Adaptations of tropical marine microphytobenthic assemblages along a gradient of light and nutrient availability in Suva Lagoon,Fiji

How are microphytobenthic biofilms adapted to the high incident irradiances and temperatures, low inorganic nutrient concentrations and high desiccation stresses on intertidal flats present in tropical environments? This study investigated biofilms subject to different environmental conditions in a range of tropical sites in Suva lagoon, Fiji. PAM fluorescence was used to measure photophysiological responses to the light climate. Biofilm colloidal carbohydrate, extracellular polymeric substances (EPS) and low molecular weight (MW) carbohydrate concentrations and diel carbohydrate production patterns were measured. Average biomass (Chl a) ranged from 15 to 36?mg?m^?2, and was highest in seagrass bed sediments, but biomass was not correlated with water column or sediment porewater nutrient concentrations. Biofilm photophysiology differed significantly along a combined gradient of light and nutrient availability, with F _v/F _m, relative ETR_max and E _k of biofilms highest in mangrove and intertidal main island sites and lowest in subtidal coral reef flats. Subtidal biofilms showed photoinhibition at irradiances > 1000?µmol?m^?2. Significant correlations between Chl a and colloidal carbohydrate concentrations were present (except on intertidal sandflats), and tropical biofilms had higher ratios of colloidal carbohydrate and EPS to Chl a than temperate estuarine biofilms, probably due to a combination of high irradiance and low nutrient availability leading to the production of excess photoassimilates. The percentage of EPS present in the colloidal fraction was highest in coral sand biofilms (42%), which had the lowest nutrient concentrations, compared with other sites (25–32%). Intertidal biofilms predominantly consisted of large motile taxa and showed strong rhythms of vertical migration. During tidal emersion, high sediment temperatures (41?°C), irradiance (>2300?µmol?m^?2?s^?1) and salinity (49‰) stimulated downward migration. In silty sediments, migration resulted in a reduction in photosynthetic activity during the midday period but, in sands with high light penetration (to a depth of > 1700?µm), high production rates of EPS (18.2?µg carbo. µg Chl a^?1 h^?1) and low MW carbohydrate exudates (40.2?µg carbo. µg Chl a^?1 h^?1) occurred. Vertical migration, high E _k and high rates of photoassimilate dumping are all adaptations to living in the tropical intertidal zone. Seagrass and reef flat biofilms consisted of a diverse non-migratory flora of motile and non-motile taxa that were not subject to such extreme temperature and irradiance conditions. Low values of photosynthetic parameters and high colloidal and EPS content indicated that these biofilms were nutrient-limited.     

Carbon translocation from symbiont to host depends on irradiance and food availability in the tropical coral Stylophora pistillata

Reef-building corals live in symbiosis with dinoflagellates that translocate a large proportion of their photosynthetically fixed carbon compounds to their coral host for its own metabolism. The carbon budget and translocation rate, however, vary depending on environmental conditions, coral host species, and symbiont clade. To quantify variability in carbon translocation in response to environmental conditions, this study assessed the effect of two different irradiance levels (120 and 250 μmol photons m^?2 s^?1) and feeding regimes (fed with Artemia salina nauplii and unfed) on the carbon budget of the tropical coral Stylophora pistillata. For this purpose, H¹³CO₃ ^?-enriched seawater was used to trace the conversion of photosynthetic carbon into symbiont and coral biomass and excrete particulate organic carbon. Results showed that carbon translocation (ca. 78 %) and utilization were similar under both irradiance levels for unfed colonies. In contrast, carbon utilization by fed colonies was dependent on the growth irradiance. Under low irradiance, heterotrophy was accompanied by lower carbon translocation (71 %), higher host and symbiont biomass, and higher calcification rates. Under high irradiance, heterotrophy was accompanied by higher rates of photosynthesis, respiration, and carbon translocation (90 %) as well as higher host biomass. Hence, levels of resource sharing within coral–dinoflagellate symbioses depend critically on environmental conditions.     

