Temporal changes in biomass specific photosynthesis during the summer: regulation by environmental factors and the importance of phytoplankton succession

Measurements of phytoplankton photosynthesis vs. irradiance relationships have been made at 3–7 day intervals in Lake Erken (central Sweden) for three years during summer stratification. Both the rate of light-limited (^B) and light-saturated (P_max ^B) photosynthesis per unit chlorophyll a showed distinct and similar temporal trends in each year. Seasonal trends were especially evident for P_max ^B, which increased in value for several weeks following the onset of thermal stratification, and then declined in the presence of the large colonial blue-green alga, Gloeotrichia echinulata. By late summer, when the biomass of G. echinulata had decreased, P_max ^B again rose to its early summer value. The covariation of biomass-specific photosynthesis with the blooming of G. echinulata was the one clear seasonal (week-month) pattern which emerged in each of 3 years. Over short (day-week) time scales, changes in ^B were related to changes in irradiance exposure on the day of sampling. However, the relationship between these two parameters was variable in time, since it was superimposed upon longer term trends controlled by changes in phytoplankton species composition. Increases in G. echinulata biomass corresponded with a deepening of the thermocline, which both increased internal phosphorus loading and the transport of resting G. echinulata colonies into the epilimnion. The timing and magnitude of the yearly G. echinulata bloom was as a result related to the seasonal development of thermal stratification. These results illustrate the importance of seasonal changes in the phytoplankton community as a factor regulating rates of biomass specific photosynthesis, particularly when the successional changes involve species with very different life strategies.     

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.     

A new technique for estimating rates of carboxylation and electron transport in leaves of C3 plants for use in dynamic global vegetation models

The possible responses of the terrestrial biosphere to future CO₂ increases and associated climatic change are being investigated using dynamic global vegetation models (DG VMs) which include the Farquhar et al. (1980) biochemical model of leaf assimilation as the primary means of carbon capture. This model requires representative values of the maximum rates of Rubisco activity, V_max, and electron transport, J_max, for different vegetation types when applied at the global scale. Here, we describe an approach for calculating these values based on measurements of the maximum rate of leaf photosynthesis (A_max) ¹³C discrimination. The approach is tested and validated by comparison with measurements of Rubisco activity assayed directly on wild-type and transgenic Nicotiana tabacum (tobacco) plants with altered Rubisco activity grown under ambient and elevated CO₂ mole fractions with high and low N-supply. V_max and J_max values are reported for 18 different vegetation types with global coverage. Both variables were linearly related reinforcing the idea of optimal allocation of resources to photosynthesis (light harvesting vs. Rubisco) at the global scale. The reported figures should be of value to the further development of vegetation and ecosystem models employing mechanistic DGVMs.     

Brassinosteroids promote photosynthesis and growth by enhancing activation of Rubisco and expression of photosynthetic genes in <Emphasis Type="Italic">Cucumis sativus</Emphasis>

Brassinosteroids (BRs) are a new group of plant growth substances that promote plant growth and productivity. We showed in this study that improved growth of cucumber (Cucumis sativus) plants after treatment with 24-epibrassinolide (EBR), an active BR, was associated with increased CO₂ assimilation and quantum yield of PSII (Φ_PSII). Treatment of brassinazole (Brz), a specific inhibitor for BR biosynthesis, reduced plant growth and at the same time decreased CO₂ assimilation and Φ_PSII. Thus, the growth-promoting activity of BRs can be, at least partly, attributed to enhanced plant photosynthesis. To understand how BRs enhance photosynthesis, we have analyzed the effects of EBR and Brz on a number of photosynthetic parameters and their affecting factors, including the contents and activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco). Northern and Western blotting demonstrated that EBR upregulated, while Brz downregulated, the expressions of rbcL, rbcS and other photosynthetic genes. In addition, EBR had a positive effect on the activation of Rubisco based on increased maximum Rubisco carboxylation rates (V _c,max), total Rubisco activity and, to a greater extent, initial Rubisco activity. The accumulation patterns of Rubisco activase (RCA) based on immunogold-labeling experiments suggested a role of RCA in BR-regulated activation state of Rubisco. Enhanced expression of genes encoding other Calvin cycle genes after EBR treatment may also play a positive role in RuBP regeneration (J _max), thereby increasing maximum carboxylation rate of Rubisco (V _c,max). Thus, BRs promote photosynthesis and growth by positively regulating synthesis and activation of a variety of photosynthetic enzymes including Rubisco in cucumber.     

