Acclimation of photosynthesis to increasing atmospheric CO2: The gas exchange perspective

The nature of photosynthetic acclimation to elevated CO₂ is evaluated from the results of over 40 studies focusing on the effect of long-term CO₂ enrichment on the short-term response of photosynthesis to intercellular CO₂ (the A/C_i response). The effect of CO₂ enrichment on the A/C_i response was dependent on growth conditions, with plants grown in small pots (< 5 L) or low nutrients usually exhibiting a reduction of A at a given C_i, while plants grown without nutrient deficiency in large pots or in the field tended to exhibit either little reduction or an enhancement of A at a given C_i following a doubling or tripling of atmospheric CO₂ during growth. Using theoretical interpretations of A/C_i curves to assess acclimation, it was found that when pot size or nutrient deficiency was not a factor, changes in the shape of A/C_i curves which are indicative of a reallocation of resources within the photosynthetic apparatus typically were not observed. Long-term CO₂ enrichment usually had little effect or increased the value of A at all C_i. However, a minority of species grown at elevated CO₂ exhibited gas exchange responses indicative of a reduced amount of Rubisco and an enhanced capacity to metabolize photosynthetic products. This type of response was considered beneficial because it enhanced both photosynthetic capacity at high CO₂ and reduced resource investment in excessive Rubisco capacity. The ratio of intercellular to ambient CO₂ (the C_i/C_a ratio) was used to evaluate stomatal acclimation. Except under water and humidity stress, C_i/C_a exhibited no consistent change in a variety of C₃ species, indicating no stomatal acclimation. Under drought or humidity stress, C_i/C_a declined in high-CO₂ grown plants, indicating stomata will become more conservative during stress episodes in future high CO₂ environments.Abbreviations A net CO₂ assimilation rate - C_i (C_a) intercellular (ambient) partial pressure of CO₂ - operational C_i intercellular partial pressure of CO₂ at a given ambient partial pressure of CO₂ - g_s stomatal conductance - normal CO₂ current atmospheric mole fraction of CO₂ (330 to 355 mol mol^–1) - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase     

Cardiorespiratory responses of the woodchuck and porcupine to CO2 and hypoxia

Summary The burrow-dwelling woodchuck (Marmota monax) (mean body wt.=4.45±1 kg) was compared to a similar-sized (5.87±1.5 kg) but arboreal rodent, the porcupine (Erithrizon dorsatum), in terms of its ventilatory and heart rate responses to hypoxia and hypercapnia, and its blood characteristics.V _T,f,T _I andT _E were measured by whole-body plethysmography in four awake individuals of each species. The woodchuck has a longerT _E/T _TOT (0.76±0.03) than the porcupine (0.61±0.03). The woodchuck had a higher threshold and significantly smaller slope to its CO₂ ventilatory response compared to the porcupine, but showed no difference in its hypoxic ventilatory response. The woodchuck P₅₀ of 27.8 was hardly different from the porcupine value of 30.7, but the Bohr factor, –0.72, was greater than the porcupine's, –0.413. The woodchuck breathing air has PaCO₂=48 (±2) torr, PaO₂=72 (±6), pHa=7.357 (±0.01); the porcupine blood gases are PaCO₂=34.6 (±2.8), PaO₂=94.9 (±5), pHa=7.419 (±0.03), suggesting a difference in PaCO₂/pH set points. The woodchuck exhibited no reduction in heart rate with hypoxia, nor did it have the low normoxic heart rate observed in other burrowing mammals.     

Soil drying and its effect on leaf conductance and CO2 assimilation of Vigna unguiculata (L.) Walp

