Co-culture with Synechococcus facilitates growth of Prochlorococcus under ocean acidification conditions

Anthropogenic CO₂ emissions are projected to lower the pH of the ocean 0.3 units by 2100. Previous studies suggested that Prochlorococcus and Synechococcus, the numerically dominant phytoplankton in the oceans, have different responses to elevated CO₂ that may result in a dramatic shift in their relative abundances in future oceans. Here we showed that the exponential growth rates of these two genera respond to future CO₂ conditions in a manner similar to other cyanobacteria, but Prochlorococcus strains had significantly lower realized growth rates under elevated CO₂ regimes due to poor survival after exposure to fresh culture media. Despite this, a Synechococcus strain was unable to outcompete a Prochlorococcus strain in co-culture at elevated CO₂. Under these conditions, Prochlorococcus' poor response to elevated CO₂ disappeared, and Prochlorococcus' relative fitness showed negative frequency dependence, with both competitors having significant fitness advantages when initially rare. These experiments suggested that the two strains should be able to coexist indefinitely in co-culture despite sharing nearly identical nutritional requirements. We speculate that negative frequency dependence exists due to reductive Black Queen evolution that has resulted in a passively mutualistic relationship analogous to that connecting Prochlorococcus with the ‘helper’ heterotrophic microbes in its environment.     

Seasonal patterns of photosynthesis in Douglas fir seedlings during the third and fourth year of exposure to elevated CO2 and temperature

The interactive effects of elevated atmospheric CO₂ and temperature on seasonal patterns of photosynthesis in Douglas fir (Psuedotsuga menziesii (Mirb.) Franco) seedlings were examined. Seedlings were grown in sunlit chambers controlled to track either ambient (~400 p.p.m.) CO₂ or ambient +200 p.p.m. CO₂, and either ambient temperature or ambient +4 °C. Light‐saturated net photosynthetic rates were measured approximately monthly over a 21 month period. Elevated CO₂ increased net photosynthetic rates by an average of 21% across temperature treatments during both the 1996 hydrologic year, the third year of exposure, and the 1997 hydrologic year. Elevated mean annual temperature increased net photosynthetic rates by an average of 33% across CO₂ treatments during both years. Seasonal temperature changes also affected net photosynthetic rates. Across treatments, net photosynthetic rates were highest in the spring and autumn, and lowest in July, August and December–January. Seasonal increases in temperature were not correlated with increases in the relative photosynthetic response to elevated CO₂. Seasonal shifts in the photosynthetic temperature optimum reduced temperature effects on the relative response to elevated CO₂. These results suggest that the effects of elevated CO₂ on net photosynthetic rates in Douglas fir are largely independent of temperature.     

Effects of elevated CO2 and temperature on photosynthesis and leaf traits of an understory dwarf bamboo in subalpine forest zone,China

The dwarf bamboo (Fargesia rufa Yi), growing understory in subalpine dark coniferous forest, is one of the main foods for giant panda, and it influences the regeneration of subalpine coniferous forests in southwestern China. To investigate the effects of elevated CO₂, temperature and their combination, the dwarf bamboo plantlets were exposed to two CO₂ regimes (ambient and double ambient CO₂ concentration) and two temperatures (ambient and +2.2°C) in growth chambers. Gas exchange, leaf traits and carbohydrates concentration were measured after the 150‐day experiment. Elevated CO₂ significantly increased the net photosynthetic rate (A_net), intrinsic water‐use efficiency (WUE_i) and carbon isotope composition (δ¹³C) and decreased stomatal conductance (g_s) and total chlorophyll concentration based on mass (Chl_m) and area (Chl_a). On the other hand, elevated CO₂ decreased specific leaf area (SLA), which was increased by elevated temperature. Elevated CO₂ also increased foliar carbon concentration based on mass (C_m) and area (C_a), nitrogen concentration based on area (N_a), carbohydrates concentration (i.e. sucrose, sugar, starch and non‐structural carbohydrates) and the slope of the A_net–N_a relationship. However, elevated temperature decreased C_m, C_a and N_a. The combination of elevated CO₂ and temperature hardly affected SLA, C_m, C_a, N_m, N_a, Chl_m and Chl_a. Variables A_net and N_a had positive linear relationships in all treatments. Our results showed that photosynthetic acclimation did not occur in dwarf bamboo at elevated CO₂ and it could adjust physiology and morphology to enable the capture of more light, to increase WUE and improve nutritional conditions.     