Alterations in Rubisco activity and in stomatal behavior induce a daily rhythm in photosynthesis of aerial leaves in the amphibious-plant Nuphar lutea

Nuphar lutea is an amphibious plant with submerged and aerial foliage, which raises the question how do both leaf types perform photosynthetically in two different environments. We found that the aerial leaves function like terrestrial sun-leaves in that their photosynthetic capability was high and saturated under high irradiance (ca. 1,500 μmol photons m⁻² s⁻¹). We show that stomatal opening and Rubisco activity in these leaves co-limited photosynthesis at saturating irradiance fluctuating in a daily rhythm. In the morning, sunlight stimulated stomatal opening, Rubisco synthesis, and the neutralization of a night-accumulated Rubisco inhibitor. Consequently, the light-saturated quantum efficiency and rate of photosynthesis increased 10-fold by midday. During the afternoon, gradual closure of the stomata and a decrease in Rubisco content reduced the light-saturated photosynthetic rate. However, at limited irradiance, stomatal behavior and Rubisco content had only a marginal effect on the photosynthetic rate, which did not change during the day. In contrast to the aerial leaves, the photosynthesis rate of the submerged leaves, adapted to a shaded environment, was saturated under lower irradiance. The light-saturated quantum efficiency of these leaves was much lower and did not change during the day. Due to their low photosynthetic affinity for CO₂ (35 μM) and inability to utilize other inorganic carbon species, their photosynthetic rate at air-equilibrated water was CO₂-limited. These results reveal differences in the photosynthetic performance of the two types of Nuphar leaves and unravel how photosynthetic daily rhythm in the aerial leaves is controlled.     

Photosynthetic activity of phytoplankton and its relation to environmental factors in hypersaline Mono Lake,California

The photosynthetic activity of phytoplankton in hypersaline Mono Lake, California was measured over the three year period, 1983–1985. The maximum chlorophyll-specific rate of carbon uptake (P_m ^B) and the light-limited slope (alpha) were derived from laboratory measurements of photosynthesis vs. irradiance (P-I) relationships. Annual estimates of primary production were 340–540 g C m^-2 yr^-1. Production was two to three times higher during the spring of 1983 than in the springs of 1984 and 1985; higher standing biomass of algae occurred in 1983. While P_m ^B rates followed water temperatures and varied over 40-fold over the year, integral primary production varied less since periods of high P_m ^B occurred when algal biomass was low. Sixty-eight percent of the seasonal variation in the P_m ^B was explained by a regression on temperature (53%), chlorophyll a (12%), and the carbon:chlorophyll a ratio (3%). Light-saturated and light-limited rates of photosynthesis generally covaried, evidenced by the strong seasonal correlation between P_m ^B and alpha. Sixty-one percent of variation in alpha was explained by a regression on P_m ^B, temperature, grazing, water column stability, and self-shading. There was no correlation of carbon uptake with ambient levels of inorganic nitrogen. The regression coefficient of the dependence of P_m ^B on the seasonal temperature trend was much larger than that determined from individual samples incubated at several different temperatures; this indicates that uptake is limited by more than low temperatures in the spring. Regression equations including only temperature, chlorophyll and depth were sufficient to estimate patterns of seasonal and year to year variation in integral primary productivity.     