Summary Well watered plants of Vigna unguiculata (L.) Walp cv. California Blackeye No. 5 had maximum photosynthetic rates of 16 mol m^-2 s^-1 (at ambient CO₂ concentration and environmental parameters optimal for high CO₂ uptake). Leaf conductance declined with increasing water vapour concentration difference between leaf and air (w), but it increased with increasing leaf temperature at a constant small w. When light was varied, CO₂ assimilation and leaf conductance were correlated linearly. We tested the hypothesis that g was controlled by photosynthesis via intercellular CO₂ concentration (c _i). No unique relationship between (1) c _i, (2) the difference between ambient CO₂ concentration (c _a) and c _i, namely c _a-c _i, or (3) the c _i/c _a ratio and g was found. g and A appeared to respond to environmental factors fairly independently of each other. The effects of different rates of soil drying on leaf gas exchange were studied. At unchanged air humidity, different rates of soil drying were produced by using (a) different soils, (b) different irrigation schemes and (c) different soil volumes per plant. Although the soil dried to wilting point the relative leaf water content was little affected. Different soil drying rates always resulted in the same response of photosynthetic capacity (A _max) and corresponding leaf conductance (g(_Amax)) when plotted against percent relative plant-extractable soil water content (W _e %) but the relationship with relative soil water content (W _e ) was less clear. Above a range of W _e of 15%–25%, A _max and g(_Amax) were both high and responded little to decreasing W _e . As soon as W _e fell below this range, A _max and g(_Amax) declined. The data suggest root-to-leaf communication not mediated via relative leaf water content. However, g(_Amax) was initially more affected than A _max.List of abbreviations A CO₂ assimilation - A _max photosynthetic capacity at favourable ambient conditions - c _a CO₂ concentration of the air in the leaf chamber - c _i intercellular - CO₂ concentration - E transpiration - g leaf conductance - g(_Amax) leaf conductance corresponding to photosynthetic capacity - I photon flux rate - T _l leaf temperature - W _e relative plant-extractable soil water content - W _e absolute plant-extractable soil water content - W _l relative leaf water content - W _s relative soil water content - w difference in water vapour mole fraction between leaf and air - leaf water potential     

Photosynthetic response of Podophyllum hexandrum Royle from different altitudes in Himalayan ranges

Plants of Podophyllum hexandrum, collected from lower, mid, and upper distribution limits in alpine Himalaya were studied under greenhouse conditions to evaluate the photosynthetic response. Net photosynthetic rates (P _N), stomatal conductance (g _s), and efficiency of carbon uptake increased with altitude. The maximum P _N and g _s were measured in the considered population during the 3–6^th week of development. P _N and g _s decreased on an average by 58 and 48 % from maximum rates reached around 4^th week to the 10^th week of growth, respectively. The photosynthetic response in the three ecotypes appeared to be genetically controlled.     

A new species of Botryocladia from the West Indies

A very small species of Botryocladia (Rhodophyta-Rhodymeniales), notable for the spinose projections on the surface cells of the vesicles, is here described as B. spinulifera sp. nov. from the Bahamas and the American Virgin Islands.     

Physiological responses of carbon-sequestering microalgae to elevated carbon regimes

In order to identify a high carbon-sequestering microalgal strain, the physiological effect of different concentrations of carbon sources on microalgae growth was investigated. Five indigenous strains (I-1, I-2, I-3, I-4 and I-5) and a reference strain (I-0: Coccolithus pelagicus 913/3) were subjected to CO₂ concentrations of 0.03–15% and NaHCO₃ of 0.05–2 g CO₂ l^–1. The logistic model was applied for data fitting, as well as for estimation of the maximum growth rate (μ_max) and the biomass carrying capacity (B_max). Amongst the five indigenous strains, I-3 was similar to the reference strain with regards to biomass production values. The B_max of I-3 significantly increased from 214 to 828 mg l^–1 when CO₂ concentration was increased from 0.03 to 15% (r = 0.955, P = 0.012). Additionally, the B_max of I-3 increased with increasing NaHCO₃ (r = 0.885, P = 0.046) and was recorded at 153 mg l^–1 (at 0.05 g CO₂ l^–1) and 774 mg l^–1 at (2 g CO₂ l^–1). Relative electron transport rate (rETR) and maximum quantum yield (F_v/F_m) were also applied to assess the impact of elevated carbon sources on the microalgal cells at the physiological level. Isolate I-3 displayed the highest rETR confirming its tolerance to higher quantities of carbon. Additionally, the decline in F_v/F_m with increasing carbon was similar for strains I-3 and the reference strain. Based on partial 28s ribosomal RNA gene sequencing, strain I-3 was homologous to the ribosomal genes of Chlorella sp.     

