_m) leaves and the corresponding variable fluorescence (F_v and F_v) clearly decreased. The indicators of PSII quantum yield (F_v/F_m) in dark-adapted leaves, and the potential PSII excitation capture efficiency (F'_v/F'_m) and the quantum yield of PSII (q_p [F'_v/F'_m]) in the light were also significantly lower in curled leaves. Decreasing tendencies were also noted for the PSII photochemical yield (photochemical quenching, q_p) and in the energy status of the chloroplast (non-photochemical quenching, q_N, and Stern-Vollmer value, NPQ) although the differences were not always significant. In curled leaves the main alteration documented is the imbalance between the drastic inhibition of CO₂ fixation and the moderate decrease in photochemical reactions (i.e. F_v/F_m and ΔF/F'_m), indicating changes in the energy flux.     

Beech trees exposed to high CO2 and to simulated summer ozone levels: Effects on photosynthesis, chloroplast components and leaf enzyme activity

Young trees of European beech ( Fagus sylvatica L.) were exposed in a phytotron to different levels of ozone and CO₂ under the climatic simulation of one vegetation period. High ozone levels were simulated similar to high ozone concentration in the field (up to 110 ppb), while CO₂ was added as 300 ppm to the present level of ca 380 ppm. Our study describes different aspects of photosynthesis from the leaf level to the reactions of selected thylakoid components at different harvest times during growth of the beech trees under the different fumigation regimes. Ozone effects appeared in the first weeks of the treatment as a stimulation of chlorophyll fluorescence (F_v/F_m), in oxygen production and in ribulose-1,5-bisphosphate carboxylase/oxygenase activity, while the summer and early autumn harvests showed strong reductions in these parameters. Only phosphoenolpyruvate carboxylase (PEPcase) activity remained higher under high ozone. The effects of high CO₂ appeared in general as a small stimulation in enzyme activity like PEPcase in spring. However, with increasing time of fumigation, reductions of all parameters were observed. Especially chlorophylls showed strong reductions under high CO₂. The combined treatment with high ozone plus high CO₂ resulted mostly in an amelioration of the negative ozone effects, although control levels were not reached.     

Chlorophyll fluorescence measured using the Fraunhofer line-depth principle and relationship to photosynthetic rate in the field

Abstract. A field study was conducted to determine the relationship of solar-excited chlorophyll a fluorescence to net CO₂ assimilation rate in attached leaves. The Fraunhofer line-depth principle was used to measure fluorescence at 656.3 nm wavelength while leaves remained exposed to full sunlight and normal atmospheric pressures of CO₂ and O₂. Fluorescence induction kinetics were observed when leaves were exposed to sunlight after 10 min in darkness. Subsequently, fluorescence varied inversely with assimilation rate. In the C₄ Zea mays , fluorescence decreased from 2.5 to 0.8 mW m^-2 nm^-1 as CO₂ assimilation rate increased from 1 to 8 μmol m^-2 s^-1 (r²= 0.520). In the C₃ Liquidambar styraciflua and Pinus taeda , fluorescence decreased from 6 to 2 mW m^-2 nm^-1 as assimilation rate increased from 2 to 5 or 0 to 2 μmol m^-2 s^-1 (r²= 0.44 and 0.45. respectively). The Fraunhofer line-depth principle enables the simultaneous measurement of solar-excited fluorescence and CO₂ assimilation rate in individual leaves, but also at larger scales. Thus, it may contribute significantly to field studies of the relationship of fluorescence to photosynthesis.     

Effect of light and gibberellic acid on photosynthesis during leaf senescence of alstroemeria cut flowers.

The functioning of the photosynthetic apparatus during leaf senescence was investigated in alstroemeria cut flowers by a combination of gas-exchange measurements and analysis of in vivo chlorophyll fluorescence. Chlorophyll loss in leaves of alstroemeria cut flowers is delayed by light and by a treatment of the cut flowers with gibberellic acid (GA₃). The maximal photosynthesis of the leaves was approximately 6 μmol CO₂ m⁻² s⁻¹ at I 350 μmol m⁻² s⁻¹ (PAR) which is relatively low for intact C₃ leaves. Qualitatively the gas-exchange rates followed the decline in chlorophyll content for the various treatments, i.e. light and GA₃-treatment delayed the decline in photosynthetic rates. However, when chlorophyll loss could not yet be observed in the leaves, photosynthetic rates were already strongly decreased. In vivo fluorescence measurements revealed that the decrease in CO₂ uptake is (partly) due to a decreased electron flow through photosystem II. Furthermore, analysis of the fluorescence data showed a high nonphotochemical quenching under all experimental conditions, indicating that the consumption of reducing power in the Calvin cycle is very low. The chlorophyll, remaining after 9 days incubation of leaves with GA₃ in the dark should be considered as a 'cosmetic' pigment without any function in the supply of assimilates to the flowers.     