Effects of Cerium on Key Enzymes of Carbon Assimilation of Spinach Under Magnesium Deficiency

The mechanism of the fact that cerium improves the photosynthesis of plants under magnesium deficiency is poorly understood. The main aim of the study was to determine the role of cerium in the amelioration of magnesium deficiency effects in CO₂ assimilation of spinach. Spinach plants were cultivated in Hoagland’s solution. They were subjected to magnesium deficiency and to cerium chloride administered in the magnesium-present Hoagland’s media and magnesium-deficient Hoagland’s media. The results showed that the chlorophyll synthesis and oxygen evolution was destroyed, and the activities of Rubisco carboxylasae and Rubisco activase and the expression of Rubisco large subunit (rbcL), Rubisco small subunit (rbcS), and Rubisco activase subunit (rca) were significantly inhibited, then plant growth was inhibited by magnesium deficiency. However, cerium promotes the chlorophyll synthesis, the activities of two key enzymes in CO₂ assimilation, and the expression of rbcL, rbcS, and rca, thus leading to the enhancement of spinach growth under magnesium-deficient conditions.     

PHOTOSYNTHETIC RESPONSE OF NATURAL ASSEMBLAGES OF MARINE BENTHIC MICROALGAE TO SHORT-AND LONG-TERM VARIATIONS OF INCIDENT IRRADIANCE IN BAFFIN BAY,TEXAS1

This study was designed to understand the high variability characterizing primary production rates of microphytobenthos. The photosynthetic efficiency (α^B) and photosynthetic capacity (P^B_max) of the microphytobenthos were measured at different times of the day on two different dates (8 May and 7 July 1990). In July, unusually low light conditions were caused by the development of a brown tide (chrysophytes). Both light-limited and light-saturated photosynthesis changed at hourly and monthly scales. There was a linear relationship between α^B and P^B_max, suggesting a common response to environmental factors [α^B= 0.0075(±0.00063)·P^B_max+ 0.00097(±0.0071), R²= 0.94]. Incident irradiance at the sediment-water interface was the primary physical factor that explained variability of both α^B (84%) and P^B_max (92%). Temperature had a negative but minor effect that explained an extra 8% and 2% of the variance, respectively. There was no diel rhythm of α^B and P^B_max and incident irradiance was regulated by wind-induced currents. Therefore, microphytobenthos photosynthesis seemed to be primarily controlled by wind events in Baffin Bay.     

Widespread positive selection in the photosynthetic Rubisco enzyme

Background  

Rubisco enzyme catalyzes the first step in net photosynthetic CO₂ assimilation and photorespiratory carbon oxidation and is responsible for almost all carbon fixation on Earth. The large subunit of Rubisco is encoded by the chloroplast rbcL gene, which is widely used for reconstruction of plant phylogenies due to its conservative nature. Plant systematicists have mainly used rbcL paying little attention to its function, and the question whether it evolves under Darwinian selection has received little attention. The purpose of our study was to evaluate how common is positive selection in Rubisco among the phototrophs and where in the Rubisco structure does positive selection occur.     

Primary production of epipsammic algal communities in Lake Balaton (Hungary)

Benthic algal communities can play an important role in matter and energy flux of shallow lakes. Their contribution to total primary production of lakes has been largely unexplored. The aim of this study was to estimate the primary production of the epipsammic algal communities at different water depths in Lake Balaton (Hungary) with photosynthetic measurements performed in laboratory. The photosynthesis of the benthic algae of different origin was studied at nine different irradiance levels, in three replicates. The maximum photosynthetic rate (P _max) was always higher in samples from the shallow parts than those from the deeper regions of the lake. Along the west–east longitudinal axis of the lake P _max decreased in the southern part and increased in the middle of the lake as a consequence of differences in the chlorophyll-a concentrations. Knowing P _max, I _k, global radiation and extinction coefficient, the primary production (mg C m⁻² day⁻¹) of the epipsammic algal community was calculated at different water depths. In the shallow regions at 0.5 and 1 m water depth 75–95% and 60–85% of the production was attributable to the epipsammon. The percentage contribution of epipsammon was at 2 m water depth 20–65%. In the deeper pelagic region (>3 m) more than 85% of the primary production originated from the phytoplankton.     