Leaf cavity CO2 concentrations and CO2 exchange in onion,Allium cepa L.

Onion (Allium cepa L.) plants were examined to determine the photosynthetic role of CO₂ that accumulates within their leaf cavities. Leaf cavity CO₂ concentrations ranged from 2250 L L^–1 near the leaf base to below atmospheric (<350 L L^–1) near the leaf tip at midday. There was a daily fluctuation in the leaf cavity CO₂ concentrations with minimum values near midday and maximum values at night. Conductance to CO₂ from the leaf cavity ranged from 24 to 202 mol m^–2 s^–1 and was even lower for membranes of bulb scales. The capacity for onion leaves to recycle leaf cavity CO₂ was poor, only 0.2 to 2.2% of leaf photosynthesis based either on measured CO₂ concentrations and conductance values or as measured directly by ¹⁴CO₂ labeling experiments. The photosynthetic responses to CO₂ and O₂ were measured to determine whether onion leaves exhibited a typical C₃-type response. A linear increase in CO₂ uptake was observed in intact leaves up to 315 L L^–1 of external CO₂ and, at this external CO₂ concentration, uptake was inhibited 35.4±0.9% by 210 mL L^–1 O₂ compared to 20 mL L^–1 O₂. Scanning electron micrographs of the leaf cavity wall revealed degenerated tissue covered by a membrane. Onion leaf cavity membranes apparently are highly impermeable to CO₂ and greatly restrict the refixation of leaf cavity CO₂ by photosynthetic tissue.Abbreviations C_a external CO₂ concentration - C_i intercellular CO₂ concentration - CO₂ compensation concentration - PPFR photosynthetic photon fluence rate     

Photosynthesis and yield responses of ozone-polluted winter wheat to drought

Winter wheat (Triticum aestivum L. cv. Jingdong 8) was exposed to short-term high ozone treatment after anthesis and then was either well irrigated with soil water content (SWC) of 80–85 % (O₃+W) or drought treated (SWC 35–40 %, O₃+D). Short-term ozone exposure significantly decreased irradiance-saturated net photosynthetic rate (P _N) of winter wheat. Under good SWC, P _N of the O₃-treated plant was similar to that of control on 2 d after O₃-exposure (6 DAA), but decreased significantly after 13 DAA, indicating that O₃ exposure accelerated leaf senescence. Meanwhile, green flag leaf area was reduced faster than that of control. As a result, grain yield of O₃+W was significantly decreased. P _N of O₃+D was further notably decreased and green flag leaf area was reduced more than that in O₃+W. Consequently, substantial yield loss of O₃+D was observed compared to that of O₃+W. Although P _N was significantly positively correlated with stomatal conductance, it also had notable positive correlation with the maximum photochemical efficiency in the dark adapted leaves (F_v/F_m), electron transport rate (ETR), photochemical quenching (q_P), as well as content of chlorophyll, suggesting that the depression of P _N was mainly caused by non-stomatal limitation. Hence optimal soil water condition should be considered in order to reduce the yield loss caused by O₃ pollution.     

Effect of doubled CO<Subscript>2</Subscript> on morphology: Inhibition of stomata development in growing birch (<Emphasis Type="Italic">Betula platyphylla</Emphasis> Suk.) leaves

Two-year old birch (Betula platyphylla Suk.) seedlings were grown in climatic chambers for 7 weeks under various conditions: (1) ambient CO₂ concentration (350 ppm) and an ordinary nitrogen content in soil (2 mM NH₄NO₃); (2) ambient CO₂ concentration and a high nitrogen rate (16 mM NH₄NO₃); (3) doubled CO₂ concentration (700 ppm) and ordinary nitrogen content, and (4) doubled CO₂ concentration and a high nitrogen rate. Doubled CO₂ concentration in combination with the high nitrogen rate activated mostly seedling growth, e.g., stem thickening and leaf initiation. In this treatment, the maximum rate of apparent photosynthesis (A _max) was twice as high as in control seedlings. At doubled CO₂ concentration and ordinary nitrogen content, we observed the phenomenon of stomata absence from the upper leaf surface and doubling their number on the lower surface, whereas, at doubled CO₂ concentration and a high nitrogen rate, stomata partition was essentially similar as in control leaves. The conclusion is that, when the balance between CO₂ concentration and nitrogen rate is shifted, doubled CO₂ concentration exerts a morphotropic effect on differentiation of young epidermal tissue.Translated from Fiziologiya Rastenii, Vol. 52, No. 2, 2005, pp. 198–202.Original Russian Text Copyright © 2005 by Mao, Y.-J. Wang, X.-W. Wang, Voronin.This revised version was published online in April 2005 with a corrected cover date.     