Drought-induced shifts in daily CO2 uptake patterns for leafy cacti

For cacti with persistent, relatively large leaves, most shoot CO₂ uptake under well-watered conditions occurs by the leaves using the C₃ pathway. For three species in the primitive subfamily Pereskioideae, droughts of 7 or 14 days decreased leaf daytime net CO₂ uptake by an average of 49 and 88%, respectively; these species always had a net CO₂ release at night by the leaves and both at night and during the day by the stems. For three leafy species in subfamily Opuntioideae, 7 and 14 days of drought reduced leaf daytime net CO₂ uptake by 90 and 100%, respectively. Although drought reduced the total CO₂ uptake over 24 h, the average percentage occurring at night by the leaves of these species increased from 5% under wet conditions to 71% after 7 days of drought to 99% after 14 days of drought. For two of the three species of Opuntioideae, 7 days of drought caused the small net CO₂ uptake by the sterns to shift from the daytime to the nighttime, while for the third species drought caused a reduction of its stem nocturnal net CO₂ uptake. Thus, shifts from predominantly daytime to predominantly nighttime net CO₂ uptake can be induced by drought for the leaves and the stems of leafy cacti in subfamily Opuntioideae, indicating a high degree of biochemical versatility.     

Role of light in changes in free amino acid and polyamine contents at chilling temperature in maize (Zea mays)

The effect of irradiance on changes in photosynthesis, free amino acids and polyamines was investigated. Two-week-old maize ( Zea mays L.) plants were chilled at 5°C in the light (250 μmol m⁻² s⁻¹ PAR) or dark. The chlorophyll fluorescence ratio, F_v/F_m, decreased in the light by ca 50% but did not change in the dark. Similarly to the F_v/F_m, there was no change in the transpiration rate or the stomatal conductance in the dark, while these parameters decreased by ca 55% in the light. The net photosynthesis rate declined in both cases, but to a far greater extent in the light (73%) than in the dark (40%). The intercellular CO₂ partial pressure increased by ca 50% in all cases. The free amino acid contents increased compared to the control during the cold treatment. In most cases this increase was more pronounced in the light than in the dark. There was a continuous increase in the putrescine level, which was more pronounced in the light than in the dark. The spermidine content increased one and a half times after one day in the light but decreased by 70% in the dark compared to the control values. From the second day a 50% decline in the spermidine content was observed in the light and an 80% decline in the dark. These results suggest that light has an influence not only on the photosynthetic processes during chilling stress but also on other stress markers such as polyamines and free amino acids.     

Physiological effects of long-term exposure to low and moderate concentrations of atmospheric NH3 on poplar leaves

Abstract. Poplar shoots ( Populus euramericana L.) obtained from cuttings were exposed for 6 or 8 weeks to NH₃ concentrations of 50 and 100 μgm⁻³ or filtered air in fumigation chambers. After this exposure the rates of NH₃ uptake, transpiration, CO₂ assimilation and respiration of leaves were measured using a leaf chamber. During the long-term exposure also modulated chlorophyll fluorescence measurements were carried out to obtain information about the photosynthetic performance of individual leaves. Both fluorescence and leaf chamber measurements showed a higher photosynthetic activity of leaves exposed to 100 μg NH₃ m⁻³. These leaves showed also a larger leaf conductance and a larger uptake rate of NH₃ than leaves exposed to 50 μg m⁻³ NH₃ or filtered air. The long-term NH₃ exposure did not induce an internal resistance against NH₃ transport in the leaf, nor did it affect the leaf cuticle. So, not only at a short time exposure, but also at a long-term exposure NH₃ uptake into leaves can be calculated from data on the boundary layer and stomatal resistance for H₂O and ambient NH₃-concentration. Furthermore, the NH₃ exposure had no effect on the relation between CO₂-assimilation and stomatal conductance, indicating that NH₃ in concentrations up to 100 μg m⁻³ has no direct effect on stomatal behaviour; for example, by affecting the guard or contiguous cells of the stomata.     

