Comparative effects of light during growth on the photosynthetic properties of NADP-ME type C4 grasses from open and shaded habitats. II. Photosynthetic enzyme activities and metabolism*

Abstract The influences of shading during growth upon the activities of several photosynthetic enzymes were examined in NADP-ME type C₄ grasses from open (Zea mays L.) and shaded (Paspalum conjugation Berg.) habitats. The substantial species-difference in maximum photosynthetic rate observed under a high light regime was correlated with large differences in both enzyme activities and leaf protein contents. With the exception of RuBP carboxylase activity, other photosynthetic enzyme activities in Z. mays were reduced by shading to a similar extent as maximum photosynthetic rate. In contrast, only PEP carboxylase and pyruvate, P_i dikinase activities were decreased by shading in P. conjugatum. As with maximum photosynthetic rate, other photosynthetic enzyme activities in P. conjugatum were relatively insensitive to irradiance during growth. Under a low photon flux density of photosynthetically active radiation (50 μmol m^?2 s^?1), the flow of [¹⁴C] label through photosynthetic intermediates in intact, shade-grown leaves of P. conjugatum was typical of C₄ metabolism. This provides incontrovertible proof for the occurrence of C₄ photosynthesis in shaded habitats.     

Influence of phosphorus and nitrogen on photosynthetic parameters and growth in Vicia faba L.

The influence of phosphorus (P) and nitrogen (N) supply on biomass, leaf area, photon saturated photosynthetic rate (P_max), quantum yield efficiency (), intercellular CO₂ concentration (C_i), and carboxylation efficiency (CE) was investigated in Vicia faba. The influence of P on N accumulation, biomass, and leaf area production was also investigated. An increase in P supply was consistently associated with an increase in N accumulation and N productivity in terms of biomass and leaf area production. Furthermore, P increased the photosynthetic N use efficiency (NUE) in terms of P_max and . An increase in P supply was also associated with an increase in CE and a decrease in C_i. Under variable daily meteorological conditions specific leaf nitrogen content (N_L), specific leaf phosphorus content (P_L), specific leaf area (_L), root mass fraction (R_f), P_max, and remained constant for a given N and P supply. A monotonic decline in the steady-state value of R_f occurred with increasing N supply. _L increased with increasing N supply or with increasing N_L. We tested also the hypothesis that P supply positively affects both N demand and photosynthetic NUE by influencing the upper limit of the asymptotic values for P_max and CE, and the lower limit for C_i in response to increasing N.This revised version was published online in March 2005 with corrections to the page numbers.     

干旱和遮荫对马尾松幼苗生长和光合特性的影响

为探究马尾松对干旱和遮荫胁迫的生理响应规律和适应机制,以2年生马尾松幼苗为对象,设置对照(CK)、模拟干旱(DR)、遮荫(LL)以及干旱与遮荫的交互处理(DRLL)4种环境,研究干旱和遮荫对马尾松幼苗的生长和光合生理特性的影响。结果表明:(1)在干旱、遮荫和二者的交互处理下,马尾松幼苗的基径和株高增长量均显著减小,且二者的交互处理加重了干旱和遮荫单一处理下的减小趋势,二者交互作用的影响符合"相互作用理论"。(2)在干旱处理下,针叶长度和比叶面积减小,净光合速率、蒸腾速率、气孔导度和细胞间隙CO₂浓度显著降低,水分利用效率显著增加,光合色素含量基本不变。(3)在遮荫处理下,针叶长度和比叶面积增大,净光合速率、蒸腾速率、气孔导度显著降低,光合色素含量显著增加,水分利用效率和细胞间隙CO₂浓度基本未发生改变。(4)二者的交互处理下,针叶长度、比叶面积减小,净光合速率、蒸腾速率和气孔导度显著降低,且降低程度大于干旱单一处理,光合色素含量显著增加,但其增加程度小于遮荫单一处理。说明干旱和遮荫均能抑制马尾松植株的生长,但其光合生理特性在干旱和遮荫胁迫下分...     