Predicting responses of photosynthesis and root fraction to elevated [CO2]a: interactions among carbon,nitrogen, and growth*

At elevated atmospheric CO₂ concentrations ([CO₂]_a), photosynthetic capacity (A_max) and root fraction (η_R, the ratio of root to plant dry mass) increased in some studies and decreased in others. Here, we have explored possible causes of this, focusing on the relative magnitudes of the effects of elevated [CO₂]_a on specific leaf (n_m) and plant (n_p) nitrogen concentrations, leaf mass per unit area (h), and plant nitrogen productivity (α). In our survey of 39 studies with 35 species, we found that elevated [CO₂]_a led to decreased n_m and n_p in all the studies and to increased h and α in most of the studies. The magnitudes of these changes varied with species and with experimental conditions. Based on a model that integrated [CO₂]_a-induced changes in leaf nitrogen into a biochemically based model of leaf photosynthesis, we predicted that, to a first approximation, photosynthesis will be upregulated (A_max will increase) when growth at increased [CO₂]_a leads to increases in h that are larger than decreases in n_m. Photosynthesis will be downregulated (A_max will decrease) when increases in h are smaller than decreases in n_m. The model suggests that photosynthetic capacity increases at elevated [CO₂]_a only when additional leaf mesophyll more than compensates the effects of nitrogen dilution. We considered two kinds of regulatory paradigms that could lead to varying responses of η_R to elevated [CO₂]_a, and compared the predictions of each with the data. A simple static model based on the functional balance concept predicts that η_R should increase when neither n_p nor h is very responsive to elevated [CO₂]_a. The quantitative and qualitative agreement of the predictions with data from the literature, however, is poor. A model that predicts η_R from the relative sensitivities of photosynthesis and relative growth rate to elevated [CO₂]_a corresponds much more closely to the observations. In general, root fraction increases if the response of photosynthesis to [CO₂]_a is greater than that of relative growth rate.     

EFFECTS OF CO2 ENRICHMENT ON THE BLOOM‐FORMING CYANOBACTERIUM MICROCYSTIS AERUGINOSA (CYANOPHYCEAE): PHYSIOLOGICAL RESPONSES AND RELATIONSHIPS WITH THE AVAILABILITY OF DISSOLVED INORGANIC CARBON1

Microcystis aeruginosa Kütz. 7820 was cultured at 350 and 700 μL·L ^{? 1} CO₂ to assess the impacts of doubled atmospheric CO₂ concentration on this bloom‐forming cyanobacterium. Doubling of CO₂ concentration in the airflow enhanced its growth by 52%–77%, with pH values decreased and dissolved inorganic carbon (DIC) increased in the medium. Photosynthetic efficiencies and dark respiratory rates expressed per unit chl a tended to increase with the doubling of CO₂. However, saturating irradiances for photosynthesis and light‐saturated photosynthetic rates normalized to cell number tended to decrease with the increase of DIC in the medium. Doubling of CO₂ concentration in the airflow had less effect on DIC‐saturated photosynthetic rates and apparent photosynthetic affinities for DIC. In the exponential phase, CO₂ and HCO₃ ^? levels in the medium were higher than those required to saturate photosynthesis. Cultures with surface aeration were DIC limited in the stationary phase. The rate of CO₂ dissolution into the liquid increased proportionally when CO₂ in air was raised from 350 to 700 μL·L ^{? 1}, thus increasing the availability of DIC in the medium and enhancing the rate of photosynthesis. Doubled CO₂ could enhance CO₂ dissolution, lower pH values, and influence the ionization fractions of various DIC species even when the photosynthesis was not DIC limited. Consequently, HCO₃ ^? concentrations in cultures were significantly higher than in controls, and the photosynthetic energy cost for the operation of CO₂ concentrating mechanism might decrease.     

Exposures to elevated CO2, elevated temperature and enhanced UV-B radiation modify activities of polyphenol oxidase and guaiacol peroxidase and concentrations of chlorophylls,polyamines and soluble proteins in the leaves of Betula pendula seedlings