Unimodal size scaling of phytoplankton growth and the size dependence of nutrient uptake and use

Phytoplankton size structure is key for the ecology and biogeochemistry of pelagic ecosystems, but the relationship between cell size and maximum growth rate (μ_max) is not yet well understood. We used cultures of 22 species of marine phytoplankton from five phyla, ranging from 0.1 to 10⁶ μm³ in cell volume (V_cell), to determine experimentally the size dependence of growth, metabolic rate, elemental stoichiometry and nutrient uptake. We show that both μ_max and carbon‐specific photosynthesis peak at intermediate cell sizes. Maximum nitrogen uptake rate (V_maxN) scales isometrically with V_cell, whereas nitrogen minimum quota scales as V_cell^0.84. Large cells thus possess high ability to take up nitrogen, relative to their requirements, and large storage capacity, but their growth is limited by the conversion of nutrients into biomass. Small species show similar volume‐specific V_maxN compared to their larger counterparts, but have higher nitrogen requirements. We suggest that the unimodal size scaling of phytoplankton growth arises from taxon‐independent, size‐related constraints in nutrient uptake, requirement and assimilation.     

Spatial and temporal variation of photosynthetic parameters in natural phytoplankton assemblages in the Beaufort Sea, Canadian Arctic

During summer 2008, as part of the Circumpolar Flaw Lead system study, we measured phytoplankton photosynthetic parameters to understand regional patterns in primary productivity, including the degree and timescale of photoacclimation and how variability in environmental conditions influences this response. Photosynthesis–irradiance measurements were taken at 15 sites primarily from the depth of the subsurface chlorophyll a (Chl a) maximum (SCM) within the Beaufort Sea flaw lead polynya. The physiological response of phytoplankton to a range of light levels was used to assess maximum rates of carbon (C) fixation (P _m^*), photosynthetic efficiency (α ^*), photoacclimation (E _k), and photoinhibition (β ^*). SCM samples taken along a transect from under ice into open water exhibited a >3-fold increase in α ^* and P _m^*, showing these parameters can vary substantially over relatively small spatial scales, primarily in response to changes in the ambient light field. Algae were able to maintain relatively high rates of C fixation despite low light at the SCM, particularly in the large (>5 μm) size fraction at open water sites. This may substantially impact biogenic C drawdown if species composition shifts in response to future climate change. Our results suggest that phytoplankton in this region are well acclimated to existing environmental conditions, including sea ice cover, low light, and nutrient pulses. Furthermore, this photoacclimatory response can be rapid and keep pace with a developing SCM, as phytoplankton maintain photosynthetic rates and efficiencies in a narrow “shade-acclimated” range.     

Leaf carbon gain simulation for tree seedlings acclimatized to microsite light regimes of a temperate pine forest

Leaf carbon gain simulation was performed forQuercus serrata seedlings with previously reported 6 day photosynthetic photon flux density (PPFD) histograms from 20 understorey microsites of a pine forest (Washitani & Tang 1991). This simulation was performed with or without an assumption of the acclimatization of photosynthetic capacity (P_max) to microsite light availability. A constant ratio of respiration rate to P_max, within, the range of 0.07–0.1, was assumed as a constraint. In relatively well illuminated microsites with a diffuse site factor above 0.1, predicted optimal P_max was about 5 μmol m⁻² s⁻¹, with the predicted mean daily net carbon gain being about 50 mmol m⁻² day⁻¹. Each of the predicted optimal P_max and the simulated mean daily net carbon gains with a constant P_max (5 μmol m⁻² s⁻¹) or the predicted optimal P_max was linearly related to the microsite light availability index, diffuse site factor. Simulated net carbon gain was negative at diffuse site factors below 0.04, if the constant of P_max was assumed. The predicted linear relationship between net carbon gain and diffuse site factor could provide an ecophysiological basis for the observed linear dependency of the relative growth rate of biomass ofQ. serrata seedlings on the microsite diffuse site factor (Washitani & Tang 1991).     