遮光对香果树幼苗光合特性及叶片解剖结构的影响

设置了全光照(L0)、透光率60%(L1)、透光率25%(L2)、透光率10%(L3)4种光环境,研究了不同遮光处理对香果树幼苗光合作用及叶片解剖结构的影响。结果表明:(1)遮光对香果树幼苗净光合速率(P_n)、蒸腾速率(T_r)、气孔导度(G_s)、气孔限制值(L_s)、胞间CO_2浓度(C_i)、蒸汽压亏缺(V_(pdl))均产生了显著影响。全光和L1处理下Pn和Tr日变化呈明显的双峰曲线,中午出现了明显的"午休",其余处理的变化相对缓和,双峰和"午休"现象不明显;G_s的日变化规律与Pn基本一致;C_i的日变化整体呈先期缓慢降低后期抬升的趋势,与对照相比遮光显著提高了叶片胞间CO_2浓度;Ls和V_(pdl)呈现出先升后降的弧形变化,并且遮光下L_s和V_(pdl)日均值显著低于全光照。一天中P_n和T_r的日均值大小排序为L1L0L2L3。(2)不同光环境下香果树幼苗的光合响应特征明显不同。全光下,香果树幼苗能够维持较高的净光合作用速率而未呈现明显的光抑制,表现出一定的光忍耐和喜光性。L1处理下的最大净光合速率(P_(max))、暗呼吸速率(R_d)和表观量子效率(AQY)显著增加,说明香果树对光强具有一定的可塑性,轻度遮光有利于光合效率提高;强度遮光(L2、L3)导致光补偿点(LCP)和光饱和点(LSP)显著降低,有利于充分利用弱光,但P_(max)、R_d明显下降,光合效率受到抑制。(3)遮光显著影响了香果树叶片的气孔分布。遮光条件下香果树叶片的气孔密度显著减小,气孔器长度、宽度与单个气孔器面积也呈现类似的变化。(4)遮光条件下香果树叶片厚度减小,上表皮增厚,下表/ST增加;维管束内单列导管数目和导管直径均有明显增大,促进了叶片水分传导运输。(5)在育苗生产中,为了适应香果树幼苗生长发育的需求,建议有效辐射强度保持在自然光强的60%以上。     

Influence of phosphorus and nitrogen on photosynthetic parameters and growth in <Emphasis Type="Italic">Vicia faba</Emphasis> L.

The influence of phosphorus (P) and nitrogen (N) supply on biomass, leaf area, photon saturated photosynthetic rate (P_max), quantum yield efficiency (α), intercellular CO₂ concentration (C_i), and carboxylation efficiency (CE) was investigated in Vicia faba. The influence of P on N accumulation, biomass, and leaf area production was also investigated. An increase in P supply was consistently associated with an increase in N accumulation and N productivity in terms of biomass and leaf area production. Furthermore, P increased the photosynthetic N use efficiency (NUE) in terms of P_max and α. An increase in P supply was also associated with an increase in CE and a decrease in C_i. Under variable daily meteorological conditions specific leaf nitrogen content (N_L), specific leaf phosphorus content (P_L), specific leaf area (δ_L), root mass fraction (R_f), P_max, and α remained constant for a given N and P supply. A monotonic decline in the steady-state value of R_f occurred with increasing N supply. δ_L increased with increasing N supply or with increasing N_L. We tested also the hypothesis that P supply positively affects both N demand and photosynthetic NUE by influencing the upper limit of the asymptotic values for P_max and CE, and the lower limit for C_i in response to increasing N.     