The activity of polyphenol oxidase (PPO) and guaiacol peroxidase (POD) and the concentrations of chlorophylls, free polyamines and soluble proteins were determined from the leaves of six genotypes of silver birch (Betula pendula Roth) seedlings exposed to short-term elevated carbon dioxide (CO₂), temperature (T), ultraviolet-B irradiation (UV-B, 280-315 nm) and their combinations. Results showed that the activity of PPO in the leaves was low but increased by elevated CO₂ and elevated T. The POD activity varied between the genotypes due to an interactive effect of CO₂ × UV-B. The soluble proteins were clearly decreased by elevated CO₂, but the level of response varied among the genotypes. The concentrations of chl a and total chlorophylls were lower in the leaves treated with elevated CO₂ than in leaves grown at ambient CO₂. An interactive effect of CO₂ × UV-B on the chl a/b ratio was found. Elevated T increased chl b concentration and decreased chl a/b ratio. Temperature treatments also caused variation in the concentrations of chl a, chl b and total chlorophylls among the genotypes. Polyamine analyses showed that the concentrations of putrescine were increased and spermine decreased in leaves treated with elevated T. However, the change in putrescine by elevated T was clearer at ambient CO₂ than in eCO₂ environment (significant effect of T × CO₂). In conclusion, the defensive enzymes, photosynthetic pigments, soluble proteins and growth-regulating polyamines in silver birch leaves were not susceptible to enhanced UV-B radiation. In contrast, all the variables responded to elevated T and/or elevated CO₂, reflecting the enhancive effects of climate change conditions not only on leaf productivity, but also on leaf turn-over rate. Most of these climate-driven changes were not regulated by UV-B radiation.     

BENTHIC AND PLANKTONIC ALGAL COMMUNITIES IN A HIGH ARCTIC LAKE: PIGMENT STRUCTURE AND CONTRASTING RESPONSES TO NUTRIENT ENRICHMENT1

We investigated the fine pigment structure and composition of phytoplankton and benthic cyanobacterial mats in Ward Hunt Lake at the northern limit of High Arctic Canada and the responses of these two communities to in situ nutrient enrichment. The HPLC analyses showed that more than 98% of the total pigment stocks occurred in the benthos. The phytoplankton contained Chrysophyceae, low concentrations of other protists and Cyanobacteria (notably picocyanobacteria), and the accessory pigments chl c₂, fucoxanthin, diadinoxanthin, violaxanthin, and zeaxanthin. The benthic community contained the accessory pigments chl b, chl c₂, and a set of carotenoids dominated by glycosidic xanthophylls, characteristic of filamentous cyanobacteria. The black surface layer of the mats was rich in the UV‐screening compounds scytonemin, red scytonemin‐like, and mycosporine‐like amino acids, and the blue‐green basal stratum contained high concentrations of light‐harvesting pigments. In a first bioassay of the benthic mats, there was no significant photosynthetic or growth response to inorganic carbon or full nutrient enrichment over 15 days. This bioassay was repeated with increased replication and HPLC analysis in a subsequent season, and the results confirmed the lack of significant response to added nutrients. In contrast, the phytoplankton in samples from the overlying water column responded strongly to enrichment, and chl a biomass increased by a factor of 19.2 over 2 weeks. These results underscore the divergent ecophysiology of benthic versus planktonic communities in extreme latitudes and show that cold lake ecosystems can be dominated by benthic phototrophs that are nutrient sufficient despite their ultraoligotrophic overlying waters.     

PHOTOACCLIMATION OF PHOTOSYNTHESIS IRRADIANCE RESPONSE CURVES AND PHOTOSYNTHETIC PIGMENTS IN MICROALGAE AND CYANOBACTERIA1

The photosynthesis‐irradiance response (PE) curve, in which mass‐specific photosynthetic rates are plotted versus irradiance, is commonly used to characterize photoacclimation. The interpretation of PE curves depends critically on the currency in which mass is expressed. Normalizing the light‐limited rate to chl a yields the chl a‐specific initial slope (α^chl). This is proportional to the light absorption coefficient (a^chl), the proportionality factor being the photon efficiency of photosynthesis (φ_m). Thus, α^chl is the product of a^chl and φ_m. In microalgae α^chl typically shows little (<20%) phenotypic variability because declines of φ_m under conditions of high‐light stress are accompanied by increases of a^chl. The variation of α^chl among species is dominated by changes in a^chl due to differences in pigment complement and pigment packaging. In contrast to the microalgae, α^chl declines as irradiance increases in the cyanobacteria where phycobiliproteins dominate light absorption because of plasticity in the phycobiliprotein:chl a ratio. By definition, light‐saturated photosynthesis (P_m) is limited by a factor other than the rate of light absorption. Normalizing P_m to organic carbon concentration to obtain P_m^C allows a direct comparison with growth rates. Within species, P_m^C is independent of growth irradiance. Among species, P_m^C covaries with the resource‐saturated growth rate. The chl a:C ratio is a key physiological variable because the appropriate currencies for normalizing light‐limited and light‐saturated photosynthetic rates are, respectively, chl a and carbon. Typically, chl a:C is reduced to about 40% of its maximum value at an irradiance that supports 50% of the species‐specific maximum growth rate and light‐harvesting accessory pigments show similar or greater declines. In the steady state, this down‐regulation of pigment content prevents microalgae and cyanobacteria from maximizing photosynthetic rates throughout the light‐limited region for growth. The reason for down‐regulation of light harvesting, and therefore loss of potential photosynthetic gain at moderately limiting irradiances, is unknown. However, it is clear that maximizing the rate of photosynthetic carbon assimilation is not the only criterion governing photoacclimation.     