Effects of temperature,irradiance, and daylength on the marine diatom leptocylindrus danicus cleve. I. Photosynthesis and cellular composition

Rates of ¹⁴C uptake and cellular composition of C, N, and Chl a in the marine diatom Leptocylindrus danicus Cleve were measured in axenic batch culture under 49 combinations of temperature (5, 10, 15, 20 °C), daylength (15: 9, 12: 12, 9: 15 LD), and irradiance (at least four irradiances per daylength). ¹⁴C uptake exhibited a temperature-dependent daylength effect. Similar P-I curves characterized cells grown under 15: 9 and 12: 12 LD; P_max values were 17.2, 11.2, 4.3, and 1.8 pg C. pg Chl a^?1. h^?1 at 20, 15, 10, and 5°C, respectively. Under 9:15 LD at 20 and 15°C, the lightsaturated photosynthetic rate was ≈50% that in cells grown under longer daylengths. ¹⁴C uptake was independent of daylength at 10 and 5°C. The initial slope, a, of cells grown under long daylengths increased by five-fold between 5 and 20 °C. α values of cells grown under 9: 15 LD at 15 and 20 °C were depressed relative to longer daylengths. Chl a was inversely related to irradiance, and increased with temperature from 10 to 20 °C, whereas cell carbon and nitrogen showed a similar temperature dependence but was not influenced by irradiance or daylength. The C : N ratio and cell volume were independent of temperature, irradiance, and daylength. Both the C : Chl a and N : Chl a ratios increased with irradiance by greater amounts at lower temperatures.     

Seasonal variation in pigmentation of the dinoflagellate Peridinium gatunense (Dinophyceae) in Lake Kinneret, Israel

1. Pigment composition was measured in natural phytoplankton samples from Lake Kinneret, Israel. From March through June 1998, the dinoflagellate Peridinium gatunense Nygaard mostly contributed more than 95% of the algal biomass. Peak densities were found in April, close to the water surface, with >10⁹ cells m^?3, chlorophyll (Chl) a concentration of 380 mg m^?3 and areal Chl‐a density of >1300 mg m^?2. 2. Cellular concentrations of Chl‐a changed between 201 and 282 pg cell^?1, but did not show a defined temporal fluctuation. 3. The mass ratio of Chl‐c to Chl‐a changed from March to June between 0.16 and 0.22, and the peridinin to Chl‐a ratio changed from 0.25 to 0.41. Neither ratio showed a clear pattern of seasonal change. Conversely, there was a progressive increase in diadinoxanthin and β‐carotene ratios to Chl‐a through the season, parallel to the increase in photon flux impinging upon the lake surface. The diadinoxanthin to Chl‐a ratio changed from 0.11 to 0.28 and the β‐carotene to Chl‐a ratio varied from 0.03 to 0.08 from March through June. 4. Diatoxanthin was not detected in natural samples. However, it was present in experiments with P. gatunense cultures, when concentration of diatoxanthin increased rapidly, concurrent with a decrease in diadinoxanthin and β‐carotene concentrations, while Chl‐c and peridinin ratios to Chl‐a were almost stable with photon flux increase. 5. The seasonal variation in cellular pigmentation of P. gatunense in Lake Kinneret suggests that accumulation of photoprotective pigments is essential for optimisation of photosynthetic activity of this large dinoflagellate.     

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

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

Combined effects of light and temperature on growth, photosynthesis, and pigment content in the mat-forming cyanobacterium Geitlerinema amphibium

