Nitrogen supply as a factor influencing photoinhibition and photosynthetic acclimation after transfer of shade-grown Solanum dulcamara to bright light

We have compared the ability of shadegrown clones of Solamum dulcamara L. from shade and sun habitats to acclimate to bright light, as a function of nitrogen nutrition before and after transfer to bright light. Leaves of S. dulcamara grown in the shade with 0.6 mM NO ₃ ^- have similar photosynthetic properties as leaves of plants grown with 12.0 mM NO ₃ ^- . When transferred to bright light for 1–2 d the leaves of these plants show substantial photoinhibition which is characterized by about 50% decrease in apparent quantum yield and a reduction in the rate of photosynthesis in air at light saturation. Photoinhibition of leaf photosynthesis is associated with reduction in the variable component of low-temperature fluorescence emission, and with loss of in-vitro electron transport, especially of photosystem II-dependent processes.We find no evidence for ecotypic differentiation in the potential for photosynthetic acclimation among shade and sun clones of S. dulcamara, or of differentiation with respect to nitrogen requirements for acclimation. Recovery from photoinhibition and subsequent acclimation of photosynthesis to bright light only occurs in leaves of plants provided with 12.0 mM NO ₃ ^- . In these, apparent quantum yield is fully restored after 14 d, and photosynthetic acclimation is shown by an increase in light-saturated photosynthesis in air, of light-and CO₂-saturated photosynthesis, and of the initial slope of the CO₂-response curve. The latter changes are highly correlated with changes in ribulose-bisphosphate-carboxylase activity in vitro. Plants supplied with 0.6 mM NO ₃ ^- show incomplete recovery of apparent quantum yield after 14 d, but CO₂-dependent leaf photosynthetic parameters return to control levels.Symbols and abbreviations F_o initial level of fluorescence at 77 K - F_m maximum level of fluorescence at 77 K - F_v variable components of fluorescence at 77 K (F_v=F_m-F_o) - PSI, PSII photosystem I and II, respectively - RuBP ribulose-1,5-bisphosphate - RuBPCase ribulose-1,5-bisphosphate carboxylase-oxygenase (EC 4.1.1.39)     

Photosynthetic acclimation in relation to nitrogen allocation in cucumber leaves in response to changes in irradiance

Leaves deep in canopies can suddenly be exposed to increased irradiances following e.g. gap formation in forests or pruning in crops. Studies on the acclimation of photosynthesis to increased irradiance have mainly focused on the changes in photosynthetic capacity (A_max), although actual irradiance often remains below saturating level. We investigated the effect of changes in irradiance on the photosynthesis irradiance response and on nitrogen allocation in fully grown leaves of Cucumis sativus. Leaves that fully developed under low (50 µmol m^?2 s^?1) or moderate (200 µmol m^?2 s^?1) irradiance were subsequently exposed to, respectively, moderate (LM‐leaves) or low (ML‐leaves) irradiance or kept at constant irradiance level (LL‐ and MM‐leaves). Acclimation of photosynthesis occurred within 7 days with final A_max highest in MM‐leaves, lowest in LL‐leaves and intermediate in ML‐ and LM‐leaves, whereas full acclimation of thylakoid processes underlying photosystem II (PSII) efficiency and non‐photochemical quenching occurred in ML‐ and LM‐leaves. Dark respiration correlated with irradiance level, but not with A_max. Light‐limited quantum efficiency was similar in all leaves. The increase in photosynthesis at moderate irradiance in LM‐leaves was primarily driven by nitrogen import, and nitrogen remained allocated in a similar ratio to Rubisco and bioenergetics, while allocation to light harvesting relatively decreased. A contrary response of nitrogen was associated with the decrease in photosynthesis in ML‐leaves. Net assimilation of LM‐leaves under moderate irradiance remained lower than in MM‐leaves, revealing the importance of photosynthetic acclimation during the leaf developmental phase for crop productivity in scenarios with realistic, moderate fluctuations in irradiance that leaves can be exposed to.     

