Do the capacity and kinetics for modification of xanthophyll cycle pool size depend on growth irradiance in temperate trees?

The present study investigated the interaction of growth irradiance (Q_int) with leaf capacity for and kinetics of adjustment of the pool size of xanthophyll cycle carotenoids (sum of violaxanthin, antheraxanthin and zeaxanthin; VAZ) and photosynthetic electron transport rate (J_max) after changes in leaf light environment. Individual leaves of lower‐canopy/lower photosynthetic capacity species Tilia cordata Mill. and upper canopy/higher photosynthetic capacity species Populus tremula L. were either illuminated by additional light of 500–800 µmol m^?2 s^?1 for 12 h photoperiod or enclosed in shade bags. The extra irradiance increased the total amount of light intercepted by two‐fold for the upper and 10–15‐fold for the lower canopy leaves, whereas the shade bags transmitted 45% of incident irradiance. In control leaves, VAZ/area, VAZ/Chl and J_max were positively associated with leaf growth irradiance (Q_int). After 11 d extra illumination, VAZ/Chl increased in all cases due to a strong reduction in foliar chlorophyll, but VAZ/area increased in the upper canopy leaves of both species, and remained constant or decreased in the lower canopy leaves of T. cordata. The slope for VAZ/area changes with cumulative extra irradiance was positively associated with Q_int only in T. cordata, but not in P. tremula. Nevertheless, all leaves of P. tremula increased VAZ/area more than the most responsive leaves of T. cordata. Shading reduced VAZ content only in P. tremula, but not in T. cordata, again demonstrating that P. tremula is a more responsive species. Compatible with the hypothesis of the role of VAZ in photoprotection, the rates of photosynthetic electron transport declined less in P. tremula than in T. cordata after the extra irradiance treatment. However, foliar chlorophyll contents of the exposed leaves declined significantly more in the upper canopy of P. tremula, which is not consistent with the suggestion that the leaves with the highest VAZ content are more resistant to photoinhibition. This study demonstrates that previous leaf light environment may significantly affect the adaptation capacity of foliage to altered light environment, and also that species differences in photosynthetic capacity and acclimation potentials importantly alter this interaction.     

Shape of leaf photosynthetic electron transport versus temperature response curve is not constant along canopy light gradients in temperate deciduous trees

Responses of foliar light-saturated net assimilation rate (A_max), capacity for photosynthetic electron transport (J_max) and mitochondrial respiration rate (R_d) to long-term canopy light and temperature environment were investigated in a temperate deciduous canopy composed of Populus tremula L. in the upper (17–28 m) and of Tilia cordata Mill. in the lower canopy layer (4–17 m). Climatic measurements indicated that seasonal average daily maximum air temperature (T_max) was 5·5 °C (range 0·7–10·5 °C) higher in the top than in the bottom of the canopy, and strong positive correlations were observed between T_max and seasonal average integrated quantum flux density (Q_int), as well as between seasonal average daily mean temperature and Q_int. Because of changes in leaf dry mass and nitrogen per unit area, A_max, J_max, and R_d scaled positively with Q_int in both species at a common leaf temperature (T). According to J_max versus T response curves and dark chlorophyll fluorescence transients, photosynthetic electron transport was less heat resistant in P. tremula with optimum temperature of J_max, T_opt, of 33·5 ± 0·6 °C than in T. cordata with T_opt of 40·7 ± 0·6 °C. This difference was suggested to manifest evolutionary adaptation of photosynthetic electron transport to cooler environments in P. tremula, the range of which extends farther north than that in T. cordata. Possibly because of acclimation to long-term canopy temperature environment, T_opt was positively related to Q_int in P. tremula, foliage of which was also exposed to higher irradiances and temperatures, but not in T. cordata, in the canopy of which quantum flux densities and temperatures were lower, and gradients in the environmental factors less pronounced. Parallel to changes in T_opt, the activation energy for photosynthetic electron transport decreased with increasing Q_int in P. tremula, indicating that J_max of leaves acclimated to colder environment was more responsive to T in lower temperatures than that of high T acclimated leaves. Similar alterations in the activation energy for mitochondrial respiration rate were also observed, indicating that acclimation to temperature of mitochondrial and chloroplastic electron transport proceeds in a co-ordinated manner, and possibly involves long-term changes in membrane fluidity properties. We conclude that, because of correlations between temperature and light, the shapes of J_max versus T, and R_d versus T response curves vary within tree canopies, and this needs to be taken account in modelling whole canopy photosynthesis.     