Highly purified intact chloroplasts from mesophyll protoplasts of the C4 plant Digitaria sanguinalis. Inhibition of phosphoglycerate reduction by orthophosphate and by phosphoenolpyruvate

Purified mesophyll protoplasts from the C₄ plant Digitaria sanguinalis were used to prepare intact mesophyll chloroplasts with low cytoplasmic contamination. The procedure involved breakage of protoplasts, differential centrifugation, partition in a dextran-polyethylene glycol two-phase system, and Percoll density gradient centrifugation. The final chloroplast preparation contained about 80% intact chloroplasts with a phosphoenolpyruvate carboxylase contamination of 0.2–1% of the original protoplast activity, corresponding to 1–6 μmol ¹⁴CO₂ fixed/mg Chl h. The purified chloroplasts showed substrate-dependent oxygen evolution in the range of 40–150 μmol substrate reduced/mg Chl h, with phosphoglycerate or oxaloacetate as substrate. Both reactions were stimulated 1.5 fold by pyruvate and further by addition of the other substrate. These measurements indicated that phosphoglycerate reduction was limited by substrate transport across the chloroplast envelope. Without added substrate, the chloroplasts consumed oxygen via pseudo-cyclic electron transport in the light. Also this reaction was stimulated by pyruvate. Phosphoglycerate-dependent oxygen evolution was inhibited by P_i and by phosphoenolpyruvate to about the same extent with purified chloroplasts, but only by P_i with protoplast extracts. This suggests that phosphoglycerate, P_i and phosphoenolpyruvate share a common carrier, similar to the P_i-translocator in C₃ chloroplasts, and that the lack of inhibition obtained with phosphoenolpyruvate and unpurified chloroplasts is artefactual, possibly due to oxaloacetate formation from added phosphoenolpyruvate and concomitant stimulation of oxygen evolution by oxaloacetate reduction. Furthermore, the results suggest that phosphoenolpyruvate is transported with a K_m similar to that of P_i in C₄ mesophyll chloroplasts.     

Does the photosynthetic light‐acclimation need change in leaf anatomy?

There is a strong correlation between leaf thickness and the light‐saturated rate of photosynthesis per unit leaf area (P_max). However, when leaves are exposed to higher light intensities after maturation, P_max often increases without increasing leaf thickness. To elucidate the mechanism with which mature leaves increase P_max, the change in anatomical and physiological characteristics of mature leaves of Chenopodium album, which was transferred from low to high light condition, were examined. When compared with leaves subjected to low light continuously (LL leaves), the leaves transferred from low to high light (LH leaves) significantly increased P_max. The transfer also increased the area of chloroplasts facing the intercellular space (S_c) and maintained a strong correlation between P_max and S_c. The mesophyll cells of LL leaves had open spaces along cell walls where chloroplasts were absent, which enabled the leaves to increase P_max when they were exposed to high light (LH). However, the LH leaves were not thick enough to allow further increase in P_max to the level in HH leaves. Thus leaf thickness determines an upper limit of P_max of leaves subjected to a change from low to high light conditions. Shade leaves would only increase P_max when they have open space to accommodate chloroplasts which elongate after light conditions improve.     

Influence of Phosphorus and Endomycorrhiza (Glomus Intraradices) on gas exchange and plant growth of Chile Ancho Pepper (Capsicum Annuum L. cv. San Luis)

Seedlings of chile ancho pepper were grown in pots containing a pasteurized mixture of sand and a low phosphorus (P) sandy loam soil, and either inoculated (VAM) or not inoculated (NVAM) with the endomycorrhizal fungus Glomus intraradices. Long Ashton nutrient solution (LANS) was modified to supply P to the seedlings at 0, 11, and 44 g(P) m^-3 (P₀, P11, P₄₄, respectively). Low P depressed net photosynthetic rate (P_N), stomatal conductance (gs), phosphorus use efficiency (P_N/P), and internal CO₂ concentration (C_i). The mycorrhiza alleviated low P effects by increasing P_N, g_s, P_N/P, and decreasing C_i. At P₀, C_i of NVAM plants was equal to or higher than that of VAM plants, suggesting nonstomatal inhibition of photosynthesis. Gas exchange of VAM plants at P₀ was similar to that of NVAM plants at P11. Endomycorrhiza increased leaf number, leaf area, shoot, root and fruit mass at P₀ and P₁₁ compared to NVAM plants. Reproductive growth was enhanced by 450 % in mycorrhizal plants at P₄₄. Root colonization (arbuscules, vesicles, internal and extraradical hyphae development) was higher at lower P concentrations, while sporulation was unaffected. The enhanced growth and gas exchange of mycorrhizal plants was in part due to greater uptake of P and greater extraradical hyphae development. This revised version was published online in June 2006 with corrections to the Cover Date.     