Geitlerinema amphibium (BA-13), mat-forming cyanobacterium from the southern Baltic Sea, was grown at three irradiances [5, 65, and 125 μmol(photon) m^?2 s^?1] and three temperatures (15, 22.5, and 30°C). To determine the effect of the investigated factors and their interaction on culture concentration, pigment content, and photosynthetic parameters of cyanobacterium, factorial experiments and two-way analysis of variance (ANOVA) were carried out. Both chlorophyll (Chl) a and phycobilins (PB) were influenced by the irradiance and temperature, but stronger effect was noted in the case of the former one. Chl a and PB concentration per 100 μm of filament dropped above 4-fold with the increasing irradiance. The ratios between individual carotenoids [β-carotene, zeaxanthin, and myxoxanthophyll (Myx)] and Chl a increased significantly with an increase in the irradiance. The greatest fluctuations were observed in the ratio of Myx to Chl a (above 10-fold). Thus, Myx was suggested as the main photoprotective carotenoid in G. amphibium. Based on photosynthetic light response (PI) curves, two mechanisms of photoacclimation in G. amphibium were recognized: a change of photosynthetic units (PSU) number and a change of PSU size. These two mechanisms constituted the base of significant changes in photosynthetic rate and its parameters, such as the compensation point (P _C), the initial slope of photosynthetic curve (α), saturation irradiance (E _K), maximal photosynthetic rate (P _max), and dark respiration rate (R _D). The greatest changes were observed in P _C values (about 15-fold within the range of the factors tested). Studied parameters showed a wide range of changes, which might indicate G. amphibium ability to acclimatize well to irradiance and temperature, and indirectly might explain the successful growth of cyanobacterium in dynamically changing environmental conditions.     

AMMONIUM AND UV RADIATION STIMULATE THE ACCUMULATION OF MYCOSPORINE‐LIKE AMINO ACIDS IN PORPHYRA COLUMBINA (RHODOPHYTA) FROM PATAGONIA,ARGENTINA1

The combined effects of ammonium concentration and UV radiation on the red alga Porphyra columbina (Montagne) from the Patagonian coast (Chubut, Argentina) was determined using short‐term (less than a week) experimentation. Discs of P. columbina were incubated with three ammonium concentrations (0, 50, and 300 μM NH₄Cl) in anilluminated chamber (PAR=300 μmol photons·m^?2·s^?1, UVA=15 W·m^?2, UVB=0.7 W·m^?2) at 15°C. Algae incubated at 300 μM ammonium showed a significant increase (P<0.05) in the concentration of mycosporine‐like amino acids (MAAs) compared with the initial value or with the other ammonium treatments. The increase of MAAs was, however, a function of the quality of irradiance received, with a higher increase in samples exposed to UVA compared with UVB (29% and 5% increase, respectively). However, UVB radiation was more effective in inducing MAA synthesis per unit energy received by the algae. Samples exposed to PAR only had an intermediate increase in MAA concentration of 16%. Chl a concentration decreased through the incubation with the greatest decrease at high ammonium concentration. Phycobiliprotein (BP) decreased through time with the smallest decrease occurring at high ammonium concentration. Photoinhibition (as a decrease of optimal quantum yield) was significantly greater under nitrogen‐deprived conditions than that under replete ammonium levels. Maximal gross photosynthesis (GP_max), as oxygen evolution, and maximal electron transport rate (ETR_max), as chl fluorescence, increased with the ammonium concentration. Positive relationships between maximal GP or ETR and pigment ratios (BP/chl a and MAAs/chl a) and negative relationships with chl a concentration were found.     

Seasonal variation in light-shade adaptation of natural populations of the symbiotic sea anemone Aiptasiapulchella (Carlgren, 1943) in Hawaii

The productivity and biomass parameters of the symbiotic anemone Aiptasia pulchella (Carlgren, 1943) from a shaded mangrove lagoon (maximum summer irradiance of 100 μE m⁻² · s⁻¹) and a sunlit reef flat (maximum summer irradiance of 1400 μE · m⁻² · s⁻¹) were examined in Hawaii. Light-shade adaptation was evident in the summer populations (1981) but not observed during the fall (1982). In the summer, zooxanthellae from the lagoon A. pulchella (shade anemones) contained 2.97 pg Chl a cell ⁻¹ and those from the reef flat (sun anemones) contained 1.70 pg Chl a · cell⁻¹; but Chl a : c₂ ratios were 2.5 in zooxanthellae from both shade and sun anemones. During the fall, there were no significant differences in Chl a and c₂ of zooxanthellae (2.25 pg Chl a · cell⁻¹) in shade and sun anemones, but Chl a : c₂ ratios averaged 3.9. During both seasons, shade anemones were larger and contained higher densities of zooxanthellae than sun anemones. In addition to differences between shade and sun habitats, there was localized photoadaptation of zooxanthellae within individual anemones due to microhabitat variations in ambient irradiance. Growth rates of zooxanthellae in A. pulchella differed in shade and sun anemones. Specific growth rates for zooxanthellae in situ were the same for shade populations in both summer and fall (0.016 day⁻¹). However, zooxanthellae in sun anemones grew four times faster in the fall (0.033 day⁻¹) than during the summer (0.008 day⁻¹). These results suggest that growth of zooxanthellae in these anemones was independent of ambient irradiance. Photosynthesis-irradiance (P-I) responses of shade and sun anemones during the summer showed that shade anemones had greater photosynthetic efficiencies (α) but lower photosynthetic capacities (P_max) than sun anemones. Dark-respiration rates of sun anemones were twice those obtained with shade anemones. In the fall, these populations of anemones did not exhibit P-I responses characteristic of light-shade adaptation. Both α and P_max of shade and sun anemones were higher in the fall, indicating that zooxanthellae in A. pulchella adapted to seasonal reduction in irradiance.     