Phytoplankton biomass and production during late austral spring (1991) and summer (1993) in the Bransfield Strait

Phytoplankton biomass and productivity were measured during two cruises in the Bransfield Strait in December 1991 (D91) and January/February 1993 (J93). Strong seasonal variability in productivity values was observed due to differences in the physiological response of phytoplankton. However, although the photosynthetic capacity of phytoplankton was markedly lower in D91 [P _m ^B =0.61 ± 0.25 mg C (mg Chla)⁻¹ h⁻¹] than in J93 [P _m ^B =2.18 ± 0.91 mg C (mg Chla)⁻¹ h⁻¹], average water column chlorophyll values in different areas of the strait were approximately similar in D91 (49–78 mg Chla m⁻²) and J93 (22–76 mg Chla m⁻²). The spatial distribution of chlorophyll was patchy and generally associated with the influence of the different water masses that meet together in the Bransfield Strait. No correlation was found between the mixed layer depth and either the integrated chlorophyll or the productivity. Our results suggest that major phytoplankton blooms in the Bransfield Strait are advected from the nearby Gerlache Strait or Bellingshausen Sea following the main eastward surface currents. Accepted: 5 July 1998     

Stomatal development and associated photosynthetic performance of capsicum in response to differential light availabilities

The mechanisms of capsicum growth in response to differential light availabilities are still not well elucidated. Hereby, we analyzed differential light availabilities on the relationship between stomatal characters and leaf growth, as well as photosynthetic performance. We used either 450–500 μmol m⁻² s⁻¹ as high light (HL) or 80–100 μmol m⁻² s⁻¹ as low light (LL) as treatments for two different cultivars. Our results showed that the stomatal density (SD) and stomatal index (SI) increased along with the leaf area expansion until the peak of the correlation curve, and then decreased. SD and SI were lower under the LL condition after three days of leaf expansion. For both cultivars, downregulation of photosynthesis and electron transport components was observed in LL-grown plants as indicated by lower light- and CO₂-saturated photosynthetic rate (P _max and RuBP_max), quantum efficiency of photosystem II (PSII) photochemistry (Φ_PSII), electron transport rate (ETR) and photochemical quenching of fluorescence (q_p). The observed inhibition of the photosynthesis could be explained by the decrease of SD, SI, Rubisco content and by the changes of the chloroplast. The low light resulted in lower total biomass, root/shoot ratio, and the thickness of the leaf decreased. However, the specific leaf area (SLA) and the content of leaf pigments were higher in LL-treatment. Variations in the photosynthetic characteristics of capsicum grown under different light conditions reflected the physiological adaptations to the changing light environments.     

Photosynthesis and UV-B Tolerance of the Marine Alga Fucus vesiculosus at Different Sea Water Salinities

The marine algal species in the Baltic Sea are few due to the low sea water salinity. One of the few species that can be found is Fucus vesiculosus. Even this species is affected by the low salinity and becomes smaller in size in the Baltic. In present work the photosynthesis of F. vesiculosus in the northern Baltic (Bothnian Sea) was compared to the photosynthesis of F. vesiculosus in the Atlantic. Oxygen evolution was measured before and after exposure to 2.3 W of UV-B (280–320 nm) radiation for 5 h, as well as after 48 h recovery in low light. The plants were kept in their own sea water salinity as well as in a changed salinity, this to examine possible correlations between salinity and photosynthesis. The results show a significant higher initial maximal photosynthesis (P _max) for Atlantic plants (10.3 nmol O₂ g⁻¹ FW s⁻¹) compared to Baltic plants (4.0 nmol O₂ g⁻¹ FW s⁻¹). The Baltic plants were found more sensitive to UV-B with a 40–50% decrease of P _max as well as a lower degree of recovery (60–70% compared to 75–95% for the Atlantic plants). The higher salinity (35 psu) had a positive effect on the Baltic F. vesiculosus with increased P _max as well as increased tolerance to UV-B. The lower salinity (5 psu) had a negative effect on the Atlantic plants with a decreased P _max as well as a lower tolerance to UV-B. Pigment content was found higher in Atlantic F. vesiculosus. The pigment content decreased then the Atlantic plants were transferred to 5 psu. The concentration of Chl a as well as the total content of violaxanthin, diadinoxanthin and zeaxanthin in Baltic plants increased when transferred to 35 psu. The Atlantic F. vesiculosus can not survive the low salinity in the northern Baltic (died within 8 weeks). It is likely that a long time acclimation or adaptation to low salinity has taken place for F. vesiculosus in northern Baltic. If this is an ecotypic or genotypic development it is too early to say.     