Intracellular localization of certain photosynthetic enzymes in bundle sheath cells of plants possessing the C4 pathway of photosynthesis.

Bundle sheath cells were enzymatically isolated from representatives of three groups of C₄ plants: Zea mays (NADP malic enzyme type), Panicum miliaceum (NAD malic enzyme type), and Panicum maximum (phosphoenolpyruvate (PEP) carboxykinase type). Cellular organelles from bundle sheath homogenates were partially resolved by differential centrifugation and on isopycnic sucrose density gradients in order to study compartmentation of photosynthetic enzymes. A 48-h-dark pretreatment of the leaves allowed the isolation of relatively intact chloroplasts. Enzymes that decarboxylate C₄ acids and furnish CO₂ to the Calvin cycle are localized as follows: NADP malic enzyme, chloroplastic in Z. mays; NAD malic enzyme, mitochondrial in all three species; PEP carboxykinase, chloroplastic in P. maximum. The activity of NAD malic enzyme in the three species was in the order of P. miliaceum > P. maximum > Z. mays. There were high levels of aspartate and alanine aminotransferases in bundle sheath extracts of P. miliaceum and P. maximum and substantial activity in Z. mays. In all three species, aspartate aminotransferase was mitochondrial whereas alanine aminotransferase was cytoplasmic. Based on the activity and localization of certain enzymes, the concept for aspartate and malate as transport metabolites from mesophyll to bundle sheath cells in C₄ species of the three C₄ groups is discussed.     

Functional consequences of stenophylly for leaf productivity: comparison of the anatomy and physiology of a rheophyte, Farfugium japonicum var. luchuence, and a related non-rheophyte, F. japonicum (Asteraceae)

We investigated the anatomical and physiological characteristics of stenophyllous leaves of a rheophyte, Farfugium japonicum var. luchuence, and sun and shade leaves of a non-rheophyte, F. japonicum, comparing three different populations from coastal, forest floor, and riparian habitats. Light adaptation resulted in smaller leaves, and riparian adaptation resulted in narrower leaves (stenophylly). The light-saturated rate of photosynthesis (P _max) per unit leaf area corresponded to the light availability of the habitat. Irrespective of leaf size, the P _max per unit leaf mass was similar for sun and shade leaves. However, the P _max per mass of stenophyllous leaves was significantly lower than that of sun and shade leaves. This was because the number and size of mesophyll cells were greater than that required for intercellular CO₂ diffusion, which resulted in a larger leaf mass per unit leaf area. Higher cell density increases contact between mesophyll cells and enhances leaf toughness. Stenophyllous leaves of the rheophyte are frequently exposed to a strong water flow when the water level rises, suggesting a mechanical constraint caused by physical stress.     

Gas exchanges of three co-occurring species of Cypripedium in a scrubland in the Hengduan Mountains

Gas exchanges and related leaf traits of three co-occurring species of genus Cypripedium (C. yunnanense Franch., C. guttatum SW., and C. flavum P.F. Hunt et Summerch.) were investigated in a scrubland at 3 460 m a.s.l. in the Hengduan Mountains. The considered species had similar photosynthetic responses to photosynthetic photon flux density (PPFD) and air temperature. The photosynthetic capacity (P _max), carboxylation efficiency (CE), apparent quantum efficiency (AQE), PPFD-saturated rate of electron transport (J_max), respiration rate (R _D), and leaf nitrogen content per unit area (LNC) of C. guttatum were higher than those of C. yunnanense and C. flavum. The highest P _max of C. guttatum was related to the highest LNC and the lowest ratio of intercellular CO₂ concentration to atmospheric CO₂ concentration (C _i/C _a). However, no significant differences in stomatal conductance (g _s) and relative stomatal limitations (RSL) were observed among the three species. Hence biochemical limitation had a dominant role in P _max differences among the considered species.     