Photosynthetic acclimation of plants to growth irradiance: the relative importance of specific leaf area and nitrogen partitioning in maximizing carbon gain

Changes in specific leaf area (SLA, projected leaf area per unit leaf dry mass) and nitrogen partitioning between proteins within leaves occur during the acclimation of plants to their growth irradiance. In this paper, the relative importance of both of these changes in maximizing carbon gain is quantified. Photosynthesis, SLA and nitrogen partitioning within leaves was determined from 10 dicotyledonous C₃ species grown in photon irradiances of 200 and 1000 µmol m^?2 s^?1. Photosynthetic rate per unit leaf area measured under the growth irradiance was, on average, three times higher for high‐light‐grown plants than for those grown under low light, and two times higher when measured near light saturation. However, light‐saturated photosynthetic rate per unit leaf dry mass was unaltered by growth irradiance because low‐light plants had double the SLA. Nitrogen concentrations per unit leaf mass were constant between the two light treatments, but plants grown in low light partitioned a larger fraction of leaf nitrogen into light harvesting. Leaf absorptance was curvilinearly related to chlorophyll content and independent of SLA. Daily photosynthesis per unit leaf dry mass under low‐light conditions was much more responsive to changes in SLA than to nitrogen partitioning. Under high light, sensitivity to nitrogen partitioning increased, but changes in SLA were still more important.     

Leaf Factors Affecting Light-Saturated Photosynthesis in Ecotypes of Solidago virgaurea from Exposed and Shaded Habitats

Plants of Solidago virgaurea L. from exposed and shaded habitats differ with respect to the response of the photosynthetic apparatus to the level of irradiance during growth. An analysis was carried out on leaf characteristies which might be responsible for the differences established in the rates of Hght-saturated CO₂ uptake. The clones were grown in controlled environment chambers at high and low levels of irradiance. Light-saturated rates of photosynthesis and transpiration were measured at natural and lower ambient CO₂ concentrations. A low temperature dependence of light-saturated CO₂ uptake at natural CO₂ concentrations, and a strong response to changes in stomatal width, suggested that the rate of CO₂ transfer from ambient air towards reaetion sites in chloroplasts was mainly limiting the pholosynthetic rate. Resistances to transfer of CO₂ for different parts of the pathway were calculated. There was a weak but significant correlation between stomatal conductance and the product stomatal frequency ± pore length. Mesopbyll conductance and dry weight per unit area were highly correlated in leaves not damaged by high irradiance. This suggests that mesophyll conductance increases with increasing cross sectional area (per unit leaf area) of the pathways of CO₂ transfer in the mesophyll from cell surfaces to reaction sites. The higher light-saturated photosynthesis in clones from exposed habitats when grown at high irradiance than when grown at low irradiance was attributable mainly to a lower mesophyll resistance. In shade clones the effect upon CO₂ uptake of the increase in leaf thickness when grown at high irradiance was counteracted by the associated inactivation of the photosynthetic apparatus. The difference in CO₂ uptake present between clones from exposed and shaded habitats when preconditioned to high irradiance resulted from differences in both mesophyll and stomatal resistances. A few hybrid clones of an F₁-population from a cross between a clone from an exposed habitat and a clone from a shaded habitat reacted, on the whole, in the same way as the exposed habitat parent. When grown at high irradiance, the hybrid clones showed higher photosynthetic rates than either parent; this was largely attributable to the unusually low stomatal resistance of the hybrid leaves.     

Influence of Etherel and Gibberellic Acid on Carbon Metabolism,Growth, and Essential Oil Accumulation in Spearmint (Mentha Spicata)

Controlled environment chamber and glasshouse studies were conducted on six herbaceous annual species grown at 350 (AC) and 700 (EC) mol(CO₂) mol^-1 to determine whether growth at EC resulted in acclimation of the apparent quantum yield of photosynthesis (QY) measured at limiting photosynthetic photon flux density (PPFD), or in acclimation of net photosynthetic rate (P _N) measured at saturating PPFD. It was also determined whether acclimation in P _N at limiting PPFD was correlated with acclimation of carboxylation efficiency or ribulose-1,5-bisphosphate (RuBP) regeneration rate measured at saturating PPFD. Growth at EC reduced both the QY and P _N at limiting PPFD in three of the six species. The occurrence of photosynthetic acclimation measured at a rate limiting PPFD was independent of whether photosynthetic acclimation was apparent at saturating measurement PPFD. At saturating measurement PPFD, acclimation to EC in the apparent carboxylation efficiency and RuBP regeneration capacity also occurred independently. Thus at least three components of the photosynthetic system may adjust independently when leaves are grown at EC. Estimates of photosynthetic acclimation at both high and low PPFD are necessary to accurately predict photosynthesis at the whole plant or canopy level as [CO₂] increases.     