Photoinhibition at chilling temperatures and effects of freezing stress on cold acclimated spinach leaves in the field. A fluorescence study

The role of high light stress in a natural environment was studied on spinach plants ( Spinacia oleracea L. cv. Wolter) grown in the field during the winter season. Fluorescence induction (at 293 K and 77 K) of leaves was used to characterize the stress effects. Night frost with minimum temperatures between – 1.5°C and –7.5°C (i.e. above the'frost killing point'at ca. –11.5°C) led to impaired photosynthesis. This was seen as increased initial fluorescence (F_o), decreased ratio of variable to maximum fluorescence (F_V/F_M) and lowered rates of O₂ evolution. The freezing injury was reversible within several frostless days. Exposure to high light (about 900 mol m_–2 s_–1) at chilling temperatures in the field caused photoinhibition, manifested as decreased variable fluorescence (F_V) and F_V/F_M ratio without changes in F_O. The photoinhibitory fluorescence quenching was not stronger after frost than after frostless nights; synergism between light stress and preceding freezing stress was not observed. Fluorescence induction signals at 77 K showed that F_V of photosystems I and II decreased to the same extent, indicating increased thermal deactivation of excited chlorophyll. Photoinhibition was fully reversible at +4°C within 1 h in low light, but only partially in moderate light. Preceding night frosts did not affect the recovery. The photoinhibition observed here is regarded as a protective system of thermal dissipation of excess light energy.     

Photosynthesis of two poplar clones under long‐term exposure to ozone

The photosynthetic response was studied in two clones ( Populus deltoides × maximowiczii Eridano and Populus × euramericana I‐214), known for their differential response to ozone (O₃) in terms of visible symptoms, when exposed to O₃ (60 nl l⁻¹ 5 h day⁻¹, 7 and 15 days). The photosynthetic ability was tested using gas exchange and chlorophyll fluorescence analysis. O₃ caused a decrease in the CO₂ assimilation rate at light saturation level in mature leaves of both clones. Alterations of Chl fluorescence parameters, in particular the F_v/F_m ratio and non‐photochemical quenching were also observed. The effects were similar for both clones and it could not be concluded that differential effects on electron transport capacity were responsible for the observed reduction in photosynthesis. The reduction of photosynthetic rate in Eridano was due mainly to a reduced mesophyll activity, as evidenced by the increase in intercellular CO₂ concentration and the minimal changes in stomatal conductance. In contrast, in I‐214, stomatal effects were primarily responsible, although effects on the mesophyll cannot be excluded. Data obtained indicate that the effects observed at the mesophyll level may be attributed to indirect effects caused by membrane disorders.     

Inhibition of photosynthesis and export in geranium grown at two CO2 levels and infected with Xanthomonas campestris pv. Pelargonii

The effects of CO₂ enrichment on growth of Xanthomonas campestris pv. pelargonii and the impact of infection on the photosynthesis and export of attached, intact, 'source' leaves of geranium ( Pelargonium x domesticum, 'Scarlet Orbit Improved' ) are reported. Two experiments were performed, one with plants without flower buds, and another with plants which were flowering. Measurements were made on healthy and diseased leaves at the CO₂ levels (35 Pa or 90 Pa) at which the plants were grown. There were no losses of chlorophyll, or any signs of visible chlorosis or necrosis due to infection. Lower numbers of bacteria were found in leaves at high CO₂, suggesting growth at elevated CO₂ created a less favourable condition in the leaf for bacterial growth. Although high CO₂ lowered the bacterial number in infected leaves, reductions in photosynthesis and export were greater than at ambient CO₂. The capacity of infected source leaves to export photoassimilates at rates observed in the controls was reduced in both light and darkness. In summary, the severity of infection on source leaf function by the bacteria was increased, rather than reduced by CO₂ enrichment, underscoring the need for further assessment of plant diseases and bacterial virulence in plants growing under varying CO₂ levels.     

Interactive effects of water table and precipitation on net CO2 assimilation of three co-occurring Sphagnum mosses differing in distribution above the water table

Sphagnum cuspidatum , S. magellanicum and S. rubellum are three co-occurring peat mosses, which naturally have a different distribution along the microtopographical gradient of the surface of peatlands. We set out an experiment to assess the interactive effects of water table (low: −10 cm and high: −1 cm) and precipitation (present or absent) on the CO₂ assimilation and evaporation of these species over a 23-day period. Additionally, we measured which sections of the moss layer were responsible for light absorption and bulk carbon uptake. Thereafter, we investigated how water content affected carbon uptake by the mosses. Our results show that at high water table, CO₂ assimilation of all species gradually increased over time, irrespective of the precipitation. At low water table, net CO₂ assimilation of all species declined over time, with the earliest onset and highest rate of decline for S. cuspidatum . Precipitation compensated for reduced water tables and positively affected the carbon uptake of all species. Almost all light absorption occurred in the first centimeter of the Sphagnum vegetation and so did net CO₂ assimilation. CO₂ assimilation rate showed species-specific relationships with capitulum water content, with narrow but contrasting optima for S. cuspidatum and S. rubellum . Assimilation by S. magellanicum was constant at a relatively low rate over a broad range of capitulum water contents. Our study indicates that prolonged drought may alter the competitive balance between species, favoring hummock species over hollow species. Moreover, this study shows that precipitation is at least equally important as water table drawdown and should be taken into account in predictions about the fate of peatlands with respect to climate change.     