The Photosynthetic Response to Elevated CO2 in High Altitude Potato Species (Solanum curtilobum)

Plants of Solanum curtilobum (from high altitude) and Solanum tuberosum (from low altitude) were grown in open-top chambers in a greenhouse at either ambient (AC, 360 µmol mol^–1) or ca. twice ambient (EC, 720 µmol mol^–1) CO₂ concentrations for 30 d. CO₂ treatments started at the reproductive stage of the plants. There were similar patterns in the physiological response to CO₂ enrichment in the two species. Stomatal conductance was reduced by 59 % in S. tuberosum and by 55 % in S. curtilobum, but such a reduction did not limit the net photosynthetic rate (P _N), which was increased by approximately 56 % in S. curtilobum and 53 % in S. tuberosum. The transpiration rate was reduced by 16 % in both potato species while instantaneous transpiration efficiency increased by 80 % in S. tuberosum and 90 % in S. curtilobum. Plants grown under EC showed 36 and 66 % increment in total dry biomass, whereas yields (dry mass of tubers) were increased by 40 and 85 % in S. tuberosum and S. curtilobum, respectively. EC promoted productivity by increasing P _N. Thus S. tuberosum, cultivated around the world at low altitudes, and S. curtilobum, endemic of the highland Andes, respond positively to EC during the tuberisation stage.     

Effect of Foliar Application of Chitin and Chitosan Oligosaccharides on Photosynthesis of Maize and Soybean

On the first day after foliar application, chitosan pentamer (CH5) and chitin pentamer (CHIT5) decreased net photosynthetic rate (P _N) of soybean and maize, however, on subsequent days there was an increase in P _N in some treatments. CH5 caused an increase in maize P _N on day 3 at 10^–5 and 10^–7 M; the increases were 18 and 10 % over the control plants. This increase was correlated with increases in stomatal conductance (g _s) and transpiration rate (E), while the intercellular CO₂ concentration (C _i) was not different from the control plants. P _N of soybean plants did not differ from the control plants except for treatment CH5 (10^–7 M) which caused an 8 % increase on day 2, along with increased g _s, E, and C _i. On days 5 and 6 the CHIT5 treatment caused a 6–8 % increase in P _N of maize, which was accompanied by increases in g _s, E, and C _i. However, there was no such increase for soybean plants treated with CHIT5. In general, foliar application of high molecular mass chitin (CHH) resulted in decreased P _N, particularly for 0.010 % treated plants, both in maize and soybean. Foliar applications of chitosan and chitin oligomers did not affect (p > 0.05) maize or soybean height, root length, leaf area, shoot or root or total dry mass.     

Response of tussock tundra to elevated carbon dioxide regimes: analysis of ecosystem CO2 flux through nonlinear modeling

Summary The response of tussock tundra to elevated atmospheric concentrations of CO₂ was measured at Toolik Lake, Alaska in the summer of 1983. Computer-controlled greenhouses were used to determine diurnal ecosystem flux of CO₂ under four treatments: 340 ppm, 510 ppm, and 680 ppm CO₂, as well as 680 ppm CO₂ with a four degree centrigrade increase in temperature. For the seven days of data analyzed, net daily CO₂ flux was significantly different between treatments. Net uptake was positively correlated with CO₂ concentration in the chamber and negatively correlated with temperature. A nonlinear model was used to analyze this data set and to determine some of the reasons for different net CO₂ flux. This model allowed an estimation of light utilization efficiency, total conductance of CO₂, and a comparable measure of total respiration. From this analysis we conclude that nutrient limitations in the arctic decrease the capacity of tundra plants to make use of elevated CO₂ concentrations. The plants respond by decreasing conductance in the presence of elevated CO₂, which results in approximately equal gross uptake rates for the three CO₂ treatments. Apparent changes in system respiration result in higher net uptake under elevated CO₂ but this may be due to biases in the data. The treatment with increased temperature exhibited higher conductances and, consequently, higher gross uptake of CO₂ than the other treatments. Higher temperatures, however, also increase respiration with the result being lower net uptake than would be expected in the absence of temperature inscreases.     