Does Rubisco control the rate of photosynthesis and plant growth? An exercise in molecular ecophysiology

Experiments are described in which tobacco (Nicotiana tabacum L.) transformed with antisense rbcS to decrease expression of ribulose-1,5-bisphosphate carboxylase-oxygenase (Rubisco) was used to evaluate the contribution of Rubisco to the control of photosynthetic rate, and the impact of a changed rate of photosynthesis on whole plant composition, allocation and growth. (1) The concept of flux control coefficients is introduced. It is discussed how, with adequate precautions, a set of wild-type and transgenic plants with varying expression of an enzyme can be used to obtain experimental values for its flux control coefficient. (2) The flux control coefficient of Rubisco for photosynthesis depends on the short-term conditions. It increases in high light, or low CO₂. (3) When plants are grown under constant irradiance, the flux control coefficient in the growth conditions is low (<0.2) at irradiances of up to 1000μmol quanta m⁻² s⁻¹. In a natural irradiance regime exceeding 1500μmol quanta m⁻² s⁻² over several hours the flux coefficient rose to 0.8–0.9. It is concluded that plants are able to adjust the balance between Rubisco and the remainder of the photosynthetic machinery, and thereby avoid a one-sided limitation of photosynthesis by Rubisco over a wide range of ambient growth irradiance regimes. (4) When the plants were grown on limiting inorganic nitrogen, Rubisco had a higher flux control coefficient (0.5). It is proposed that, in many growth conditions, part of the investment in Rubisco may be viewed as a nitrogen store, albeit bringing additional marginal advantages with respect to photosynthetic rate and water use efficiency. (5) A change in the rate of photosynthesis did not automatically translate into a change in growth rate. Several factors are identified which contribute to this buffering of growth against a changed photosynthetic rate. (6) There is an alteration in whole plant allocation, resulting in an increase in the leaf area ratio. The increase is mainly due to a higher leaf water content, and not to changes in shoot/root allocation. This increased investment in whole plant leaf area partly counteracts the decreased efficiency of photosynthesis at the biochemical level. (7) Plants with decreased Rubisco have a lower intrinsic water use efficiency and contain high levels of inorganic cations and anions. It is proposed that these are a consequence of the increased rate of transpiration, and that the resulting osmotic potential might be a contributory factor to the increased water content and expansion of the leaves. (8) Starch accumulation in source leaves is decreased when unit leaf photosynthesis is reduced, allowing a more efficient use of the fixed carbon. (9) Decreased availability of carbohydrates leads to a down-regulation of nitrate assimilation, acting via a decrease in nitrate reductase activity.     

Growth response of barley and tomato to nitrogen stress and its control by abscisic acid,water relations and photosynthesis

Barley (Hordeum vulgare L.) and tomato Lycopersicon esculentum Mill.) were grown hydroponically and examined 2, 5, and 10 d after being deprived of nitrogen (N) supply. Leaf elongation rate declined in both species in response to N stress before there was any reduction in rate of dryweight accumulation. Changes in water transport to the shoot could not explain reduced leaf elongation in tomato because leaf water content and water potential were unaffected by N stress at the time leaf elongation began to decline. Tomato maintained its shoot water status in N-stressed plants, despite reduced water absorption per gram root, because the decline in root hydraulic conductance with N stress was matched by a decline in stomatal conductance. In barley the decline in leaf elongation coincided with a small (8%) decline in water content per unit area of young leaves; this decline occurred because root hydraulic conductance was reduced more strongly by N stress than was stomatal conductance. Nitrogen stress caused a rapid decline in tissue NO ₃ ^- pools and in NO ₃ ^- flux to the xylem, particularly in tomato which had smaller tissue NO ₃ ^- reserves. Even in barley, tissue NO ₃ ^- reserves were too small and were mobilized too slowly (60% in 2 d) to support maximal growth for more than a few hours. Organic N mobilized from old leaves provided an additional N source to support continued growth of N-stressed plants. Abscisic acid (ABA) levels increased in leaves of both species within 2 d in response to N stress. Addition of ABA to roots caused an increase in volume of xylem exudate but had no effect upon NO ₃ ^- flux to the xylem. After leaf-elongation rate had been reduced by N stress, photosynthesis declined in both barley and tomato. This decline was associated with increased leaf ABA content, reduced stomatal conductance and a decrease in organic N content. We suggest that N stress reduces growth by several mechanisms operating on different time scales: (1) increased leaf ABA content causing reduced cell-wall extensibility and leaf elongation and (2) a more gradual decline in photosynthesis caused by ABA-induced stomatal closure and by a decrease in leaf organic N.Abbreviation and symbols ABA abscisic acid - c_i leaf internal CO₂ concentration - L_p root hydraulic conductance     

Effects of irradiance on growth,photosynthesis, and water use efficiency of seedlings of the chaparral shrub,Ceanothus megacarpus