Direct and indirect effects of elevated atmospheric CO2 on net ecosystem production in a Chesapeake Bay tidal wetland

The rapid increase in atmospheric CO₂ concentrations (C_a) has resulted in extensive research efforts to understand its impact on terrestrial ecosystems, especially carbon balance. Despite these efforts, there are relatively few data comparing net ecosystem exchange of CO₂ between the atmosphere and the biosphere (NEE), under both ambient and elevated C_a. Here we report data on annual sums of CO₂ (NEE_net) for 19 years on a Chesapeake Bay tidal wetland for Scirpus olneyi (C₃ photosynthetic pathway)‐ and Spartina patens (C₄ photosynthetic pathway)‐dominated high marsh communities exposed to ambient and elevated C_a (ambient + 340 ppm). Our objectives were to (i) quantify effects of elevated C_a on seasonally integrated CO₂ assimilation (NEE_net = NEE_day + NEE_night, kg C m^?2 y^?1) for the two communities; and (ii) quantify effects of altered canopy N content on ecosystem photosynthesis and respiration. Across all years, NEE_net averaged 1.9 kg m^?2 y^?1 in ambient C_a and 2.5 kg m^?2 y^?1 in elevated C_a, for the C₃‐dominated community. Similarly, elevated C_a significantly (P < 0.01) increased carbon uptake in the C₄‐dominated community, as NEE_net averaged 1.5 kg m^?2 y^?1 in ambient C_a and 1.7 kg m^?2 y^?1 in elevated C_a. This resulted in an average CO₂ stimulation of 32% and 13% of seasonally integrated NEE_net for the C₃‐ and C₄‐dominated communities, respectively. Increased NEE_day was correlated with increased efficiencies of light and nitrogen use for net carbon assimilation under elevated C_a, while decreased NEE_night was associated with lower canopy nitrogen content. These results suggest that rising C_a may increase carbon assimilation in both C₃‐ and C₄‐dominated wetland communities. The challenge remains to identify the fate of the assimilated carbon.     

Effects of climate change on labile and structural carbon in Douglas-fir needles as estimated by δ13C and Carea measurements

Models of photosynthesis, respiration, and export predict that foliar labile carbon (C) should increase with elevated CO₂ but decrease with elevated temperature. Sugars, starch, and protein can be compared between treatments, but these compounds make up only a fraction of the total labile pool. Moreover, it is difficult to assess the turnover of labile carbon between years for evergreen foliage. Here, we combined changes in foliar C_area (C concentration on an areal basis) as needles aged with changes in foliar isotopic composition (δ¹³C) caused by inputs of ¹³C‐depleted CO₂ to estimate labile and structural C in needles of different ages in a four‐year, closed‐chamber mesocosm experiment in which Douglas‐fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings were exposed to elevated temperature (ambient + 3.5 °C) and CO₂ (ambient + 179 ppm). Declines in δ ¹³C of needle cohorts as they aged indicated incorporation of newly fixed labile or structural carbon. The δ ¹³C calculations showed that new C was 41 ± 2% and 28 ± 3% of total needle carbon in second‐ and third‐year needles, respectively, with higher proportions of new C in elevated than ambient CO₂ chambers (e.g. 42 ± 2% vs. 37 ± 6%, respectively, for second‐year needles). Relative to ambient CO₂, elevated CO₂ increased labile C in both first‐ and second‐year needles. Relative to ambient temperature, elevated temperature diminished labile C in second‐year needles but not in first‐year needles, perhaps because of differences in sink strength between the two needle age classes. We hypothesize that plant‐soil feedbacks on nitrogen supply contributed to higher photosynthetic rates under elevated temperatures that partly compensated for higher turnover rates of labile C. Strong positive correlations between labile C and sugar concentrations suggested that labile C was primarily determined by carbohydrates. Labile C was negatively correlated with concentrations of cellulose and protein. Elevated temperature increased foliar %C, possibly due to a shift of labile constituents from low %C carbohydrates to relatively high %C protein. Decreased sugar concentrations and increased nitrogen concentrations with elevated temperature were consistent with this explanation. Because foliar constituents that vary in isotopic signature also vary in concentrations with leaf age or environmental conditions, inferences of c_i/c_a values from δ ¹³C of bulk leaf tissue should be done cautiously. Tracing of ¹³C through foliar carbon pools may provide new insight into foliar C constituents and turnover.     