Genotypic variation in the photosynthetic competence of Sorghum bicolor seedlings subjected to polyethylene glycol-mediated drought stress

Eleven varieties of Sorghum bicolor, subjected to PEG-mediated drought stress were compared for their photosynthetic performance. The varieties differed in their relative water content over a range of PEG concentrations (0-25%). CO2 assimilation, stomatal conductance and the quantum yield of PSII electron transport decreased with increasing PEG concentrations in all varieties. However the intercellular CO2 concentration showed a nonlinear PEG concentration-dependent change. At lower PEG concentrations there was a decrease in the levels of intercellular CO2 concentration in all varieties that could be attributed to stomatal closure. At higher PEG concentrations, some varieties showed an increase in the intercellular CO2 concentration, indicating an inhibition of photosynthetic activity due to non-stomatal effects, while others did not. It was seen that the varieties differed in the stress thresholds at which stomatal and metabolic limitations to photosynthesis occur. These differences in the photosynthetic adaptation of Sorghum varieties could be useful in identifying genotypes showing large differences in photosynthetic adaptation, which could be useful in mapping photosynthetic traits for drought stress tolerance.     

Photosynthetic responses of Populus przewalski subjected to drought stress

Cuttings of P. przewalski were exposed to two different watering regimes which were watered to 100 and 25 % of field capacity (WW and WS, respectively). Drought stress not only significantly decreased net photosynthetic rate (P _N), transpiration rate (E), stomatal conductance (g _s), efficiency of photosystem 2 (PS2) (F_v/F_m and yield), and increased intrinsic water use efficiency (WUE_i) under controlled optimal conditions, but also altered the diurnal changes of gas exchange, chlorophyll fluorescence, and WUE_i. On the other hand, WS also affected the P _N-photosynthetically active radiation (PAR) response curve. Under drought stress, P _N peak appeared earlier (at about 10:30 of local time) than under WW condition (at about 12:30). At midday, there was a depression in P _N for WS plants, but not for WW plants, and it could be caused by the whole microclimate, especially high temperature, low relative humidity, and high PAR. There were stomatal and non-stomatal limitations to photosynthesis. Stomatal limitation dominated in the morning, and low P _N at midday was caused by both stomatal and non-stomatal limitations, whereas non-stomatal limitation dominated in the afternoon. In addition, drought stress also increased compensation irradiance and dark respiration rate, and decreased saturation irradiance and maximum net photosynthetic rate. Thus drought stress decreased plant assimilation and increased dissimilation through affected gas exchange, the diurnal pattern of gas exchange, and photosynthesis-PAR response curve, thereby reducing plant growth and productivity.     

Responses of photosynthesis to irradiance and temperature in soybean,Glycine max (L.) Merr

The response of photosynthesis to irradiance and temperature during growth was investigated in two soybean genotypes. Soybean is a species that can modify its structure and metabolism so as to adapt to differing light conditions; its responses to rapid changes in irradiance are characterized by their flexibility. However, the temperature during growth can change the response to irradiance: moreover, there may be a marked interaction with genotype.The response of photosynthesis to irradiance consists of changes in leaf thickness, which bring about variations in the mesophyll resistance to CO₂ transfer. The increase in net photosynthesis per unit of leaf area is due to the increase in the amount of assimilating material beneath unit of area, as corroborated by the stability of the net photosynthesis per unit volume. Moreover, the response of photosynthesis to temperature is due to the mesophyll diffusion constant which decreases with the growth temperature.     