Summary The effects of irradiance during growth on biomass allocation, growth rates, leaf chlorophyll and protein contents, and on gas exchange responses to irradiance and CO₂ partial pressures of the evergreen, sclerophyllous, chaparral shrub, Ceanothus megacarpus were determined. Plants were grown at 4 irradiances for the growth experiments, 8, 17, 25, 41 nE cm^-2 sec^-1, and at 2 irradiances, 9 and 50 nE cm^-2 sec^-1, for the other comparisons.At higher irradiances root/shoot ratios were somewhat greater and specific leaf weights were much greater, while leaf area ratios were much lower and leaf weight ratios were slightly lower than at lower irradiances. Relative growth rates increased with increasing irradiance up to 25 nE cm^-2 sec^-1 and then leveled off, while unit leaf area rates increased steeply and unit leaf weight rates increased more gradually up to the highest growth irradiance.Leaves grown at 9 nE cm^-2 sec^-1 had less total chlorophyll per unit leaf area and more per unit leaf weight than those grown at 50 nE cm^-2 sec^-1. In a reverse of what is commonly found, low irradiance grown leaves had significantly higher chlorophyll a/b than high irradiance grown leaves. High irradiance grown leaves had much more total soluble protein per unit leaf area and per unit dry weight, and they had much higher soluble protein/chlorophyll than low irradiance grown leaves.High irradiance grown leaves had higher rates of respiration in very dim light, required higher irradiances for photosynthetic saturation and had higher irradiance saturated rates of photosynthesis than low irradiance grown leaves. CO₂ compensation irradiances for leaves of both treatments were very low, <5 nE cm^-2 sec^-1. Leaves grown under low and those grown under high irradiances reached 95% of their saturated photosynthetic rates at 65 and 85 nE cm^-2 sec^-1, respectively. Irradiance saturated rates of photosynthesis were high compared to other chaparral shrubs, 1.3 for low and 1.9 nmol CO₂ cm^-2 sec^-1 for high irradiance grown leaves. A very unusual finding was that leaf conductances to H₂O were significantly lower in the high irradiance grown leaves than in the low irradiance grown leaves. This, plus the differences in photosynthetic rates, resulted in higher water use efficiencies by the high irradiance grown leaves. High irradiance grown leaves had higher rates of photosynthesis at any particular intercellular CO₂ partial pressure and also responded more steeply to increasing CO₂ partial pressure than did low irradiance grown leaves. Leaves from both treatments showed reduced photosynthetic capability after being subjected to low CO₂ partial pressures (100 bars) under high irradiances. This treatment was more detrimental to leaves grown under low irradiances.The ecological implications of these findings are discussed in terms of chaparral shrub community structure. We suggest that light availability may be an important determinant of chaparral community structure through its effects on water use efficiencies rather than on net carbon gain.     

Effect of irradiance during growth of Glycine max on photosynthetic capacity and percent activation of ribulose 1,5-bisphosphate carboxylase

The initial (in vivo) and total (activity present after preincubation with CO₂ and Mg²⁺) activities of ribulose bisphosphate carboxylase were both assayed in extracts of leaves of soybean (Glycine max) plants which had been grown under 4 different irradiance levels. The total carboxylase activity per unit leaf area decreased with decreased irradiance during growth but was not different on a dry weight basis. The initial activity as a percentage of the total activity was unchanged (approximately 95%) except in leaves of plants grown at the lowest irradiance (74%). When the plants grown at the lowest irradiance were exposed to high irradiance, the initial activity was increased to 93% of the total. Light saturated rates of photosynthesis per unit leaf area were lower and saturated at lower irradiance for plants grown at lower irradiances. Initial carboxylase activity was correlated closely (r²=0.84) with leaf photosynthesis rate on a dry weight basis.     

Influence of temperature and nitrogen concentration on photosynthesis ofDunaliella viridis Teodoresco

The photosynthetic behaviour ofDunaliella viridis has been studied under a combination of three variables: irradiance (0–900 mol m^–2 s^–1), temperature (15, 23, 31, 38, 42 °C) and nitrogen concentration (0.05, 0.5, 1.5, 5, 10 mM NO ₃ ^- ) at a salinity of 2 M NaCl.The highest rates of photosynthesis have been found at 31 °C and a nitrate concentration of 10 mM. There exists a synergistic effect between temperature and nitrogen availability on the photosynthesis ofD. viridis; under nitrogen deficiency oxygen evolution is low, even null at high temperature. The interaction between these two variables of control occurs in a multiplicative way. There is also a general increase in photosynthetic pigments following the increase in nitrogen concentration in the culture medium. The normalization of net photosynthesis data in relation to chlorophylla shows that nitrogen concentration makes an indirect control of the photosynthetic rate ofD. viridis through the variation of pigment concentration.     