Within-canopy variation in the rate of development of photosynthetic capacity is proportional to integrated quantum flux density in temperate deciduous trees

The present study was undertaken to test for the hypothesis that the rate of development in the capacity for photosynthetic electron transport per unit area (J_max;A), and maximum carboxylase activity of Rubisco (V_cmax;A) is proportional to average integrated daily quantum flux density (Q_int) in a mixed deciduous forest dominated by the shade‐intolerant species Populus tremula L., and the shade‐tolerant species Tilia cordata Mill. We distinguished between the age‐dependent changes in net assimilation rates due to modifications in leaf dry mass per unit area (M_A), foliar nitrogen content per unit dry mass (N_M), and fractional partitioning of foliar nitrogen in the proteins of photosynthetic electron transport (F_B), Rubisco (F_R) and in light‐harvesting chlorophyll‐protein complexes (V_cmax;A ∝ M_AN_MF_R; J_max;A ∝ M_AN_MF_B). In both species, increases in J_max;A and V_cmax;A during leaf development were primarily determined by nitrogen allocation to growing leaves, increases in leaf nitrogen partitioning in photosynthetic machinery, and increases in M_A. Canopy differences in the rate of development of leaf photosynthetic capacity were mainly controlled by the rate of change in M_A. There was only small within‐canopy variation in the initial rate of biomass accumulation per unit Q_int (slope of M_A versus leaf age relationship per unit Q_int), suggesting that canopy differences in the rate of development of J_max;A and V_cmax;A are directly proportional to Q_int. Nevertheless, M_A, nitrogen, J_max;A and V_cmax;A of mature leaves were not proportional to Q_int because of a finite M_A in leaves immediately after bud‐burst (light‐independent component of M_A). M_A, leaf chlorophyll contents and chlorophyll : N ratio of mature leaves were best correlated with the integrated average quantum flux density during leaf development, suggesting that foliar photosynthetic apparatus, once developed, is not affected by day‐to‐day fluctuations in Q_int. However, for the upper canopy leaves of P. tremula and for the entire canopy of T. cordata, there was a continuous decline in N contents per unit dry mass in mature non‐senescent leaves on the order of 15–20% for a change of leaf age from 40 to 120 d, possibly manifesting nitrogen reallocation to bud formation. The decline in N contents led to similar decreases in leaf photosynthetic capacity and foliar chlorophyll contents. These data demonstrate that light‐dependent variation in the rate of developmental changes in M_A determines canopy differences in photosynthetic capacity, whereas foliar photosynthetic apparatus is essentially constant in fully developed leaves.     

Interactive effects of nitrogen and phosphorus on the acclimation potential of foliage photosynthetic properties of cork oak,Quercus suber,to elevated atmospheric CO2 concentrations

Leaf gas-exchange and chemical composition were investigated in seedlings of Quercus suber L. grown for 21 months either at elevated (700 μmol mol^–1) or normal (350 μmol mol^–1) ambient atmospheric CO₂ concentrations, [CO₂], in a sandy nutrient-poor soil with either ‘high’ N (0.3 mol N m^–3 in the irrigation solution) or with ‘low’ N (0.05 mol N m^–3) and with a constant suboptimal concentration of the other macro- and micronutrients. Although elevated [CO₂] yielded the greatest total plant biomass in ‘high’ nitrogen treatment, it resulted in lower leaf nutrient concentrations in all cases, independent of the nutrient addition regime, and in greater nonstructural carbohydrate concentrations. By contrast, nitrogen treatment did not affect foliar N concentrations, but resulted in lower phosphorus concentrations, suggesting that under lower N, P use-efficiency in foliar biomass production was lower. Phosphorus deficiency was evident in all treatments, as photosynthesis became CO₂ insensitive at intercellular CO₂ concentrations larger than ≈ 300 μmol mol^–1, and net assimilation rates measured at an ambient [CO₂] of 350 μmol mol^–1 or at 700 μmol mol^–1 were not significantly different. Moreover, there was a positive correlation of foliar P with maximum Rubisco (Ribulose-1,5-bisphosphate carboxylase/oxygenase) carboxylase activity (V_cmax), which potentially limits photosynthesis at low [CO₂], and the capacities of photosynthetic electron transport (J_max) and phosphate utilization (P_max), which are potentially limiting at high [CO₂]. None of these potential limits was correlated with foliar nitrogen concentration, indicating that photosynthetic N use-efficiency was directly dependent on foliar P availability. Though the tendencies were towards lower capacities of potential limitations of photosynthesis in high [CO₂] grown specimens, the effects were statistically insignificant, because of (i) large within-treatment variability related to foliar P, and (ii) small decreases in P/N ratio with increasing [CO₂], resulting in balanced changes in other foliar compounds potentially limiting carbon acquisition. The results of the current study indicate that under P-deficiency, the down-regulation of excess biochemical capacities proceeds in a similar manner in leaves grown under normal and elevated [CO₂], and also that foliar P/N ratios for optimum photosynthesis are likely to increase with increasing growth CO₂ concentrations. Symbols: A, net assimilation rate (μmol m^–2 s^–1); A_max, light-saturated A (μmol m^–2 s^–1); α, initial quantum yield at saturating [CO₂] and for an incident Q (mol mol^–1); [CO₂], atmospheric CO₂ concentration (μmol mol^–1); C_i, intercellular CO₂ concentration (μmol mol^–1); C_a, CO₂ concentration in the gas-exchange cuvette (μmol mol^–1); F_B, fraction of leaf N in ‘photoenergetics’; F_L, fraction of leaf N in light harvesting; F_R, fraction of leaf N in Rubisco; Γ*, CO₂ compensation concentration in the absence of R_d (μmol mol^–1); J_max*, capacity for photosynthetic electron transport; J_mc, capacity for photosynthetic electron transport per unit cytochrome f (mol e^–[mol cyt f]^–1 s^–1); K_c, Michaelis-Menten constant for carboxylation (μmol mol^–1); K_o, Michaelis-Menten constant for oxygenation (mmol mol^–1); M_A, leaf dry mass per area (g m^–2); O, intercellular oxygen concentration (mmol mol^–1); [P_i], concentration of inorganic phosphate (mM); P_max*, capacity for phosphate utilization; Q, photosynthetically active quantum flux density (μmol m^–2 s^–1); R_d*, day respiration (CO₂ evolution from nonphotorespiratory processes continuing in the light); Rubisco, ribulose-1,5-bisphosphate carboxylase/oxygenase; RUBP, ribulose-1,5-bisphosphate; T_l, leaf temperature (°C); U_TPU*, rate of triose phosphate utilization; V_cmax*, maximum Rubisco carboxylase activity; V_cr, specific activity of Rubisco (μmol CO₂[g Rubisco]^–1 s^–1] *given in either μmol m^–2 s^–1 or in μmol g^–1 s^–1 as described in the text.     