A comparative analysis of photosynthetic characteristics of hulless barley at two altitudes on the Tibetan Plateau

To determine the photosynthetic characteristics of C₃ plants and their sensitivity to CO₂ at different altitudes on the Tibetan Plateau, hulless barley (Hordeum vulgare L. ssp. vulgare) was grown at altitudes of 4,333 m and 3,688 m. Using gas-exchange measurements, photosynthetic parameters were simulated, including the maximum net photosynthesis (P _max) and the apparent quantum efficiency (α). Plants growing at higher altitude had higher net photosynthetic rates (P _N), photosynthesis parameters (P _max and α) and sensitivities to CO₂ enhancement than plants growing at lower altitude on the Tibetan Plateau. The enhancements of P _N, P _max, and α for plants growing at higher altitude, corresponding with 10 μmol(CO₂) mol⁻¹ increments, were approximately 0.20∼0.45%, 0.05∼0.20% and 0.12∼0.36% greater, respectively, than for plants growing at lower altitude, respectively, where CO₂ levels rose from 10 to 170 μmol(CO₂) mol⁻¹. Therefore, on the Tibetan Plateau, the changes in the photosynthetic capacities and the photosynthetic sensitivities to CO₂ observed in the C₃ plants grown above 3,688 m are likely to increase with altitude despite the decreasing CO₂ partial pressure.     

Localization of photosynthetic metabolism in the parasitic angiosperm Cuscuta reflexa

Cells capable of photosynthesis in the parasitic angiosperm Cuscuta reflexa Roxb. (dodder) are highly localized. Immunolocalization of ribulose-1,5 bisphosphate carboxylase-oxygenase (Rubisco) and autofluorescence of chlorophyll in transverse sections of stems showed that they were largely restricted to a band of cells adjacent to the vascular bundles, consequently, the concentrations of Rubisco and chlorophyll were low per unit area or fresh weight. When ¹⁴CO₂ was supplied to stem segments of C. reflexa it preferentially accumulated in these cells adjacent to the vasculature. Although the conductance for CO₂ movement to the cells containing chlorophyll and Rubisco was very low, both the light reactions and dark reactions of photosynthesis appeared to be functional. De-epoxidation of the xanthophyll-cycle pigments after exposure to high light, and the chlorophyll fluorescence parameters, photochemical quenching (qP), non-photochemical quenching (NPQ) and the quantum efficiency of photosystem II (φPSII) responded normally to changes in photon flux density, indicating functional light-driven electron transport. The response of CO₂ exchange to photon flux density followed a typical hyperbolic curve, and positive rates of CO₂ fixation occurred when external CO₂ was increased to 5%. We propose that CO₂ for carbon assimilation is derived from internally respired CO₂ and that this layer of photosynthetic cells makes a positive contribution to the carbon budget of C. reflexa. Received: 23 October 1997 / Accepted: 16 December 1997     

Intra- and inter-specific variation in canopy photosynthesis in a mixed deciduous forest