Phosphorus recycling in photorespiration maintains high photosynthetic capacity in woody species

Leaf photosynthetic CO₂ responses can provide insight into how major nutrients, such as phosphorus (P), constrain leaf CO₂ assimilation rates (A_net). However, triose‐phosphate limitations are rarely employed in the classic photosynthesis model and it is uncertain as to what extent these limitations occur in field situations. In contrast to predictions from biochemical theory of photosynthesis, we found consistent evidence in the field of lower A_net in high [CO₂] and low [O₂] than at ambient [O₂]. For 10 species of trees and shrubs across a range of soil P availability in Australia, none of them showed a positive response of A_net at saturating [CO₂] (i.e. A_max) to 2 kPa O₂. Three species showed >20% reductions in A_max in low [O₂], a phenomenon potentially explained by orthophosphate (P_i) savings during photorespiration. These species, with largest photosynthetic capacity and P_i > 2 mmol P m^?2, rely the most on additional P_i made available from photorespiration rather than species growing in P‐impoverished soils. The results suggest that rarely used adjustments to a biochemical photosynthesis model are useful for predicting A_max and give insight into the biochemical limitations of photosynthesis rates at a range of leaf P concentrations. Phosphate limitations to photosynthetic capacity are likely more common in the field than previously considered.     

Canopy gradients in leaf intercellular CO2 mole fractions revisited: interactions between leaf irradiance and water stress need consideration

Intercellular CO₂ mole fractions (C_i) are lower in the upper canopy relative to the lower canopy leaves. This canopy gradient in C_i has been associated with enhanced rates of carbon assimilation at high light, and concomitant greater draw‐downs in C_i. However, increases in irradiance in the canopy are generally also associated with decreases in leaf water availability. Thus, stress effects on photosynthesis rates (A) and stomatal conductance (G), may provide a further explanation for the observed C_i gradients. To test the hypotheses of the sources of canopy variation in C_i, and quantitatively assess the influence of within‐canopy differences in stomatal regulation on A, the seasonal and diurnal variation in G was studied in relation to seasonal average daily integrated quantum flux density (Q_int) in tall shade‐intolerant Populus tremula L. trees. Daily time‐courses of A were simulated using the photosynthesis model of Farquhar et al. (Planta 149, 78–90, 1980). Stable carbon isotope composition of a leaf carbon fraction with rapid turnover rate was used to estimate canopy gradient in C_i during the simulations. Daily maximum G (G_max) consistently increased with increasing Q_int. However, canopy differences in G_max decreased as soil water availability became limiting during the season. In water‐stressed leaves, there were strong mid‐day decreases in G that were poorly associated with vapour pressure deficits between the leaf and atmosphere, and the magnitude of the mid‐day decreases in G occasionally interacted with long‐term leaf light environment. Simulations indicated that the percentage of carbon lost due to mid‐day stomatal closure was of the order of 5–10%, and seasonal water stress increased this percentage up to 20%. The percentage of carbon lost due to stomatal closure increased with increasing Q_int. Canopy differences in light environment resulted in a gradient of daily average C_i of approximately 20 µmol mol^?1. The canopy variation in seasonal and diurnal reductions in G led to a C_i gradient of approximately 100 µmol mol^?1, and the actual canopy C_i gradient was of the same magnitude according to leaf carbon isotope composition. This study demonstrates that stress effects influence C_i more strongly than within‐canopy light gradients, and also that leaves acclimated to different irradiance and water stress conditions may regulate water use largely independent of foliar photosynthetic potentials.     