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.     

Effects of light intensity and temperature on the photosynthetic irradiance response curves and chlorophyll fluorescence in three picocyanobacterial strains of Synechococcus

Chrococcoid cyanobacteria of the genus Synechococcus are the important component of marine and freshwater ecosystems. Picocyanobacteria comprise even 80% of total cyanobacterial biomass and contribute to 50% of total primary cyanobacterial bloom production. Chlorophyll (Chl) fluorescence and photosynthetic light response (P-I) curves are commonly used to characterize photoacclimation of Synechococcus strains. Three brackish, picocyanobacterial strains of Synechococcus (BA-132, BA-124, BA-120) were studied. They were grown under 4 irradiances [10, 55, 100, and 145 μmol(photon) m^?2 s^?1] and at 3 temperatures (15, 22.5, and 30°C). Photosynthetic rate was measured by Clark oxygen electrode, whereas the Chl fluorescence was measured using Pulse Amplitude Modulation fluorometer. Based on P-I, two mechanisms of photoacclimation were recognized in Synechococcus. The maximum value of maximum rate of photosynthesis (P _max) expressed per biomass unit at 10 μmol(photon) m^?2 s^?1 indicated a change in the number of photosynthetic units (PSU). The constant values of initial slope of photosynthetic light response curve (α) and the maximum value of P _max expressed per Chl unit at 145 μmol(photon) m^?2 s^?1 indicated another mechanism, i.e. a change in PSU size. These two mechanisms caused changes in photosynthetic rate and its parameters (compensation point, α, saturation irradiance, dark respiration, P _max) upon the influence of different irradiance and temperature. High irradiance had a negative effect on fluorescence parameters, such as the maximum quantum yield and effective quantum yield of PSII photochemistry (φ_PSII), but it was higher in case of φ_PSII.     