Carbon dioxide exchange in lichens: Partition of total CO2 resistances at different thallus water contents into transport and carboxylation components

The total resistances to CO₂ uptake by Sticta latifrons Rich, and Pseudocyphellaria amphisticta Kremp. were separated into transport and carboxylation components by calculation after transformation of net photosynthesis rate against CO₂ concentration curves into a linear form. The use of this technique circumvented the problem of measuring the internal CO₂ concentration of the lichen thalli. Both species exhibited an increase in transport resistance at high thallus water contents and an increase in both transport and carboxylation resistances at low water contents. At low and intermediate water contents internal transport resistances were larger than carboxylation resistances when measured at limiting CO₂ concentrations. However, at ambient CO₂ concentrations carboxylation processes were the dominant factors limiting photosynthesis at all, except the high, water contents.     

An integrated portable apparatus for the simultaneous field measurement of photosynthetic CO2 and water vapour exchange, light absorption and chlorophyll fluorescence emission of attached leaves

Abstract. A portable apparatus has been constructed to measure simultaneously the quantum yield of CO₂ assimilation, light absorption, chlorophyll fluorescence emission and water vapour exchange of attached intact leaves in the field. The core of the instrument is a light-integrating spherical leaf chamber which includes ports for a light source, photosynthetically active radiation sensor, fluorescence probes and gas inlet and outlet manifolds. Measurement of the quantum flux inside the empty chamber and with a leaf present allows determination of leaf absorptance. An open gas exchange system is employed using an infra-red analyser to measure leaf CO₂ exchange. Using a DC white light source the quantum yield of CO₂ assimilation based on absorbed light (φ_abs) may be determined rapidly in either ambient air or artificial gas mixtures. Inclusion of capacitance humidity probes into the gas inlet and outlet ports allows simultaneous determination of water vapour exchange and subsequent estimation of stomatal conductance to CO₂ and intercellular CO₂ concentration. Measurement of fluorescence emission by the sample leaf exposed to white light is achieved by a modulated fluorescence detection system. In addition to determination of the minimal, maximal and variable fluorescence levels, a further analysis allows the photochemical and non-photochemical components of fluorescence quenching, to be estimated. The theory and design of this apparatus is described in detail. The use of the apparatus in the field is demonstrated through a study of the photosynthetic performance of a maize and bean crop during the growing season and by analysis of the photosynthetic performance of crops subjected to nitrogen-stress and a herbicide treatment.     

Relationship between photosystem II activity and CO2 fixation in leaves

There is now potential to estimate photosystem II (PSII) activity in vivo from chlorophyll fluorescence measurements and thus gauge PSII activity per CO₂ fixed. A measure of the quantum yield of photosystem II, Φ_II (electron/photon absorbed by PSII), can be obtained in leaves under steady-state conditions in the light using a modulated fluorescence system. The rate of electron transport from PSII equals Φ_II times incident light intensity times the fraction of incident light absorbed by PSII. In C₄ plants, there is a linear relationship between PSII activity and CO₂ fixation, since there are no other major sinks for electrons; thus measurements of quantum yield of PSII may be used to estimate rates of photosynthesis in C₄ species. In C₃ plants, both CO₂ fixation and photorespiration are major sinks for electrons from PSII (a minimum of 4 electrons are required per CO₂, or per O₂ reacting with RuBP). The rates of PSII activity associated with photosynthesis in C₃ plants, based on estimates of the rates of carboxylation (v_o) and oxygenation (v_o) at various levels of CO₂ and O₂, largely account for the PSII activity determined from fluorescence measurements. Thus, in C₃ plants, the partitioning of electron flow between photosynthesis and photorespiration can be evaluated from analysis of fluorescence and CO₂ fixation.     