Photosynthetic performance of <Emphasis Type="Italic">Lycoris radiata</Emphasis> var. <Emphasis Type="Italic">radiata</Emphasis> to shade treatments

The effects of shade on the growth, leaf photosynthetic characteristics, and chlorophyll (Chl) fluorescence parameters of Lycoris radiata var. radiata were determined under differing irradiances (15, 65, and 100% of full irradiance) within pots. The HI plants exhibited a typical decline in net photosynthetic rate (P _N) during midday, which was not observed in MI- and LI plants. This indicated a possible photoinhibition in HI plants as the ratio of variable to maximum fluorescence (F_v/F_m) value was higher and the minimal fluorescence (F₀) was lower in the, and LI plants. Diurnal patterns of stomatal conductance (g _s) and transpiration rate (E) were remarkably similar to those of P _N at each shade treatments, and the intercellular CO₂ concentration (C _i) had the opposite change trend. Under both shading conditions, the light saturation point, light compensation point and photon-saturated photosynthetic rate (P _max) became lower than those under full sunlight, and it was the opposite for the apparent quantum yield (AQY). The higher the level of shade, the lower the integrated daytime carbon gain, stomatal and epidermis cell densities, specific leaf mass (SLM), bulb mass ratio (BMR), leaf thickness, and Chl a/b ratio. In contrast, contents of Chls per dry mass (DM), leaf area ratio (LAR), leaf mass ratio (LMR), leaf length, leaf area and total leaf area per plant increased under the same shade levels to promote photon absorption and to compensate for the lower radiant energy. Therefore, when the integrated daytime carbon gain, leaf area and total leaf area per plant, which are the main factors determining the productivity of L. radiata var. radiata plant, were taken into account together, this species may be cultivated at about 60∼70% of ambient irradiance to promote its growth.     

The effect of growth irradiance on leaf anatomy and photosynthesis in Acer species differing in light demand

Variation in light demand is a major factor in determining the growth and survival of trees in a forest. There is strong relation between the light‐demand and the effect of growth irradiance on leaf morphology and photosynthesis in three Acer species: A. rufinerve (light‐demanding), A. mono (intermediate) and A. palmatum (shade‐tolerant). The increase in mesophyll thickness and surface area of chloroplasts facing the intercellular airspaces (S_c) with growth irradiance was highest in A. rufinerve. Although the increase in light‐saturated photosynthesis (A_max) was similar among the species, the increase in water use efficiency (WUE) was much higher in A. rufinerve than that in the other species, indicating that the response to water limitation plays an important role in leaf photosynthetic acclimation to high light in A. rufinerve. The low CO₂ partial pressure at the carboxylation site (C_c) in A. rufinerve (130 µmol mol^?1) at high irradiance was caused by low stomatal and internal conductance to CO₂ diffusion, which minimized the increase in A_max in A. rufinerve despite its high Rubisco content. Under shade conditions, interspecific differences in leaf features were relatively small. Thus, difference in light demand related to leaf acclimation to high light rather than that to low light in the Acer species.     

Low conductances for CO2 diffusion from stomata to the sites of carboxylation in leaves of woody species

Concurrent measurements of leaf gas exchange and on-line ¹³C discrimination were used to evaluate the CO₂ conductance to diffusion from the stomatal cavity to the sites of carboxylation within the chloroplast (internal conductance; g_i). When photon irradiance was varied it appeared that g_i and/or the discrimination accompanying carboxylation also varied. Despite this problem, g_i, was estimated for leaves of peach (Prunus persica), grapefruit (Citrus paradisi), lemon (C. limon) and macadamia (Macadamia integrifolia) at saturating photon irradiance. Estimates for leaves of C. paradisi, C. limon and M. integrifolia were considerably lower than those previously reported for well-nourished herbaceous plants and ranged from 1.1 to2.2μmol CO₂ m^?2 s^?1 Pa^?1, whilst P. persica had a mean value of 3.5 μmol CO₂ m^?2 s^?1 Pa^?1. At an ambient CO₂ partial pressure of 33Pa, estimates of chloroplastic partial pressure of CO₂ (C_c) using measurements of CO₂ assimilation rate (A) and calculated values of g_i, and of partial pressure of CO₂ in the stomatal cavity (C_st) were as low as 11.2 Pa for C. limon and as high as 17.8Pa for peach. In vivo maximum rubisco activities (V_max) were also determined from estimates of C_c. This calculation showed that for a given leaf nitrogen concentration (area basis) C. paradisi and C. limon leaves had a lower V_max than P. persica, with C. paradisi and C. limon estimated to have only 10% of leaf nitrogen present as rubisco. Therefore, low CO₂ assimilation rates despite high leaf nitrogen concentrations in leaves of the evergreen species examined were explained not only by a low C_c but also by a relatively low proportion of leaf nitrogen being used for photosynthesis. We also show that simple one-dimensional equations describing the relationship between leaf internal conductance from stomatal cavities to the sites of carboxylation and carbon isotope discrimination (Δ) can lead to errors in the estimate of g_i. Potential effects of heterogeneity in stomatal aperture on carbon isotope discrimination may be particularly important and may lead to a dependence of g_i upon CO₂ assimilation rate. It is shown that for any concurrent measurement of A and Δ, the estimate of C_c is an overestimate of the correct photosynthetic capacity-weighted value, but this error is probably less than 1.0 Pa.     