DIEL PATTERNS OF GROWTH AND DIVISION IN MARINE PICOPLANKTON IN CULTURE

The effect of a 12:12-h light:dark (LD) cycle on the phasing of several cell parameters was explored in a variety of marine picophytoplanktonic strains. These included the photosynthetic prokaryotes Prochlorococcus (strains MED 4, PCC 9511, and SS 120) and Synechococcus (strains ALMO 03, ROS 04, WH 7803, and WH 8103) and five picoeukaryotes (Bathycoccus prasinos Eikrem et Throndsen, Bolidomonas pacifica Guillou et Chrétiennot-Dinet, Micromonas pusilla Manton et Parke, Pelagomonas calceolata Andersen et Saunders, and Pycnococcus provasolii Guillard et al.). Flow cytometric analysis was used to determine the relationship between cell light scatter, pigment fluorescence, DNA (when possible), and the LD cycle in these organisms. As expected, growth and division were tightly coupled to the LD cycle for all of these strains. For both Prochlorococcus and picoeukaryotes, chl and intracellular carbon increased throughout the light period as estimated by chl fluorescence and light scatter, respectively. In response to cell division, these parameters decreased regularly during the early part of the dark period, a decrease that either continued throughout the dark period or stopped for the second half of the dark period. For Synechococcus, the decrease of chl and scatter occurred earlier (in the middle of the light period), and for some strains these cellular parameters remained constant throughout the dark period. The timing of division was very similar for all picoeukaryotes and occurred just before the subjective dusk, whereas it was more variable between the different Prochlorococcus and Synechococcus strains. The burst of division for Prochlorococcus SS 120 and PCC 9511 was recorded at the subjective dusk, whereas the MED 4 strain divided later at night. Synechococcus ALMO 03, ROS 04, and WH 7803, which have a low phycourobilin to phycoerythrobilin (PUB:PEB) ratio, divided earlier, and their division was restricted to the light period. In contrast, the high PUB:PEB Synechococcus strain WH 8103 divided preferentially at night. There was a weak linear relationship between the FALS_max:FALS_min ratio and growth rate calculated from cell counts (r = 0.83, n = 11, P < 0.05). Because of the significance of picoplanktonic populations in marine systems, these results should help to interpret diel variations in oceanic optical properties in regions where picoplankton dominates.     

Differential effects of elevated CO2 on acorn density, weight, germination, and predation among three oak species in a scrub-oak forest

Much research on the effects of elevated CO₂ on forest trees has focused on quantitative changes in photosynthesis, secondary chemistry, and plant biomass. However, plant fitness responses to rising CO₂ should also include quantitative measures of reproduction, since most forest systems are recruitment limited. Until now, it has proved very difficult to grow forest trees to sexual maturity in a CO₂‐enriched environment. This paper is the first of its kind to address the effects of elevated CO₂ on the reproduction of hardwood trees in a natural forest. Beginning in 1996, scrub‐oak vegetation, predominantly three species of scrub‐oaks, Quercus myrtifolia, Q. chapmanii, and Q. geminata, were grown inside eight chambers with elevated CO₂ (704 parts per million (ppm)) and eight with ambient CO₂ (379 ppm) at Kennedy Space Center, Florida. In elevated CO₂, acorn production increased significantly for the dominant species Q. myrtifolia and for Q. chapmanii, but it did not increase for the subdominant, Q. geminata. Acorn weight, germination rate, and predation by weevils were unaffected by CO₂. Thus, recruitment of some forest tree species into the Florida scrub‐oak community is likely to be accelerated in an atmosphere of increased CO₂. However, because the acceleration of recruitment differs among species, over the long term, Q. myrtifolia and Q. chapmanii will be favored over Q. geminata and this is likely to change patterns of species diversity.     