Contrasting effects of long term versus short-term nitrogen addition on photosynthesis and respiration in the Arctic

We examined the effects of short (<1–4 years) and long-term (22 years) nitrogen (N) and/or phosphorus (P) addition on the foliar CO₂ exchange parameters of the Arctic species Betula nana and Eriophorum vaginatum in northern Alaska. Measured variables included: the carboxylation efficiency of Rubisco (V_cmax), electron transport capacity (J_max), dark respiration (R_d), chlorophyll a and b content (Chl), and total foliar N (N). For both B. nana and E. vaginatum, foliar N increased by 20–50 % as a consequence of 1–22 years of fertilisation, respectively, and for B. nana foliar N increase was consistent throughout the whole canopy. However, despite this large increase in foliar N, no significant changes in V_cmax and J_max were observed. In contrast, R_d was significantly higher (>25 %) in both species after 22 years of N addition, but not in the shorter-term treatments. Surprisingly, Chl only increased in both species the first year of fertilisation (i.e. the first season of nutrients applied), but not in the longer-term treatments. These results imply that: (1) under current (low) N availability, these Arctic species either already optimize their photosynthetic capacity per leaf area, or are limited by other nutrients; (2) observed increases in Arctic NEE and GPP with increased nutrient availability are caused by structural changes like increased leaf area index, rather than increased foliar photosynthetic capacity and (3) short-term effects (1–4 years) of nutrient addition cannot always be extrapolated to a larger time scale, which emphasizes the importance of long-term ecological experiments.     

No photosynthetic down-regulation in sweetgum trees (Liquidambar styraciflua L.) after three years of CO2 enrichment at the Duke Forest FACE experiment

Photosynthetic capacity and leaf properties of sun and shade leaves of overstorey sweetgum trees (Liquidambar styraciflua L.) were compared over the first 3 years of growth in ambient or ambient + 200 μL L^?1 CO₂ at the Duke Forest Free Air CO₂ Enrichment (FACE) experiment. We were interested in whether photosynthetic down‐regulation to CO₂ occurred in sweetgum trees growing in a forest ecosystem, whether shade leaves down‐regulated to a greater extent than sun leaves, and if there was a seasonal component to photosynthetic down‐regulation. During June and September of each year, we measured net photosynthesis (A) versus the calculated intercellular CO₂ concentration (C_i) in situ and analysed these response curves using a biochemical model that described the limitations imposed by the amount and activity of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Vc_max) and by the rate of ribulose‐1,5‐bisphosphate (RuBP) regeneration mediated by electron transport (J_max). There was no evidence of photosynthetic down‐regulation to CO₂ in either sun or shade leaves of sweetgum trees over the 3 years of measurements. Elevated CO₂ did not significantly affect Vc_max or J_max. The ratio of Vc_max to J_max was relatively constant, averaging 2·12, and was not affected by CO₂ treatment, position in the canopy, or measurement period. Furthermore, CO₂ enrichment did not affect leaf nitrogen per unit leaf area (N_a), chlorophyll or total non‐structural carbohydrates of sun or shade leaves. We did, however, find a strong relationship between N_a and the modelled components of photosynthetic capacity, Vc_max and J_max. Our data over the first 3 years of this experiment corroborate observations that trees rooted in the ground may not exhibit symptoms of photosynthetic down‐regulation as quickly as tree seedlings growing in pots. There was a strong sustained enhancement of photosynthesis by CO₂ enrichment whereby light‐saturated net photosynthesis of sun leaves was stimulated by 63% and light‐saturated net photosynthesis of shade leaves was stimulated by 48% when averaged over the 3 years. This study suggests that this CO₂ enhancement of photosynthesis will be sustained in the Duke Forest FACE experiment as long as soil N availability keeps pace with photosynthetic and growth processes.     