Interaction of ocean and biosphere in their transient responses to increasing atmospheric CO2

Increasing atmospheric CO₂ induces a net uptake of carbon in the ocean by a shift in chemical equilibrium in seawater, and in the terrestrial biosphere by a stimulated photosynthesis and productivity. The fractions absorbed in both biosphere and ocean decline with increasing dynamics of the release rate of CO₂ into the atmosphere. However, the relative portion of ocean absorption descends much faster with annual growth rate of CO₂ release than biospheric absorption does, due to a difference in dynamics. The equilibrium absorption capacity of the biosphere is estimated to be only one quarter of that of the ocean, but the current sink size of the biosphere is about half of that of the ocean.Apart from CO₂-stimulated carbon fixation, the biosphere releases CO₂ as a result of land use changes, in particular after deforestation. Both of these fluxes are of the order of 1–1.5 Pg of carbon per year. The CO₂-fertilization effect and regrowth together have turned the terrestrial biosphere as a whole from a source into a sink.     

Acclimation response of spring wheat in a free-air CO2 enrichment (FACE) atmosphere with variable soil nitrogen regimes. 2. Net assimilation and stomatal conductance of leaves

Atmospheric CO₂ concentration continues to rise. It is important, therefore, to determine what acclimatory changes will occur within the photosynthetic apparatus of wheat (Triticum aestivum L. cv. Yecora Rojo) grown in a future high-CO₂ world at ample and limited soil N contents. Wheat was grown in an open field exposed to the CO₂ concentration of ambient air [370 μmol (CO₂) mol⁻¹; Control] and air enriched to ∼200 μmol (CO₂) mol⁻¹ above ambient using a Free-Air CO₂ Enrichment (FACE) apparatus (main plot). A High (35 g m⁻²) or Low (7 and 1.5 g m⁻² for 1996 and 1997, respectfully) level of N was applied to each half of the main CO₂ treatment plots (split-plot). Under High-N, FACE reduced stomatal conductance (g _s) by 30% at mid-morning (2 h prior to solar noon), 36% at midday (solar noon) and 27% at mid-afternoon (2.5 h after solar noon), whereas under Low-N, g _s was reduced by as much as 31% at mid-morning, 44% at midday and 28% at mid-afternoon compared with Control. But, no significant CO₂ × N interaction effects occurred. Across seasons and growth stages, daily accumulation of carbon (A′) was 27% greater in FACE than Control. High-N increased A′ by 18% compared with Low-N. In contrast to results for g _s, however, significant CO₂ × N interaction effects occurred because FACE increased A′ by 30% at High-N, but by only 23% at Low-N. FACE enhanced the seasonal accumulation of carbon (A′′) by 29% during 1996 (moderate N-stress), but by only 21% during 1997 (severe N-stress). These results support the premise that in a future high-CO₂ world an acclimatory (down-regulation) response in the photosynthetic apparatus of field-grown wheat is anticipated. They also demonstrate, however, that the stimulatory effect of a rise in atmospheric CO₂ on carbon gain in wheat can be maintained if nutrients such as nitrogen are in ample supply. This revised version was published online in June 2006 with corrections to the Cover Date.     

Photosynthetic and stomatal responses of two tropical and two temperate trees to atmospheric humidity

The effects of leaf to air vapour pressure differences (ΔW) on net photosynthetic rate (P_N) and stomatal conductance (g_s) were examined in the leaves of two tropical rain forest trees, Eugenia grandis and Pongamia pinnata, and two temperate evergreen trees, Viburnum awabuki and Daphniphyllum macropodum. A single leaf was set inside a small chamber and ΔW was varied from 7 to 24 mmol mol^-1 at 25 and 500 μmol m^-2 s^-1 of photon flux density. P_N and g_s of the two tropical rain forest trees decreased with increasing ΔW, while the two temperate evergreen trees were not highly responsive to ΔW. P. pinnata was more sensitive to ΔW in its stomatal response, and had a higher stomatal density and higher stomatal index than did the two temperate trees and another tropical tree. Significant reductions i n g_s and intercellular CO₂ concentrations in the two tropical trees at high ΔW suggest that the decline of P_N was due to the decrease in g_s. The responses of P_N and g_s indicated that the tropical trees were more sensitive to ΔW than were the temperate ones. This revised version was published online in September 2006 with corrections to the Cover Date.