HIGH PHOTOSYNTHETIC RATES UNDER A COLIMITATION FOR INORGANIC PHOSPHORUS AND CARBON DIOXIDE1

Inorganic phosphorus (P_i) and carbon (here, CO₂) potentially limit the photosynthesis of phytoplankton simultaneously (colimitation). A single P_i limitation generally reduces photosynthesis, but the effect of a colimitation is not known. Therefore, photosynthesis was measured under P_i‐limited conditions and high and low CO₂, and osmo‐mixotrophic (i.e., growth in the presence of glucose) conditions that result in colimiting conditions in some cases. The green alga Chlamydomonas acidophila Negoro was used as a model organism because low P_i and CO₂ concentrations likely influence its photosynthetic rates in its natural environment. Results showed a decreasing maximum photosynthetic rate (P_max) and maximum quantum yield (Φ_II) with increasing P_i limitation. In addition, a P_i limitation enhanced the relative contribution of dark respiration to P_max (R_d:P_max) but did not influence the compensation light intensity. P_max positively correlated with the cellular RUBISCO content. Osmo‐mixotrophic conditions resulted in similar P_max, Φ_II, and RUBISCO content as in high‐CO₂ cultures. The low‐CO₂ cultures were colimited by P_i and CO₂ and had the highest P_max, Φ_II, and RUBISCO content. Colimiting conditions for P_i and CO₂ in C. acidophila resulted in an enhanced mismatch between photosynthesis and growth rates compared to the effect of a single P_i limitation. Primary productivity of colimited phytoplankton could thus be misinterpreted.     

Effects of low and elevated CO2 on C3 and C4 annuals

Abutilon theophrasti (C₃) and Amaranthus retroflexus (C₄), were grown from seed at four partial pressures of CO₂: 15 Pa (below Pleistocene minimum), 27 Pa (pre-industrial), 35 Pa (current), and 70 Pa (future) in the Duke Phytotron under high light, high nutrient, and wellwatered conditions to evaluate their photosynthetic response to historic and future levels of CO₂. Net photosynthesis at growth CO₂ partial pressures increased with increasing CO₂ for C₃ plants, but not C₄ plants. Net photosynthesis of Abutilon at 15 Pa CO₂ was 70% less than that of plants grown at 35 Pa CO₂, due to greater stomatal and biochemical limitations at 15 Pa CO₂. Relative stomatal limitation (RSL) of Abutilon at 15 Pa CO₂ was nearly 3 times greater than at 35 Pa CO₂. A photosynthesis model was used to estimate ribulose-1,5-bisphosphate carboxylase (rubisco) activity (Vc_max), electron transport mediated RuBP regeneration capacity (J _max), and phosphate regeneration capacity (PiRC) in Abutilon from net photosynthesis versus intercellular CO₂ (A–C _i) curves. All three component processes decreased by approximately 25% in Abutilon grown at 15 Pa compared with 35 Pa CO₂. Abutilon grown at 15 Pa CO₂ had significant reductions in total rubisco activity (25%), rubisco content (30%), activation state (29%), chlorophyll content (39%), N content (32%), and starch content (68%) compared with plants grown at 35 Pa CO₂. Greater allocation to rubisco relative to light reaction components and concomitant decreases in J _max and PiRC suggest co-regulation of biochemical processes occurred in Abutilon grown at 15 Pa CO₂. There were no significant differences in photosynthesis or leaf properties in Abutilon grown at 27 Pa CO₂ compared with 35 Pa CO₂, suggesting that the rise in CO₂ since the beginning of the industrial age has had little effect on the photosynthetic performance of Abutilon. For Amaranthus, limitations of photosynthesis were balanced between stomatal and biochemical factors such that net photosynthesis was similar in all CO₂ treatments. Differences in photosynthetic response to growth over a wide range of CO₂ partial pressures suggest changes in the relative performance of C₃ and C₄ annuals as atmospheric CO₂ has fluctuated over geologic time.     