Effects of elevated atmospheric [CO2] on instantaneous transpiration efficiency at leaf and canopy scales in Eucalyptus saligna

Rising atmospheric concentrations of CO₂ (C_a) can reduce stomatal conductance and transpiration rate in trees, but the magnitude of this effect varies considerably among experiments. The theory of optimal stomatal behaviour predicts that the ratio of photosynthesis to transpiration (instantaneous transpiration efficiency, ITE) should increase in proportion to C_a. We hypothesized that plants regulate stomatal conductance optimally in response to rising C_a. We tested this hypothesis with data from young Eucalyptus saligna Sm. trees grown in 12 climate‐controlled whole‐tree chambers for 2 years at ambient and elevated C_a. Elevated C_a was ambient + 240 ppm, 60% higher than ambient C_a. Leaf‐scale gas exchange was measured throughout the second year of the study and leaf‐scale ITE increased by 60% under elevated C_a, as predicted. Values of leaf‐scale ITE depended strongly on vapour pressure deficit (D) in both CO₂ treatments. Whole‐canopy CO₂ and H₂O fluxes were also monitored continuously for each chamber throughout the second year. There were small differences in D between C_a treatments, which had important effects on values of canopy‐scale ITE. However, when C_a treatments were compared at the same D, canopy‐scale ITE was consistently increased by 60%, again as predicted. Importantly, leaf and canopy‐scale ITE were not significantly different, indicating that ITE was not scale‐dependent. Observed changes in transpiration rate could be explained on the basis that ITE increased in proportion to C_a. The effect of elevated C_a on photosynthesis increased with rising D. At high D, C_a had a large effect on photosynthesis and a small effect on transpiration rate. At low D, in contrast, there was a small effect of C_a on photosynthesis, but a much larger effect on transpiration rate. If shown to be a general response, the proportionality of ITE with C_a will allow us to predict the effects of C_a on transpiration rate.     

SHORT‐ AND LONG‐TERM EFFECTS OF ELEVATED CO2 ON PHOTOSYNTHESIS AND RESPIRATION IN THE MARINE MACROALGA HIZIKIA FUSIFORMIS (SARGASSACEAE,PHAEOPHYTA) GROWN AT LOW AND HIGH N SUPPLIES1

The short‐term and long‐term effects of elevated CO₂ on photosynthesis and respiration were examined in cultures of the marine brown macroalga Hizikia fusiformis (Harv.) Okamura grown under ambient (375 μL · L^?1) and elevated (700 μL · L^?1) CO₂ concentrations and at low and high N availability. Short‐term exposure to CO₂ enrichment stimulated photosynthesis, and this stimulation was maintained with prolonged growth at elevated CO₂, regardless of the N levels in culture, indicating no down‐regulation of photosynthesis with prolonged growth at elevated CO₂. However, the photosynthetic rate of low‐N‐grown H. fusiformis was more responsive to CO₂ enrichment than that of high‐N‐grown algae. Elevation of CO₂ concentration increased the value of K_1/2(Ci) (the half‐saturation constant) for photosynthesis, whereas high N supply lowered it. Neither short‐term nor long‐term CO₂ enrichment had inhibitory effects on respiration rate, irrespective of the N supply, under which the algae were grown. Under high‐N growth, the Q₁₀ value of respiration was higher in the elevated‐CO₂‐grown algae than the ambient‐CO₂‐grown algae. Either short‐ or long‐term exposure to CO₂ enrichment decreased respiration as a proportion of gross photosynthesis (Pg) in low‐N‐grown H. fusiformis. It was proposed that in a future world of higher atmospheric CO₂ concentration and simultaneous coastal eutrophication, the respiratory carbon flux would be more sensitive to changing temperature.     

Leaf senescence of Quercus myrtifolia as affected by long-term CO2 enrichment in its native environment

The long‐term effects of elevated (ambient plus 350 μmol mol^?1) atmospheric CO₂ concentration (C_a) on the leaf senescence of Quercus myrtifolia Willd was studied in a scrub‐oak community during the transition from autumn (December 1997) to spring (April 1998). Plants were grown in large open‐top chambers at the Smithsonian CO₂ Research Site, Merritt Island Wildlife Refuge, Cape Canaveral, Florida. Chlorophyll (a + b) concentration, Rubisco activity and N concentration decreased by 75%, 82%, and 52%, respectively, from December (1997) to April (1998) in the leaves grown at ambient C_a. In contrast, the leaves of plants grown at elevated C_a showed no significant decrease in chlorophyll (a + b) concentration or Rubisco activity, and only a 25% reduction in nitrogen. These results indicate that leaf senescence was delayed during this period at elevated C_a. Delayed leaf senescence in elevated C_a had important consequences for leaf photosynthesis. In elevated C_a the net photosynthetic rate of leaves that flushed in Spring 1997 (last year's leaves) and were 13 months old was not different from fully‐expanded leaves that flushed in 1998, and were approximately 1 month old (current year's leaves). In ambient C_a the net photosynthetic rate of last year's leaves was 54% lower than for current year's leaves. When leaves were fully senesced, nitrogen concentration decreased to about 40% of the concentration in non‐senesced leaves, in both CO₂ treatments. In April, net photosynthesis was 97% greater in leaves grown in elevated C_a than in those grown at ambient. During the period when elevated C_a delayed leaf senescence, more leaves operating at higher photosynthetic rate would allow the ecosystem dominated by Q. myrtifolia to gain more carbon at elevated C_a than at ambient C_a.     