Hemp (Cannabis sativa L.) leaf photosynthesis in relation to nitrogen content and temperature: implications for hemp as a bio‐economically sustainable crop

Hemp (Cannabis sativa L.) may be a suitable crop for the bio‐economy as it requires low inputs while producing a high and valuable biomass yield. With the aim of understanding the physiological basis of hemp's high resource‐use efficiency and yield potential, photosynthesis was analysed on leaves exposed to a range of nitrogen and temperature levels. Light‐saturated net photosynthesis rate (A_max) increased with an increase in leaf nitrogen up to 31.2 ± 1.9 μmol m^?2 s^?1 at 25 °C. The A_max initially increased with an increase in leaf temperature (T_L), levelled off at 25–35 °C and decreased when T_L became higher than 35 °C. Based on a C₃ leaf photosynthesis model, we estimated mesophyll conductance (g_m), efficiency of converting incident irradiance into linear electron transport under limiting light (κ_2LL), linear electron transport capacity (J_max), Rubisco carboxylation capacity (V_cmax), triose phosphate utilization capacity (T_p) and day respiration (R_d), using data obtained from gas exchange and chlorophyll fluorescence measurements at different leaf positions and various levels of incident irradiance, CO₂ and O₂. The effects of leaf nitrogen and temperature on photosynthesis parameters were consistent at different leaf positions and among different growth environments except for κ_2LL, which was higher for plants grown in the glasshouse than for those grown outdoors. Model analysis showed that compared with cotton and kenaf, hemp has higher photosynthetic capacity when leaf nitrogen is <2.0 g N m^?2. The high photosynthetic capacity measured in this study, especially at low nitrogen level, provides additional evidence that hemp can be grown as a sustainable bioenergy crop over a wide range of climatic and agronomic conditions.     

Seasonal patterns of photosynthetic response and acclimation to elevated carbon dioxide in field-grown strawberry

Strawberry (Fragaria × ananassa) plants were grown in field plots at the current ambient [CO₂], and at ambient + 300 and ambient + 600 μmol mol⁻¹ [CO₂]. Approximately weekly measurements were made of single leaf gas exchange of upper canopy leaves from early spring through fall of two years, in order to determine the temperature dependence of the stimulation of photosynthesis by elevated [CO₂], whether growth at elevated [CO₂] resulted in acclimation of photosynthesis, and whether any photosynthetic acclimation was reduced when fruiting created additional demand for the products of photosynthesis. Stimulation of photosynthetic CO₂ assimilation by short-term increases in [CO₂] increased strongly with measurement temperature. The stimulation exceeded that predicted from the kinetic characteristics of ribulose-1,5-bisphosphate carboxylase at all temperatures. Acclimation of photosynthesis to growth at elevated [CO₂] was evident from early spring through summer, including the fruiting period in early summer, with lower rates under standard measurement conditions in plants grown at elevated [CO₂]. The degree of acclimation increased with growth [CO₂]. However, there were no significant differences between [CO₂] treatments in total nitrogen per leaf area, and photosynthetic acclimation was reversed one day after switching the [CO₂] treatments. Tests showed that acclimation did not result from a limitation of photosynthesis by triose phosphate utilization rate at elevated [CO₂]. Photosynthetic acclimation was not evident during dry periods in midsummer, when the elevated [CO₂] treatments conserved soil water and photosynthesis declined more at ambient than at elevated [CO₂]. Acclimation was also not evident during the fall, when plants were vegetative, despite wet conditions and continued higher leaf starch content at elevated [CO₂]. Stomatal conductance responded little to short-term changes in [CO₂] except during drought, and changed in parallel with photosynthetic acclimation through the seasons in response to the long-term [CO₂] treatments. The data do not support the hypothesis that source-sink balance controls the seasonal occurrence of photosynthetic acclimation to elevated [CO₂] in this species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Acclimation of photosynthesis in a boreal grass (Phalaris arundinacea L.) under different temperature, CO2, and soil water regimes