Low temperature stress modifies the photochemical efficiency of a tropical tree species <Emphasis Type="Italic">Hevea brasiliensis</Emphasis>: effects of varying concentration of CO<Subscript>2</Subscript> and photon flux density

Two clones of Hevea brasiliensis (RRII 105 and PB 235) were grown for one year in two distinct agroclimatic locations (warmer and colder, W and C) in peninsular India. We simultaneously measured gas exchange and chlorophyll (Chl) fluorescence on fully mature intact leaves at different photosynthetic photon flux densities (PPFDs) and ambient CO₂ concentrations (C _a) and at constant ambient O₂ concentration (21 %). Net photosynthetic rate (P _N), apparent quantum yield for CO₂ assimilation (Φ_c), in vivo carboxylation efficiency (CE), and photosystem 2 quantum yield (Φ_PS2) were low in plants grown in C climate and these reductions were more predominant in RRII 105 than in PB 235 which was also reflected in their growth. We estimated in these clones the partitioning of photosynthetic electrons between CO₂ reduction (J_A) and processes other than CO₂ reduction (J^*) at low and high PPFDs and C _a. At high C _a (700 µmol mol⁻¹) most of the photosynthetic electrons were used for CO₂ assimilation and negligible amount went for other processes when PPFD was low (200–300 µmol m⁻² s⁻¹) both in the C and W climates. But at high PPFD (900-1 100 µmol m⁻² s⁻¹), J^* was appreciably high even at a high C _a. Hence at normal ambient C _a and high irradiance, electrons can be generated in the photosynthetic apparatus far in excess of what can be safely utilised for photosynthetic CO₂ reduction. However, at high C _a there was increased diversion of electrons to photosynthetic CO₂ reduction which resulted in improved photosynthetic parameters even in plants grown in C climate.     

Nitrogen deposition limits photosynthetic response to elevated CO2 differentially in a dioecious species

Sexual dimorphisms of dioecious plants are important in controlling and maintaining sex ratios under changing climate environments. Yet, little is known about sex-specific responses to elevated CO₂ with soil nitrogen (N) deposition. To investigate sex-related physiological and biochemical responses to elevated CO₂ with N deposition, Populus cathayana Rehd. was employed as a model species. The cuttings were subjected to two CO₂ regimes (350 and 700???mol?mol^?1) with two N levels (0 and 5?g?N?m^?2?year^?1). Our results showed that elevated CO₂ and N deposition separately increased the total number of leaves, leaf area (LA), leaf mass, net photosynthetic rate (P _n), light saturated photosynthetic rate (P _max), chlorophyll a (Chl a), and chlorophyll a to chlorophyll b ratio (Chl a/b) in both males and females of P. cathayana. However, the effects on LA, leaf mass, P _n, P _max, Chl a and Chl a/b were weakened under the combined treatment of elevated CO₂ and N deposition. Males had higher leaf mass, P _n, P _max, apparent quantum yield (??), carboxylation efficiency (CE), Chl a, Chl a/b, leaf N, and root carbon to N ratio (C/N) than did females under elevated CO₂ with N deposition. In contrast to males, females had significantly higher levels of soluble sugars in leaves and greater starch accumulation in roots and stems under the same condition. The results of the present work imply that P. cathayana females are more responsive and suffer from greater negative effects on growth and photosynthetic capacity than do males when grown under elevated CO₂ with soil N deposition.     