天竺桂凋落叶对凤仙花生长和光合特性的影响

采用盆栽实验,通过向土壤(每盆8kg)中添加0g·pot-1(CK)、20g·pot-1(L)、40g·pot-1(M)和80g·pot-1(H)天竺桂(Cinnamomum japonicum)凋落叶,模拟其自然分解对凤仙花(Impatiens balsamina)生长和光合特性的影响。结果显示:(1)添加天竺桂凋落叶M和H处理下,凤仙花生物量和地径均显著降低,而株高无明显变化;其叶绿素含量受到显著抑制,净光合速率(Pn)和水分利用效率(WUE)显著低于CK,而气孔导度(Gs)、胞间二氧化碳浓度(Ci)和蒸腾速率(Tr)3个气体交换参数显著高于CK。(2)Pn-PAR曲线和Pn-Ci曲线拟合表明,凤仙花在光饱和以及CO2饱和状态下的最大净光合速率(Pn max)、表观量子效率(AQY)、暗呼吸速率(Rd)、RuBP羧化效率(CE)和光呼吸速率(Rp)均随添加天竺桂凋落叶处理量的增加而呈下降趋势。(3)添加天竺桂凋落叶36d和67d时对凤仙花生长影响不明显,而处理58d时有明显抑制作用。研究表明,在模拟天竺桂凋落叶自然分解的土壤环境中,凤仙花的光合色素含量降低,抑制了其光合能力,对环境适应能力降低,导致凤仙花的生长受到抑制。     

Consequences of CO2 and light interactions for leaf phenology, growth, and senescence in Quercus rubra

We investigated how light and CO₂ levels interact to influence growth, phenology, and the physiological processes involved in leaf senescence in red oak (Quercus rubra) seedlings. We grew plants in high and low light and in elevated and ambient CO₂. At the end of three years of growth, shade plants showed greater biomass enhancement under elevated CO₂ than sun plants. We attribute this difference to an increase in leaf area ratio (LAR) in shade plants relative to sun plants, as well as to an ontogenetic effect: as plants increased in size, the LAR declined concomitant with a decline in biomass enhancement under elevated CO₂ Elevated CO₂ prolonged the carbon gain capacity of shade‐grown plants during autumnal senescence, thus increasing their functional leaf lifespan. The prolongation of carbon assimilation, however, did not account for the increased growth enhancement in shade plants under elevated CO₂. Elevated CO₂ did not significantly alter leaf phenology. Nitrogen concentrations in both green and senesced leaves were lower under elevated CO₂ and declined more rapidly in sun leaves than in shade leaves. Similar to nitrogen concentration, the initial slope of A/C_i curves indicated that Rubisco activity declined more rapidly in sun plants than in shade plants, particularly under elevated CO₂. Absolute levels of chlorophyll were affected by the interaction of CO₂ and light, and chlorophyll content declined to a minimal level in sun plants sooner than in shade plants. These declines in N concentration, in the initial slope of A/C_i curves, and in chlorophyll content were consistent with declining photosynthesis, such that elevated CO₂ accelerated senescence in sun plants and prolonged leaf function in shade plants. These results have implications for the carbon economy of seedlings and the regeneration of red oak under global change conditions.