The aim of this work was to study the acclimation of photosynthesis in a boreal grass (Phalaris arundinacea L.) grown in controlled environment chambers under elevated temperature (ambient + 3.5°C) and CO₂ (700 μmol mol⁻¹) with varying soil water regimes. More specifically, we studied, during two development stages (early: heading; late: florescence completed), how the temperature response of light-saturated net photosynthetic rate (P _sat), maximum rate of ribulose-1,5-bisphosphate carboxylase/oxygenase activity (V _cmax) and potential rate of electron transport (J _max) acclimatized to the changed environment. During the early growing period, we found a greater temperature-induced enhancement of P _sat at higher measurement temperatures, which disappeared during the late stage. Under elevated growth temperature, V _cmax and J _max at lower measurement temperatures (5–15°C) were lower than those under ambient growth temperature during the early period. When the measurements were done at 20–30°C, the situation was the opposite. During the late growing period, V _cmax and J _max under elevated growth temperature were consistently lower across measurement temperatures. CO₂ enrichment significantly increased P _sat with higher intercellular CO₂ compared to ambient CO₂ treatment, however, elevated CO₂ slightly decreased V _cmax and J _max across measurement temperatures, probably due to down-regulation acclimation. For two growing periods, soil water availability affected the variation in photosynthesis and biochemical parameters much more than climatic treatment did. Over two growing periods, V _cmax and J _max were on average 36.4 and 30.6%, respectively, lower with low water availability compared to high water availability across measurement temperatures. During the late growing period, elevated growth temperature further reduced the photosynthesis under low water availability. V _cmax and J _max declined along with the decrease in nitrogen content of leaves as growing period progressed, regardless of climatic treatment and water regime. We suggest that, for grass species, seasonal acclimation of the photosynthetic parameters under varying environmental conditions needed to be identified to fairly estimate the whole-life photosynthesis.     

Thermal Acclimation of Respiration and Photosynthesis in the Marine Macroalga Gracilaria lemaneiformis (Gracilariales,Rhodophyta)

The responses of respiration and photosynthesis to temperature fluctuations in marine macroalgae have the potential to significantly affect coastal carbon fluxes and sequestration. In this study, the marine red macroalga Gracilaria lemaneiformis was cultured at three different temperatures (12, 19, and 26°C) and at high‐ and low‐nitrogen (N) availability, to investigate the acclimation potential of respiration and photosynthesis to temperature change. Measurements of respiratory and photosynthetic rates were made at five temperatures (7°C–33°C). An instantaneous change in temperature resulted in a change in the rates of respiration and photosynthesis, and the temperature sensitivities (i.e., the Q₁₀ value) for both the metabolic processes were lower in 26°C‐grown algae than 12°C‐ or 19°C‐grown algae. Both respiration and photosynthesis acclimated to long‐term changes in temperature, irrespective of the N availability under which the algae were grown; respiration displayed strong acclimation, whereas photosynthesis only exhibited a partial acclimation response to changing growth temperatures. The ratio of respiration to gross photosynthesis was higher in 12°C‐grown algae, but displayed little difference between the algae grown at 19°C and 26°C. We propose that it is unlikely that respiration in G. lemaneiformis would increase significantly with global warming, although photosynthesis would increase at moderately elevated temperatures.     

Temperature response of photosynthetic light‐ and carbon‐use characteristics in the red seaweed Gracilariopsis lemaneiformis (Gracilariales,Rhodophyta)

The red seaweed Gracilariopsis is an important crop extensively cultivated in China for high‐quality raw agar. In the cultivation site at Nanao Island, Shantou, China, G. lemaneiformis experiences high variability in environmental conditions like seawater temperature. In this study, G. lemaneiformis was cultured at 12, 19, or 26°C for 3 weeks, to examine its photosynthetic acclimation to changing temperature. Growth rates were highest in G. lemaneiformis thalli grown at 19°C, and were reduced with either decreased or increased temperature. The irradiance‐saturated rate of photosynthesis (P_max) decreased with decreasing temperature, but increased significantly with prolonged cultivation at lower temperatures, indicating the potential for photosynthesis acclimation to lower temperature. Moreover, P_max increased with increasing temperature (~30 μmol O₂ · g^?1FW · h^?1 at 12°C to 70 μmol O₂ · g^?1FW · h^?1 at 26°C). The irradiance compensation point for photosynthesis (I_c) decreased significantly with increasing temperature (28 μmol photons · m^?2 · s^?1 at high temperature vs. 38 μmol photons · m^?2 · s^?1 at low temperature). Both the photosynthetic light‐ and carbon‐use efficiencies increased with increasing growth or temperatures (from 12°C to 26°C). The results suggested that the thermal acclimation of photosynthetic performance of G. lemaneiformis would have important ecophysiological implications in sea cultivation for improving photosynthesis at low temperature and maintaining high standing biomass during summer. Ongoing climate change (increasing atmospheric CO₂ and global warming) may enhance biomass production in G. lemaneiformis mariculture through the improved photosynthetic performances in response to increasing temperature.     