Changes in photosynthesis,fluorescence, and nitrogen metabolism of hawthorn (<Emphasis Type="Italic">Crataegus pinnatifida</Emphasis>) in response to exogenous glutamic acid

Photosynthesis, chlorophyll (Chl) a fluorescence, and nitrogen metabolism of hawthorn (Crataegus pinnatifida Bge.), subjected to exogenous L-glutamic acid (GLA) (200 mg l⁻¹, 400 mg l⁻¹, and 800 mg l⁻¹) that possibly affect secondary metabolic regulation, were measured. The results indicated that photosynthetic and fluorescence characteristics of hawthorn exhibited positive responses to the application of GLA. Different concentrations of GLA caused an increase in Chl content, net photosynthetic rate (P _N) and stomatal conductance (g _s) as well as transpiration rate (E), and improved the carboxylation efficiency (CE), apparent quantum yield (AQY) and maximum carboxylation velocity of Rubisco (V_cmax). Application of GLA could also enhance the maximum ratio of quantum yields of photochemical and concurrent non-photochemical processes in PSII (F_v/F₀), the maximal quantum yield of PSII (F_v/F_m), the probability that an absorbed photon will move an electron into the electron transport chain beyond Q_A (Φ_Eo) as well as the performance index on absorption basis (PI_ABS), but decreased the intercellular CO₂ concentration (C _i) and the minimal fluorescence (F₀). Application of GLA also induced an increase in nitrate reductase (NR; EC 1.6.6.1) and glutamine synthetase (GS; EC 6.3.1.2) activities, and increased the soluble protein content, leaf nitrogen (N) content and N accumulation in leaves as well as the plant biomass. However, the effects were different among different concentrations of GLA, and 800 mg l⁻¹ GLA was better. This finding suggested that application of GLA is recommended to improve the photosynthetic capacity by increasing the light energy conversion and CO₂ transfer as well as the photochemical efficiency of PSII, and enhanced the nitrogen metabolism and growth and development of plants.     

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.     

铝胁迫对木荷幼苗光合特性的影响及添加盐基阳离子和磷的调节作用

为探讨铝(Al)胁迫对木荷(Schima superba)幼苗光合特征的影响,采用营养液水培的方法,对铝(Al)胁迫下木荷幼苗的光合响应及盐基阳离子(BC)和磷(P)的调节作用进行了研究。结果表明,在低浓度Al(0.25 mmol L–1)处理下,木荷幼苗的光合色素(Chl a、Chl b、Car)含量、光合作用参数(P n、g s、WUE、C i/C a)以及光响应特征参数(P max、AQY、R d、LSP)均呈下降趋势,添加BC或同时添加BC和P均能缓解上述参数的降低。中、高浓度Al(0.75、1.50 mmol L–1)处理,除光合色素含量呈增加趋势外,光合作用参数、光响应特征参数均下降,且下降幅度随Al浓度的升高而增大,添加P比添加BC更能有效缓解Al胁迫对木荷幼苗的影响。这揭示了BC、P在缓解木荷Al胁迫的相对重要性。     

Influence of irradiance on photosynthesis and PSII photochemical efficiency in maize during short-term exposure at high CO<Subscript>2</Subscript> concentration

This work aimed to evaluate if gas exchange and PSII photochemical activity in maize are affected by different irradiance levels during short-term exposure to elevated CO₂. For this purpose gas exchange and chlorophyll a fluorescence were measured on maize plants grown at ambient CO₂ concentration (control CO₂) and exposed for 4 h to short-term treatments at 800 μmol(CO₂) mol⁻¹ (high CO₂) at a photosynthetic photon flux density (PPFD) of either 1,000 μmol m⁻² s⁻¹ (control light) or 1,900 μmol m⁻² s⁻¹ (high light). At control light, high-CO₂ leaves showed a significant decrease of net photosynthetic rate (P _N) and a rise in the ratio of intercellular to ambient CO₂ concentration (C _i/C _a) and water-use efficiency (WUE) compared to control CO₂ leaves. No difference between CO₂ concentrations for PSII effective photochemistry (Φ_PSII), photochemical quenching (q_p) and nonphotochemical quenching (NPQ) was detected. Under high light, high-CO₂ leaves did not differ in P _N, C _i/C _a, Φ_PSII and NPQ, but showed an increase of WUE. These results suggest that at control light photosynthetic apparatus is negatively affected by high CO₂ concentration in terms of carbon gain by limitations in photosynthetic dark reaction rather than in photochemistry. At high light, the elevated CO₂ concentration did not promote an increase of photosynthesis and photochemistry but only an improvement of water balance due to increased WUE.     

Shoots grafted into the upper crowns of tall Japanese cedar (Cryptomeria japonica D. Don) show foliar gas exchange characteristics similar to those of intact shoots