Ozone treatment affects pigment precursor metabolism in pine seedlings

Five‐week‐old seedlings of Pinus halepensis Mill. and Pinus brutia Ten. were exposed to air polluted with ozone (O₃) (250 nl l^?1, 12 h day^?1 for 4 days) or to ambient air containing ca 10–20 nl l^?1 O₃, in the light (180 μmol m^?2 s^?1 photosynthetic photon flux density [PPFD], 12 h day^?1) and then fed for 24 h in the light (100 μmol m^?2 s^?1 PPFD) with various radioactive precursors of chlorophyll (Chl) and carotene biosynthesis: 5‐[4‐¹⁴C]‐aminolevulinic acid (¹⁴C‐ALA), l ‐[¹⁴C(U)]‐glutamic acid (¹⁴C‐Glu), or d ,l ‐[2‐¹⁴C]‐mevalonic acid (¹⁴C‐MVA). Pigments were then extracted from cotyledons and fully expanded needles. Chl a and carotene were separated by thin‐layer chromatography and high‐performance liquid chromatography and their specific activities were determined. ¹⁴C‐ALA and ¹⁴C‐Glu labels were incorporated into Chl a and carotene. Exposure to O₃ did not inhibit incorporation of ¹⁴C‐ALA into Chl a molecules, but hydrolysis of Chl a showed that O₃ inhibited phytol labelling of Chl a. Labelling of carotene was also inhibited by O₃, but not when ¹⁴C‐MVA was used as the label. These data suggest that O₃ treatment inhibits (directly or indirectly) the biosynthesis of isoprenoids from products of ALA and Glu metabolism in the plastid, but not from MVA in the cytosol. This inhibition was more prominent when ¹⁴C‐ALA was used as the label than when ¹⁴C‐Glu was the labelling precursor. A significant increase in pheophorbide a, a tetrapyrrole component of Chl a labelling, and a concomitant decrease in phytol labelling was observed following incubation of O₃‐treated pine seedlings with ¹⁴C‐ALA and ¹⁴C‐Glu. Stronger inhibition of carotene biosynthesis and activation of Chl a tetrapyrrole labelling by ¹⁴C‐ALA (in comparison with ¹⁴C‐Glu) indicated that exposure to O₃ inhibits the conversion of ALA to Glu as the first step in ALA catabolism. These results also suggested a more intensive Glu metabolism (in comparison with ALA) for carotene biosynthesis in the cytosol, as well as cooperation between two pathways of isopentenyl diphosphate biosynthesis.     

Chlorophyll fluorescence kinetics, photosynthetic activity, and pigment composition of blue-shade and half-shade leaves as compared to sun and shade leaves of different trees

The chlorophyll (Chl) fluorescence induction kinetics, net photosynthetic CO₂ fixation rates P _N, and composition of photosynthetic pigments of differently light exposed leaves of several trees were comparatively measured to determine the differences in photosynthetic activity and pigment adaptation of leaves. The functional measurements were carried out with sun, half-shade and shade leaves of seven different trees species. These were: Acer platanoides L., Ginkgo biloba L., Fagus sylvatica L., Platanus x acerifolia Willd., Populus nigra L., Quercus robur L., Tilia cordata Mill. In three cases (beech, ginkgo, and oak), we compared the Chl fluorescence kinetics and photosynthetic rates of blue-shade leaves of the north tree crown receiving only blue sky light but no direct sunlight with that of sun leaves. In these cases, we also determined in detail the pigment composition of all four leaf types. In addition, we determined the quantum irradiance and spectral irradiance of direct sunlight, blue skylight as well as the irradiance in half shade and full shade. The results indicate that sun leaves possess significantly higher mean values for the net CO₂ fixation rates P _N (7.8–10.7 μmol CO₂ m^?2 s^?1 leaf area) and the Chl fluorescence ratio R _Fd (3.85–4.46) as compared to shade leaves (mean P _N of 2.6–3.8 μmol CO₂ m^?2 s^?1 leaf area.; mean R _Fd of 1.94–2.56). Sun leaves also exhibit higher mean values for the pigment ratio Chl a/b (3.14–3.31) and considerably lower values for the weight ratio total chlorophylls to total carotenoids, (a + b)/(x + c), (4.07–4.25) as compared to shade leaves (Chl a/b 2.62–2.72) and (a + b)/(x + c) of 5.18–5.54. Blue-shade and half-shade leaves have an intermediate position between sun and shade leaves in all investigated parameters including the ratio F _v/F _o (maximum quantum yield of PS2 photochemistry) and are significantly different from sun and shade leaves but could not be differentiated from each other. The mean values of the Chl fluorescence decrease ratio R _Fd of blue-shade and half-shade leaves fit well into the strong linear correlation with the net photosynthetic rates P _N of sun and shade leaves, thus unequivocally indicating that the determination of the Chl fluorescence decrease ratio R _Fd is a fast and indirect measurement of the photosynthetic activity of leaves. The investigations clearly demonstrate that the photosynthetic capacity and pigment composition of leaves and chloroplasts strongly depend on the amounts and quality of light received by the leaves.     