Transitions in photosynthetic parameters of midvein and interveinal regions of leaves and their importance during leaf growth and development

Abstract: The areal development of photosynthetic efficiency and growth patterns in expanding leaves of two different dicotyledonous species - Coccoloba uvifera and Sanchezia nobilis - was investigated by imaging both processes repeatedly over 32 days. Measurements were performed using combined imaging systems for chlorophyll fluorescence and growth, with the same spatial resolution. Significant differences in potential quantum yield of photosynthesis (F_v/F_m), a parameter indicating the functional status of photosystem II, were found between midvein and interveinal tissue. Although base-tip gradients and spatial patchiness were observed in the distribution of relative growth rate, neither midvein nor interveinal tissue showed such patterns in F_v/F_m. In young leaves, F_v/F_m of the midvein was higher than F_v/F_m of interveinal tissue. This difference declined gradually with time, and upon cessation of growth, F_v/F_m of interveinal regions exceeded those of midvein tissue. Images of chlorophyll fluorescence quenching showed that ΔF/F_m' in the different tissues correlated with F_v/F_m, indicating that, in these uniformly illuminated leaves, transitions in photosynthetic electron transport activity follow those of predawn quantum efficiency. We explore the implications of these observations during leaf development, discuss effects of sucrose delivery from veins to interveinal areas on relative rates of photosynthetic development in these tissues, and propose that the initially higher photosynthetic activity in the midvein compared to the intervein tissues may supply carbohydrates and energy for leaf growth processes.     

Reversible, Water Stress-Induced Non-Uniform Chlorophyll Fluorescence Quenching in Wilting Leaves of Potentilla reptans may not be due to Patchy Stomatal Responses

Abstract: After exposure to full sunlight under natural conditions, attached leaves of the common meadow weed Potentilla reptans show non-uniform ("patchy") chlorophyll fluorescence quenching in the early stages of fluorescence transients. These areas of bright fluorescence can be readily reproduced in detached leaves that are allowed to wilt on the laboratory bench in weak light. The extent and duration of the patchiness increases with increasing water stress (higher relative saturation deficits). Images captured during saturating flashes show that the patches also display slow development of non-photochemical quenching, consistent with the possibility that photosynthetic metabolism is impaired in these areas. Wilted Potentilla leaves readily regain full turgor when petioles are placed in water, and uniform chlorophyll fluorescence is recovered with in 30mm. Epidermal impressions reveal closed stomata over areas of both low and high fluorescence in wilted leaves. Because highly fluorescent patches also persist when wilted tissues are exposed to high CO₂ (i.e., patchiness is unlikely to be due to local differences in CO₂ supply) the data suggest direct effects of water stress on metabolism in wilted leaves. Leaf transverse sections show that although major veins may isolate areas of the lamina, minor veins do not. Relationships to leaf anatomy are discussed.     

Alleviation of photoinhibition by calcium supplement in salt-treated Rumex leaves

By analysis of gas exchange and chlorophyll fluorescence, the effects of NaCl treatment and supplemental CaCl₂ on photosynthesis, photosystem II (PSII) photochemistry and photoinhibition were investigated in Rumex leaves. Photosynthesis in Rumex leaves was strongly inhibited by 200 m M NaCl treatment. Such inhibition of photosynthesis was ameliorated by CaCl₂ supplement. Neither NaCl treatment nor CaCl₂ supplement had any significant effects on the PSII primary photochemical reaction in dark-adapted leaves. In light-adapted leaves, however, 200 m M NaCl treatment significantly decreased photochemical quenching (q_p), efficiency of excitation energy capture by open PSII reaction centers (F_V'/F_M') and quantum yield of PSII electron transport (Φ_PSII). These decreases in q_p, F_V'/F_M' and Φ_PSII were mitigated by CaCl₂ supplement with the maximum of its effect appearing at a concentration of 8 m M CaCl₂. A similar mitigating effect was shown in 200 m M NaCl-treated Rumex leaves when susceptibility of PSII to photoinhibition was determined under high irradiance. It is suggested that the mitigation of photoinhibition in NaCl-treated leaves is because of the amelioration of inhibition of photosynthesis.     

Effect of carbon dioxide enrichment on chlorophyll content, starch content and starch grain structure in Trifolium subterraneum leaves

Trifolium subterraneum (cv. Dinninup) responds to enriched atmospheric CO₂ in a manner similar to that described by Madsen (1968 and 1976) for tomato. In immature leaves, the total chlorophyll content per unit dry weight and the chlorophyll a:b ratio are significantly lower in plants grown at 0.10 vol% CO₂. Although fully expanded mature leaves partially overcome the deficit in chlorophyll content, the chlorophyll a:b ratio remains substantially lower in these high CO₂ grown plants. The large amount of starch accumulated as irregularly shaped grains appears to disrupt normal chloroplast structure in clover plants grown in enriched atmospheric CO₂. These results indicate the chlorotic appearance of leaves from high CO₂ grown clover plants is due to a decrease in chlorophyll content per dry weight possibly resulting from large starch grains and starch accumulation altering normal chloroplast structure and function.