Responses of wild plants to nitrate availability

Summary Two annual species of Bromus, an invader (B. hordeaceus, ex B. mollis) and a non-invader (B. intermedius), were grown for 28 days in growth chambers, at 5 and 100 M NO ₃ ^- in flowing nutrient solution. No differences between the two species were observed at either NO ₃ ^- level, in terms of relative growth rate (RGR) or its components, dry matter partitioning, specific NO ₃ ^- absorption rate, nitrogen concentration, and other characteristics of NO ₃ ^- uptake and photosynthesis. The effects of decreasing NO ₃ ^- concentration in the solution were mainly to decrease the NO ₃ ^- concentration in the plants through decreased absorption rate, and to decrease the leaf area ratio through increased specific leaf mass and decreased leaf mass ratio. Organic nitrogen concentration varied little between the two treatments, which may be the reason why photosynthetic rates were not altered. Consequently, RGR was only slightly decreased in the 5-M treatment compared to the 100-M treatment. This is in contrast with other species, where growth is reduced at much higher NO ₃ ^- concentrations. These discrepancies may be related to differences in RGR, since a log-linear relationship was found between RGR and the NO ₃ ^- concentration at which growth is first reduced. In addition, a strong linear relationship was found between the RGR of these species and their maximum absorption rate for nitrate, suggesting that the growth of species with low maximum RGR may be partly regulated by nutrient uptake.     

Acclimation of photosynthesis to irradiance and spectral quality in British plant species: chlorophyll content, photosynthetic capacity and habitat preference

Twenty-two common British angiosperms were examined for their ability to acclimate photosynthetically to sun and shade conditions. Plants were grown under low irradiance, far-red enriched light (50 μmol m^?2 s^?1), selected to mimic as closely as possible natural canopy shade, and moderately high light of insufficient irradiance to induce photoinhibitory or photoprotective responses (300 μmol m^?2 s^?1). Light-and CO²-saturated photosynthetic rates of oxygen evolution (P_max) and chlorophyll content were measured. Large variation was found in both parameters, and two ‘strategies’ for long-term acclimation were identified: firstly a change in chlorophyll per unit leaf area which was found to correlate positively with photosynthetic capacity, and secondly changes in chlorophyll alb ratio and P_max, indicative of alterations at the chloroplast level, which were not associated with a change in chlorophyll content per unit leaf area. Combinations of these two strategies may occur, giving rise to the observed diversity in photosynthetic acclimation. The extent and nature of photosynthetic acclimation were compared with an index of shade association, calculated from the association each species has with woodland. It was found that the greatest flexibility for change at the chloroplast level was found in those species possessing an intermediate shade association, whilst acclimation in ‘sun’ species proceeded by a change in chlorophyll content; obligate shade species showed little capacity for acclimation at either the chloroplast or leaf level. A framework for explaining the variation between plant species in leaf-level photosynthetic capacity, in relation to the natural light environment, is presented. This is the first time the potential for light acclimation of photosynthesis in different plant species has been satisfactorily linked to habitat distribution.     

Relative enhancement of photosynthesis and growth at elevated CO2 is greater under sunflecks than uniform irradiance in a tropical rain forest tree seedling

The survivorship of dipterocarp seedlings in the deeply shaded understorey of South‐east Asian rain forests is limited by their ability to maintain a positive carbon balance. Photosynthesis during sunflecks is an important component of carbon gain. To investigate the effect of elevated CO₂ upon photosynthesis and growth under sunflecks, seedlings of Shorealeprosula were grown in controlled environment conditions at ambient or elevated CO₂. Equal total daily photon flux density (PFD) (～7·7 mol m^?2 d^?1) was supplied as either uniform irradiance (～170 µmol m^?2 s^?1) or shade/fleck sequences (～30 µmol m^?2 s^?1/～525 µmol m^?2 s^?1). Photosynthesis and growth were enhanced by elevated CO₂ treatments but lower under flecked irradiance treatments. Acclimation of photosynthetic capacity occurred in response to elevated CO₂ but not flecked irradiance. Importantly, the relative enhancement effects of elevated CO₂ were greater under sunflecks (growth 60%, carbon gain 89%) compared with uniform irradiance (growth 25%, carbon gain 59%). This was driven by two factors: (1) greater efficiency of dynamic photosynthesis (photosynthetic induction gain and loss, post‐irradiance gas exchange); and (2) photosynthetic enhancement being greatest at very low PFD. This allowed improved carbon gain during both clusters of lightflecks (73%) and intervening periods of deep shade (99%). The relatively greater enhancement of growth and photosynthesis at elevated CO₂ under sunflecks has important potential consequences for seedling regeneration processes and hence forest structure and composition.     