The influence of light on copper‐limited growth of an oceanic diatom,Thalassiosira oceanica (Coscinodiscophyceae)

Thalassiosira oceanica (CCMP 1005) was grown over a range of copper concentrations at saturating and subsaturating irradiance to test the hypothesis that Cu and light were interacting essential resources. Growth was a hyperbolic function of irradiance in Cu‐replete medium (263 fmol Cu′ · L^?1) with maximum rates achieved at 200 μmol photons · m^?2 · s^?1. Lowering the Cu concentration at this irradiance to 30.8 fmol Cu′ · L^?1 decreased cellular Cu quota by 7‐fold and reduced growth rate by 50%. Copper‐deficient cells had significantly slower (P < 0.0001) rates of maximum, relative photosynthetic electron transport (rETR_max) than Cu‐sufficient cells, consistent with the role of Cu in photosynthesis in this diatom. In low‐Cu medium (30.8 fmol Cu′ · L^?1), growth rate was best described as a positive, linear function of irradiance and reached the maximum value measured in Cu‐replete cells when irradiance increased to 400 μmol photons · m^?2 · s^?1. Thus, at high light, low‐Cu concentration was no longer limiting to growth: Cu concentration and light interacted strongly to affect growth rate of T. oceanica (P < 0.0001). Relative ETR_max and Cu quota of cells grown at low Cu also increased at 400 μmol photons · m^?2 · s^?1 to levels measured in Cu‐replete cells. Steady‐state uptake rates of Cu‐deficient and sufficient cells were light‐dependent, suggesting that faster growth of T. oceanica under high light and low Cu was a result of light‐stimulated Cu uptake.     

Distribution of N, Rubisco and photosynthesis in Pinus pinaster and acclimation to light

The light–nitrogen hypothesis suggests canopy photosynthesis is maximized when there is a positive relationship between irradiance received by foliage, its nitrogen content (per unit area N_area), and maximum rate of photosynthesis (A_max). Relationships among relative irradiance and N_area, allocation of nitrogen within the photosynthetic apparatus to Rubisco and chlorophyll, and A_max were examined in Pinus pinaster Ait. needles up to 6 years of age. Measurements were made before bud break in August 1998, and in May 1999 after the first ‘winter’ rains. In August, N_area in P. pinaster needles decreased from 5·1 to 5·7 g m^?2 in sunlit 1‐year‐old needles to 2·3 g m^?2 in shaded 6‐year‐old needles. In May, N_area was 5–40% less but spatial trends were the same. At both sampling dates, A_max was less in old shaded needles compared with young sunlit needles, and was thus consistent with the light–nitrogen hypothesis. Relationships between N_area and A_max were positive at both dates yet varied in strength and form. Allocation of nitrogen within the photosynthetic apparatus was qualitatively consistent with acclimation to light (i.e. Rubisco/Chl decreased with shading), but quantitatively suboptimal with respect to photosynthesis owing to consistent over‐investment in Rubisco. This over‐investment increased with height in the canopy and was greater in May than in August.     

AUTOTROPHIC GROWTH AND PHOTOACCLIMATION IN KARLODINIUM MICRUM (DINOPHYCEAE) AND STOREATULA MAJOR (CRYPTOPHYCEAE)1

We compared autotrophic growth of the dinoflagellate Karlodinium micrum (Leadbeater et Dodge) and the cryptophyte Storeatula major (Butcher ex Hill) at a range of growth irradiances (E_g). Our goal was to determine the physiological bases for differences in growth–irradiance relationships between these species. Maximum autotrophic growth rates of K. micrum and S. major were 0.5 and 1.5 div.·d^?1, respectively. Growth rates were positively correlated with C‐specific photosynthetic performance (PP^C, g C·g C^?1·h^?1) (r²=0.72). Cultures were grouped as light‐limited (LL) and high‐light (HL) treatments to allow interspecific comparisons of physiological properties that underlie the growth–irradiance relationships. Interspecific differences in the C‐specific light absorption rate (E_a^C, mol photons·g C^?1·h^?1) were observed only among HL acclimated cultures, and the realized quantum yield of C fixation (φ_C(real.), mol C·mol photons^?1) did not differ significantly between species in either LL or HL treatments. The proportion of fixed C that was incorporated into new biomass was lower in K. micrum than S. major at each E_g, reflecting lower growth efficiency in K. micrum. Photoacclimation to HL in K. micrum involved a significant loss of cellular photosynthetic capacity (P_max^cell), whereas in S. major, P_max^cell was significantly higher in HL acclimated cells. We conclude that growth rate differences between K. micrum and S. major under LL conditions relate primarily to cell metabolism processes (i.e. growth efficiency) and that reduced chloroplast function, reflected in PP^C and photosynthesis–irradiance curve acclimation in K. micrum, is also important under HL conditions.     