Effect of irradiance and temperature on the photosynthesis of a cultivated red alga,Pyropia tenera (= Porphyra tenera), at the southern limit of distribution in Japan

The effect of irradiance and temperature on the photosynthesis of the red alga, Pyropia tenera, was determined for maricultured gametophytes and sporophytes collected from a region that is known as one of the southern limits of its distribution in Japan. Macroscopic gametophytes were examined using both pulse‐amplitude modulated fluorometry and/or dissolved oxygen sensors. A model of the net photosynthesis–irradiance (P‐E) relationship of the gametophytes at 12°C revealed that the net photosynthetic rate quickly increased at irradiances below the estimated saturation irradiance of 46 μmol photons m^?2 s^?1, and the compensation irradiance was 9 μmol photons m^?2 s^?1. Gross photosynthesis and dark respiration for the gametophytes were also determined over a range of temperatures (8–34°C), revealing that the gross photosynthetic rates of 46.3 μmol O₂ mg_chl‐a^?1 min^?1 was highest at 9.3 (95% Bayesian credible interval (BCI): 2.3–14.5)°C, and the dark respiration rate increased at a rate of 0.93 μmol O₂ mg_chl‐a^?1 min^?1°C^?1. The measured dark respiration rates ranged from ?0.06 μmol O₂ mg_chl‐a^?1 min^?1 at 6°C to ?25.2 μmol O₂ mg_chl‐a^?1 min^?1 at 34°C. The highest value of the maximum quantum yield (Fv/Fm) for the gametophytes occurred at 22.4 (BCI: 21.5–23.3) °C and was 0.48 (BCI: 0.475–0.486), although those of the sporophyte occurred at 12.9 (BCI: 7.4–15.1) °C and was 0.52 (BCI: 0.506–0.544). This species may be considered well‐adapted to the current range of seawater temperatures in this region. However, since the gametophytes have such a low temperature requirement, they are most likely close to their tolerable temperatures in the natural environment.     

Post‐fire resprouting responses of native and exotic grasses from Cumberland Plain Woodland (Sydney,Australia) under elevated carbon dioxide

This study investigated the effect of elevated CO₂ on the post‐fire resprouting response of a grassland system of perennial grass species of Cumberland Plain Woodland. Plants were grown in mixtures in natural soil in mesocosms, each containing three exotic grasses (Nassella neesiana, Chloris gayana, Eragrostis curvula) and three native grasses (Themeda australis, Microlaena stipoides, Chloris ventricosa) under elevated (700 ppm) and ambient (385 ppm) CO₂ conditions. Resprouting response after fire at the community‐ and species‐level was assessed. There was no difference in community‐level biomass between CO₂ treatments; however, exotic species made up a larger proportion of the community biomass under all treatments. There were species‐level responses to elevated CO₂ but no significant interactions found between CO₂ and burning or plant status. Two exotic grasses (N. neesiana and E. curvula, a C₃ and a C₄ species respectively), and one native grass (M. stipoides, a C₃ species) significantly increased in biomass, and a native C₄ grass (C. ventricosa) significantly decreased in biomass under elevated CO₂. These results suggest that although overall productivity of this community may not change with increases in CO₂ and fire frequency, the community composition may alter due to differential species responses.     