The lower foliar photosynthetic rates seen in shoots in the upper crowns of tall trees than those in seedlings could be caused by extrinsic factors, such as hydraulic conductance, for shoots or by irreversible intrinsic change in the meristems during tree development. To clarify which is most significant, we compared foliar gas exchange characteristics and water relations among scions of Japanese cedar (Cryptomeria japonica D. Don) grafted into the upper crowns of tall trees, rooted cuttings developed from scions of the same clones, and intact shoots in the upper crowns of the tall trees. Grafted shoots had the same water regime as intact shoots, as confirmed by their similar water potentials at the turgor loss point, which were more negative than those of the rooted cuttings. No significant difference was observed between the grafted and intact shoots in their light-saturated photosynthetic rate (P_max), stomatal conductance (g_s), photosynthetic capacity, carboxylation efficiency, ratio of intercellular to ambient CO₂ concentration (C_i/C_a), and carbon isotope composition (¹³C). Compared with the rooted cuttings, the grafted shoots showed significantly lower P_max, g_s, photosynthetic capacity, and carboxylation efficiency (to 49%, 31%, 68%, and 65%, respectively). The C_i/C_a and ¹³C indicated significantly stronger instantaneous and long-term stomatal limitation in the grafted shoots than in the rooted cuttings. These indicate that changes in extrinsic factors can reduce foliar photosynthetic rates in shoots in the upper crowns of tall trees as a result of stronger stomatal limitation and reduced photosynthetic activity, without irreversible intrinsic changes in the meristems.     

Photosynthetic apparatus organization and function in the wild type and a chlorophyll b-less mutant of Chlamydomonas reinhardtii. Dependence on carbon source

 The assembly, organization and function of the photosynthetic apparatus was investigated in the wild type and a chlorophyll (Chl) b-less mutant of the unicellular green alga Chlamydomonas reinhardtii, generated via DNA insertional mutagenesis. Comparative analyses were undertaken with cells grown photoheterotrophically (acetate), photomixotrophically (acetate and HCO⁻ ₃) or photoautotrophically (HCO⁻ ₃). It is shown that lack of Chl b diminished the photosystem-II (PSII) functional Chl antenna size from 320 Chl (a and b) to about 95 Chl a molecules. However, the functional Chl antenna size of PSI remained fairly constant at about 290 Chl molecules, independent of the presence of Chl b. Western blot and kinetic analyses suggested the presence of inner subunits of the Chl a-b light-harvesting complex of PSII (LHCII) and the entire complement of the Chl a-b light-harvesting complex of PSI (LHCI) in the mutant. It is concluded that Chl a can replace Chl b in the inner subunits of the LHCII and in the entire complement of the LHCI. Growth of cells on acetate as the sole carbon source imposes limitations in the photon-use efficiency and capacity of photosynthesis. These are manifested as a lower quantum yield and lower light-saturated rate of photosynthesis, and as lower variable to maximal (F_v/F_max) chlorophyll fluorescence yield ratios. This adverse effect probably originates because acetate shifts the oxidation-reduction state of the plastoquinone pool, and also because it causes a decrease in the amount and/or activity of Rubisco in the chloroplast. Such limitations are fully alleviated upon inclusion of an inorganic carbon source (e.g. bicarbonate) in the cell growth medium. Further, the work provides evidence to show that transformation of green algae can be used as a tool by which to generate mutants exhibiting a permanently truncated Chl antenna size and a higher (per Chl) photosynthetic productivity of the cells. Received: 10 November 1999 / Accepted: 22 December 1999     

Shade does not ameliorate drought effects on the tree fern species Dicksonia antarctica and Cyathea australis

We examined the responses of two tree fern species (Dicksonia antarctica and Cyathea australis) growing under moderate and high light regimes to short-term water deficit followed by rewatering. Under adequate water supply, morphological and photosynthetic characteristics differed between species. D. antarctica, although putatively the more shade and less drought adapted species, had greater chlorophyll a/b ratio, and greater water use efficiency and less negative δ¹³C. Both species were susceptible to water deficit regardless of the light regime showing significant decreases in photosynthetic parameters (A _max, V _cmax, J _max) and stomatal conductance (g _s ) in conjunction with decreased relative frond water content (RWC) and predawn frond water potential (Ψ_predawn). During the water deficit period, decreases in g _s in both species started one day later, and were at lower soil water content, under moderate light compared with high light. D. antarctica under moderate light was more vulnerable to drought than all other plants as was indicated by greater decreases in Ψ_predawn, lowest stomatal conductance, and photosynthetic rates. Both tree fern species were able to recover after a short but severe water stress.     

The energy distribution between the photosystems and light-induced changes in the stoichiometry of system I and II reaction centers in the chlorophyll b-containing alga Mantoniella squamata (Prasinophyceae)