THE EFFECT OF GROWTH IRRADIANCE ON THE COUPLING OF CARBON AND NITROGEN METABOLISM IN CHAETOMORPHA LINUM (CHLOROPHYTA)

The influence of growth irradiance on the non-steady-state relationship between photosynthesis and tissue carbon (C) and nitrogen (N) pools in Chaetomorpha linum (Muller) Kutzing in response to abrupt changes in external nitrogen (N) availability was determined in laboratory experiments. For a given thallus N content, algae acclimated to low irradiance consistently had a higher rate of light-saturated photosynthesis (P_max normalized to dry weight) than algae acclimated to saturating irradiance; for both treatments, P_max was correlated to thallus N. Both P_max and the photosynthetic efficiency (α_dw) were correlated in C. linum grown at either saturating or limiting irradiance over the range of experimental conditions, indicating that variations in electron transport were coupled to variations in C-fixation capacity despite the large range of tissue N content from 1.1% to 4.8%. Optimizing both α and P_max and thereby acclimating to an intermediate light level may be a general characteristic of thin-structured opportunistic algae that confers a competitive advantage in estuarine environments in which both light and nutrient conditions are highly variable. Nitrogen-saturated algae had the same photosynthesis–irradiance relationship regardless of light level. When deprived of an external N supply, photosynthetic rates did not change in C. linum acclimated to low irradiance despite a two-fold decrease in tissue N content, suggesting that the active pools of chlorophyll and Rubisco remained constant. Both α and P_max decreased immediately and continuously in algae acclimated to high irradiance on removal of the N supply even though tissue N content was relatively high during most of the N-starvation period, indicating a diversion of energy and reductant away from C fixation to support high growth rates. Carbon and nitrogen assimilation were equally balanced in algae in both light treatments throughout the N-saturation and -depletion phases, except when protein synthesis was limited by the depletion of internal N reserves in severely N-starved high-light algae and excess C accumulated as starch stores. This suggests that the ability for short-term adjustment of internal allocation to acquire N andC in almost constant proportions may be especially beneficial to macroalgae living in environments characterized by high variability in light levels and nutrient supply.     

Interactive effect of salt (NaCl) and nitrogen form on growth, water relations and photosynthetic capacity of sunflower (Helianthus annum L.)

The effects of the ammonium (NH₄⁺) and nitrate (NO₃^-) forms of nitrogen and NaCl on the growth, water relations and photosynthesis performance of sunflower (Helianthus annuus L.) were examined under glasshouse conditions. Eight-day-old plants of cv. Hisun 33 were subjected for 21 days to Hoagland's nutrient solution containing 8 mol m^-3N as NH₄⁺or NO₃^-, and salinised with 0, 60, or 120 mol m^-3NaCl. Fresh weights of shoots and roots, and leaf area of NO₃^-supplied non-salinised plants were significantly greater than those of NH₄⁺-supplied non-salinised plants. But addition of NaCl to the rooting medium of these plants had more inhibitory effect on the growth of NO₃^--supplied plants than on NH₄⁺-supplied plants. Both leaf water and osmotic potentials of plants grown with NH₄⁺were lower than those of plants given NO₃^-under both non-saline and saline conditions. Chlorophylls a and b concentrations were higher in plants grown with NH₄⁺than N0₃^--supplied plants at the lower two levels of salinisation. The rate of photosynthesis in plants was considerably higher in non-salinised plants grown with NO₃^-than with NH₄⁺, but with increase in salinisation the photosynthesis rate decreased in NO₃^--supplied plants, but not in those given NH₄⁺. The rate of transpiration was increased significantly by salinisation in NO₃^--supplied plants, but not consistently so in NH₄⁺-supplied plants. The stomatal conductances were much higher in plants given NO₃^-than with NH₄⁺when grown under non-saline conditions, but not when salinised. As a consequence, water-use efficiency in NO₃^--supplied control plants was better than in NH₄⁺-supplied under non-saline conditions, but worse under saline conditions. The different forms of nitrogen and the addition of NaCl to the growing medium did not affect the relative intercellular concentrations of CO₂ (C_i/C_a). Overall, the NH₄⁺form of nitrogen inhibited the growth of sunflowers under non-saline conditions, but NO₃^-and NaCl interacted to inhibit growth more than did NH₄⁺under saline conditions.