 Within the same forest, photosynthesis can vary greatly among species and within an individual tree. Quantifying the magnitude of variation in leaf-level photosynthesis in a forest canopy will improve our understanding of and ability to model forest carbon cycling. This information requires extensive sampling of photosynthesis in the canopy. We used a 22-m-tall, four-wheel-drive aerial lift to reach five to ten leaves from the tops of numerous individuals of several species of temperate deciduous trees in central Massachusetts. The goals of this study were to measure light-saturated photosynthesis in co-occurring canopy tree species under field conditions, and to identify sampling schemes appropriate for canopy tree studies with challenging logistics. Photosynthesis differed significantly among species. Even though all leaves measured were canopy-top, sun-acclimated foliage, the more shade-tolerant species tended to have lower light-saturated photosynthetic rates (P _max) than the shade-intolerant species. Likewise, leaf mass per area (LMA) and nitrogen content (N) varied significantly between species. With only one exception, the shade-tolerant species tended to have lower nitrogen content on an area basis than the intolerant species, although the LMA did not differ systematically between these ecological types. Light-saturated P _max rates and nitrogen content, both calculated on either an area or a mass basis, and the leaf mass to area ratio, significantly differed not only among species, but also among individuals within species (P<0.0001 for both). Differences among species accounted for a greater proportion of variance in the P _max rates and the nitrogen content than the differences among individuals within a species (58.5–78.8% of the total variance for the measured parameters was attributed to species-level differences versus 5.5–17.4% of the variance was attributed to differences between individual trees of a given species). Furthermore, more variation is accounted for by differences among leaves in a single individual tree, than by differences among individual trees of a given species (10.7–30.4% versus 5.5–17.4%). This result allows us to compare species-level photosynthesis, even if the sample size of the number of trees is low. This is important because studies of canopy-level photosynthesis are often limited by the difficulty of canopy access. As an alternative to direct canopy access measurements of photosynthesis, it would be useful to find an ”easy-to-measure” proxy for light-saturated photosynthetic rates to facilitate modeling forest carbon cycling. Across all species in this study, the strongest correlation was between nitrogen content expressed on an area basis (mmol m^–2, N _area) and light-saturated P _max rate (μmol m^–2 s^–1, P _maxarea) (r ²=0.511). However, within a given species, leaf nitrogen was not tightly correlated with photosynthesis. Our sampling design minimized intra-specific leaf-level variation (i.e., leaves were taken only from the top of the canopy and at only one point in the season). This implies that easy-to-measure trends in nitrogen content of leaves may be used to predict the species-specific light-saturated P _max rates. Received: 16 March 1996 / Accepted: 16 August 1996     

Exogenous sucrose can decrease <Emphasis Type="Italic">in vitro</Emphasis> photosynthesis but improve field survival and growth of coconut (<Emphasis Type="Italic">Cocos nucifera</Emphasis> L.) <Emphasis Type="Italic">in vitro</Emphasis> plantlets

Summary Coconut (Cocos nucifera L.) plantlets grown in vitro often grow slowly when transferred to the field possibly, due to a limited photosynthetic capacity of in vitro-cultured plantlets, apparently caused by the sucrose added to growth medium causing negative feedback for photosynthesis. In this paper, we tested the hypothesis that high exogenous sucrose will decrease ribulose 1,5-bisphosphate carboxylase (Rubisco) activity and photosynthesis resulting in limited ex vitro growth. Plantlets grown with high exogenous sucrose (90 gl⁻¹) had reduced photosynthetic activity that resulted in a poor photosynthetic response to high levels of light and CO₂. These plantlets also had low amounts of Rubisco protein, low Rubisco activity, and reduced growth despite showing high survival when transferred to the field. Decreasing the medium’s sucrose concentration from 90 to 22.5 gl⁻¹ or 0 gl⁻¹ resulted in increased photosynthetic response to light and CO₂ along with increased Rubisco and phosphoenolpyruvate carboxylase (PEPC) activities and proteins. However, plantlets grown in vitro without exogenous sucrose died when transferred ex vitro, whereas those grown with intermediate exogenous sucrose showed intermediate photosynthetic response, high survival, fast growth, and ex vitro photosynthesis. Thus, exogenous sucrose at moderate concentration decreased photosynthesis but increased survival, suggesting that both in vitro photosynthesis and exogenous sucrose reserves contribute to field establisment and growth of coconut plantlets cultured in vitro.