The chlorophyll b-containing alga Mantoniella squamata was analyzed with respect to its capacity to balance the energy distribution from the light-harvesting antenna to photosystem I or photosystem II. It was shown, that this alga is unable to alter the absorption cross section of the two photosystems in terms of short-time regulations (state transitions). The energy absorbed by the LHC, which contains 60% of total photosynthetic pigments, is transferred to both photosystems without any preference. The stoichiometry of the two photosystems is found to be extremely unequal and variable during light adaptation. In high light, the molar ratio of P-680 per P-700 is found to be two, whereas under low light conditions this ratio accounts to nearly four. This very unbalanced stoichiometry of the reaction centers gives some new insights into the concept of the photosynthetic unit as well as in the importance of the regulation of the energy distribution. It is assumed that the high concentration of photosystem II can be understood as a mechanism to prevent the overexcitation of photosystem I. In addition, the changes im membrane protein pattern are not accompanied by variations in the ratio of appressed to nonappressed membranes as probed by ultrastructural analysis. It is suggested that the thylakoids are organized like a homogenous pigment bed. The lack of state transitions can be interpreted as a consequence of this unusual membrane morphology.Abbreviations Chl chlorophyll - CPa chlorophyll a-protein of PSII - CPl P-700 chlorophyll a-protein - CPD Chlorophyll packing density index - cyt f cytochrome f - FP free pigments - LHC light-harvesting complex - P_max light saturated photosynthetic rates per chlorophyll - n number of experiments - PQ plastoquinone - PS photosystem - PSU photosynthetic unit - Q_E non-photochemical quenching - Q_Q photochemical quenching     

A model separating leaf structural and physiological effects on carbon gain along light gradients for the shade-tolerant species Acer saccharum

A process-based leaf gas exchange model for C₃ plants was developed which specifically describes the effects observed along light gradients of shifting nitrogen investment in carboxylation and bioenergetics and modified leaf thickness due to altered stacking of photosynthetic units. The model was parametrized for the late-successional, shade-tolerant deciduous species Acer saccharum Marsh. The specific activity of ribulose-1,5-bisphosphate carboxylase (Rubisco) and the maximum photosynthetic electron transport rate per unit cytochrome f (cyt f) were used as indices that vary proportionally with nitrogen investment in the capacities for carboxylation and electron transport. Rubisco and cyt f per unit leaf area are related in the model to leaf dry mass per area (M_A), leaf nitrogen content per unit leaf dry mass (N_m), and partitioning coefficients for leaf nitrogen in Rubisco (P_R) and in bioenergetics (P_B). These partitioning coefficients are estimated from characteristic response curves of photosynthesis along with information on lear structure and composition. While P_R and P_B determine the light-saturated value of photosynthesis, the fraction of leaf nitrogen in thylakoid light-harvesting components (P_L) and the ratio of leaf chlorophyll to leaf nitrogen invested in light harvesting (C_B), which is dependent on thylakoid stoichiometry, determine the initial photosynthetic light utilization efficiency in the model. Carbon loss due to mitochondrial respiration, which also changes along light gradients, was considered to vary in proportion with carboxylation capacity. Key model parameters - N_m, P_R, P_B, P_LC_B and stomatal sensitivity with respect to changes in net photosynthesis (G_r) – were examined as a function of M_A, which is linearly related to irradiance during growth of the leaves. The results of the analysis applied to A. saccharum indicate that P_B and P_R increase, and G_f, P_L and C_B decrease with increasing M_A. As a result of these effects of irradiaiice on nitrogen partitioning, the slope of the light-saturated net photosynthesis rate per unit leaf dry mass (A^m_max) versus N_m relationship increased with increasing growth irradiance in mid-season. Furthermore, the nitrogen partitioning coefficients as well as the slopes of A^m_max versus N_m were independent of season, except during development of the leaf photosynthetic apparatus. Simulations revealed that the acclimation to high light increased A^m_max by 40% with respect to the low light regime. However, light-saturated photosynthesis per leaf area (A^a_max) varied 3-fold between these habitats, suggesting that the acclimation to high light was dominated by adjustments in leaf anatomy (A^a_max=A^m_maxM_A) rather than in foliar biochemistry. This differed from adaptation to low light, where the alterations in foliar biochemistry were predicted to be at least as important as anatomical modifications. Due to the light-related accumulation of photosynthetic mass per unit area, A^a_max depended on M_A and leaf nitrogen per unit area (N_a). However, N_a conceals the variation in both M_A and N_m (N_a=N_mM_A), and prevents clear separation of anatomical adjustments in foliage structure and biochemical modifications in foliar composition. Given the large seasonal and site nutrient availability-related variation in N_m, and the influences of growth irradiance on nitrogen partitioning, the relationship between A^a_max and N_a is universal neither in time nor in space and in natural canopies at mid-season is mostly driven by variability in M_A. Thus, we conclude that analyses of the effects of nitrogen investments on potential carbon acquisition should use mass-based rather than area-based expressions.     