Interactive effects of elevated CO2 and soil fertility on isoprene emissions from Quercus robur

The effects of global change on the emission rates of isoprene from plants are not clear. A factor that can influence the response of isoprene emission to elevated CO₂ concentrations is the availability of nutrients. Isoprene emission rate under standard conditions (leaf temperature: 30°C, photosynthetically active radiation (PAR): 1000 μmol photons m^?2 s^?1), photosynthesis, photosynthetic capacity, and leaf nitrogen (N) content were measured in Quercus robur grown in well‐ventilated greenhouses at ambient and elevated CO₂ (ambient plus 300 ppm) and two different soil fertilities. The results show that elevated CO₂ enhanced photosynthesis but leaf respiration rates were not affected by either the CO₂ or nutrient treatments. Isoprene emission rates and photosynthetic capacity were found to decrease with elevated CO₂, but an increase in nutrient availability had the converse effect. Leaf N content was significantly greater with increased nutrient availability, but unaffected by CO₂. Isoprene emission rates measured under these conditions were strongly correlated with photosynthetic capacity across the range of different treatments. This suggests that the effects of CO₂ and nutrient levels on allocation of carbon to isoprene production and emission under near‐saturating light largely depend on the effects on photosynthetic electron transport capacity.     

Pea aphid promotes amino acid metabolism both in Medicago truncatula and bacteriocytes to favor aphid population growth under elevated CO2

Rising atmospheric CO₂ levels can dilute the nitrogen (N) resource in plant tissue, which is disadvantageous to many herbivorous insects. Aphids appear to be an exception that warrants further study. The effects of elevated CO₂ (750 ppm vs. 390 ppm) were evaluated on N assimilation and transamination by two Medicago truncatula genotypes, a N‐fixing‐deficient mutant (dnf1) and its wild‐type control (Jemalong), with and without pea aphid (Acyrthosiphon pisum) infestation. Elevated CO₂ increased population abundance and feeding efficiency of aphids fed on Jemalong, but reduced those on dnf1. Without aphid infestation, elevated CO₂ increased photosynthetic rate, chlorophyll content, nodule number, biomass, and pod number for Jemalong, but only increased pod number and chlorophyll content for dnf1. Furthermore, aphid infested Jemalong plants had enhanced activities of N assimilation‐related enzymes (glutamine synthetase, Glutamate synthase) and transamination‐related enzymes (glutamate oxalate transaminase, glutamine phenylpyruvate transaminase), which presumably increased amino acid concentration in leaves and phloem sap under elevated CO₂. In contrast, aphid infested dnf1 plants had decreased activities of N assimilation‐related enzymes and transmination‐related enzymes and amino acid concentrations under elevated CO₂. Furthermore, elevated CO₂ up‐regulated expression of genes relevant to amino acid metabolism in bacteriocytes of aphids associated with Jemalong, but down‐regulated those associated with dnf1. Our results suggest that pea aphids actively elicit host responses that promote amino acid metabolism in both the host plant and in its bacteriocytes to favor the population growth of the aphid under elevated CO₂.     

Seasonal response of photosynthesis to elevated CO2 in loblolly pine (Pinus taeda L.) over two growing seasons

Trees growing in natural systems undergo seasonal changes in environmental factors that generate seasonal differences in net photosynthetic rates. To examine how seasonal changes in the environment affect the response of net photosynthetic rates to elevated CO₂, we grew Pinus taeda L. seedlings for three growing seasons in open-top chambers continuously maintained at either ambient or ambient + 30 Pa CO₂. Seedlings were grown in the ground, under natural conditions of light, temperature nd nutrient and water availability. Photosynthetic capacity was measured bimonthly using net photosynthetic rate vs. intercellular CO₂ partial pressure (A-C_i) curves. Maximum Rubisco activity (Vc_max) and ribulose 1,5-bisphosphate regeneration capacity mediated by electron transport (J_max) and phosphate regeneration (PiRC) were calculated from A-C_i curves using a biochemically based model. Rubisco activity, activation state and content, and leaf carbohydrate, chlorophyll and nitrogen concentrations were measured concurrently with photosynthesis measurements. This paper presents results from the second and third years of treatment. Mean leaf nitrogen concentrations ranged from 13.7 to 23.8 mg g^?1, indicating that seedlings were not nitrogen deficient. Relative to ambient CO₂ seedlings, elevated CO₂ increased light-saturated net photosynthetic rates 60–110% during the summer, but < 30% during the winter. A relatively strong correlation between leaf temperature and the relative response of net photosynthetic rates to elevated CO₂ suggests a strong effect of leaf temperature. During the third growing season, elevated CO₂ reduced Rubisco activity 30% relative to ambient CO₂ seedlings, nearly completely balancing Rubisco and RuBP-regeneration regulation of photosynthesis. However, reductions in Rubisco activity did not eliminate the seasonal pattern in the relative response of net photosynthetic rates to elevated CO₂. These results indicate that seasonal differences in the relative response of net photosynthetic rates to elevated CO₂ are likely to occur in natural systems.