Altitudinal variation in photosynthetic capacity, diffusional conductance and δ13C of butterfly bush (Buddleja davidii) plants growing at high elevations

In this study, we have examined several physiological, biochemical and morphological features of Buddleja davidii plants growing at 1300 m above sea level (a.s.l.) and 3400 m a.s.l., respectively, to identify coordinated changes in leaf properties in response to reduced CO₂ partial pressure (P_a). Our results confirmed previous findings that foliar δ¹³C, photosynthetic capacity and foliar N concentration on a leaf area basis increased, whereas stomatal conductance (g_s) decreased with elevation. The net CO₂ assimilation rate (A_max), maximum rate of electron transport (J_max) and respiration increased significantly with elevation, although no differences were found in carboxylation efficiency of Rubisco (V_cmax). Consequently, also the J_max to V_cmax ratio was significantly increased by elevation, indicating that the functional balance between Ribulose‐1,5‐biphosphate (RuBP) consumption and RuBP regeneration changes as elevation increases. Our results also indicated a homeostatic response of CO₂ transfer conductance inside the leaf (mesophyll conductance, g_m) to increasing elevation. In fact, with elevation, g_m also increased compensating for the strong decrease in g_s and, thus, in the P_i (intercellular partial pressure of CO₂) to P_a ratio, leading to similar chloroplast partial pressure of CO₂ (P_c) to P_a ratio at different elevations. Because there were no differences in V_cmax, also A measured at similar PPFD and leaf temperature did not differ statistically with elevation. As a consequence, a clear relationship was found between A and g_m, and between A and the sum of g_s and g_m. These data suggest that the higher dry mass δ¹³C of leaves at the higher elevation, indicative of lower long‐term P_c/P_a ratio, cannot be attributed to changes either in diffusional resistances or in carboxylation efficiency. We speculate that because temperature significantly decreases as the elevation increases, it dramatically affects CO₂ diffusion and hence P_c/P_a and, consequently, is the primary factor influencing ¹³C discrimination at high elevation.     

Acclimation of photosynthetic characteristics of the moss Pleurozium schreberi to among‐habitat and within‐canopy light gradients

Light availability varies strongly among moss habitats and within the moss canopy, and vertical variation in light within the canopy further interacts with the age gradient. The interacting controls by habitat and canopy light gradient and senescence have not been studied extensively. We measured light profiles, chlorophyll (Chl), carotenoid (Car) and nitrogen (N) concentrations, and photosynthetic electron transport capacity (J_max) along habitat and canopy light gradients in the widespread, temperate moss Pleurozium schreberi to separate sources of variation in moss chemical and physiological traits. We hypothesised that this species, like typical feather mosses with both apical and lateral growth, exhibits greater plasticity in the canopy than between habitats due to deeper within‐canopy light gradients. For the among‐habitat light gradient, Chl, Chl/N and Chl/Car ratio increased with decreasing light availability, indicating enhanced light harvesting in lower light and higher capacity for photoprotection in higher light. N and J_max were independent of habitat light availability. Within the upper canopy, until 50–60% above‐canopy light, changes in moss chemistry and photosynthetic characteristics were analogous to patterns observed for the between‐habitat light gradient. In contrast, deeper canopy layers reflected senescence of moss shoots, with pigment and nitrogen concentrations and photosynthetic capacity decreasing with light availability. Thus, variation in chemical and physiological traits within the moss canopy is a balance between acclimation and senescence. This study demonstrates extensive light‐dependent variation in moss photosynthetic traits, but also that between‐habitat and within‐canopy light gradient affects moss physiology and chemistry differently.     

PIGMENT AND PHOTOSYNTHETIC RESPONSES OF OSCILLATORIA AGARDHII (CYANOPHYTA) TO PHOTON FLUX DENSITY AND SPECTRAL QUALITY1

The effects of photon flux density (PFD) and spectral quality on biomass, pigment content and composition, and the photosynthetic activity of Oscillatoria agardhii Gomont were investigated in steady-state populations. For alterations of PFD, chemostat populations were exposed to 50, 130 and 230 μmol photons·m^?2·s^?1 of photosynthetic active radiation (PAR). Decreases in biomass, chlorophyll a (Chl a) and c-phycocyanin (CPC) contents, and CPC: Chl a and CPC: carotenoid content was not altered. Increases in the relative abundances of myxoxanthophyll and zeaxanthin and deceases in the relative abundances of echinenone and β-carotene within the carotenoid pigments coincided with increasing PFD. Increases in Chl a-specific photosynthetic rates and maxima and decreases in biomass-specific photosynthetic rates and maxima with increasing PFD were attributed to increased light harvesting by carotenoids per unit Chl a and reduction in total pigment content, respectively. Responses to spectral quality were tested by exposing chemostat populations to a gradient of spectral transmissions at 50 μmol photons·m^?2·s^?1 PAR. Biomass differences among populations were likely attributable to the distinct absorption of the PAR spectrum by Chl a, CPC, and carotenoids. Although pigment contents were not altered by spectral quality, relative abundances of zeaxanthin and echinenone in the carotenoid pigments increased in populations exposed to high-wavelength PAR. The population adapted to green light possessed a greater photosynthetic maximum than populations adapted to other spectral qualities.