Acclimation of photosynthetic capacity to irradiance in tree canopies in relation to leaf nitrogen concentration and leaf mass per unit area

The observation of acclimation in leaf photosynthetic capacity to differences in growth irradiance has been widely used as support for a hypothesis that enables a simplification of some soil‐vegetation‐atmosphere transfer (SVAT) photosynthesis models. The acclimation hypothesis requires that relative leaf nitrogen concentration declines with relative irradiance from the top of a canopy to the bottom, in 1 : 1 proportion. In combination with a light transmission model it enables a simple estimate of the vertical profile in leaf nitrogen concentration (which is assumed to determine maximum carboxylation capacity), and in combination with estimates of the fraction of absorbed radiation it also leads to simple ‘big‐leaf’ analytical solutions for canopy photosynthesis. We tested how forests deviate from this condition in five tree canopies, including four broadleaf stands, and one needle‐leaf stand: a mixed‐species tropical rain forest, oak (Quercus petraea (Matt.) Liebl), birch (Betula pendula Roth), beech (Fagus sylvatica L.) and Sitka spruce (Picea sitchensis (Bong.) Carr). Each canopy was studied when fully developed (mid‐to‐late summer for temperate stands). Irradiance (Q, µmol m^?2 s^?1) was measured for 20 d using quantum sensors placed throughout the vertical canopy profile. Measurements were made to obtain parameters from leaves adjacent to the radiation sensors: maximum carboxylation and electron transfer capacity (V_a, J_a, µmol m^?2 s^?1), day respiration (R_da, µmol m^?2 s^?1), leaf nitrogen concentration (N_m, mg g^?1) and leaf mass per unit area (L_a, g m^?2). Relative to upper‐canopy values, V_a declined linearly in 1 : 1 proportion with N_a. Relative V_a also declined linearly with relative Q, but with a significant intercept at zero irradiance (P < 0·01). This intercept was strongly related to L_a of the lowest leaves in each canopy (P < 0·01, r² = 0·98, n= 5). For each canopy, daily lnQ was also linearly related with lnV_a(P < 0·05), and the intercept was correlated with the value for photosynthetic capacity per unit nitrogen (PUN: V_a/N_a, µmol g^?1 s^?1) of the lowest leaves in each canopy (P < 0·05). V_a was linearly related with L_a and N_a(P < 0·01), but the slope of the V_a : N_a relationship varied widely among sites. Hence, whilst there was a unique V_a : N_a ratio in each stand, acclimation in V_a to Q varied predictably with L_a of the lowest leaves in each canopy. The specific leaf area, L_m(cm² g^?1), of the canopy‐bottom foliage was also found to predict carboxylation capacity (expressed on a mass basis; V_m, µmol g^?1 s^?1) at all sites (P < 0·01). These results invalidate the hypothesis of full acclimation to irradiance, but suggest that L_a and L_m of the most light‐limited leaves in a canopy are widely applicable indicators of the distribution of photosynthetic capacity with height in forests.     

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。     

Distribution and photobiology of <Emphasis Type="Italic">Symbiodinium</Emphasis> types in different light environments for three colour morphs of the coral <Emphasis Type="Italic">Madracis pharensis</Emphasis>: is there more to it than total irradiance?

The role of symbiont variation in the photobiology of reef corals was addressed by investigating the links among symbiont genetic diversity, function and ecological distribution in a single host species, Madracis pharensis. Symbiont distribution was studied for two depths (10 and 25 m), two different light habitats (exposed and shaded) and three host colour morphs (brown, purple and green). Two Symbiodinium genotypes were present, as defined by nuclear internal transcribed spacer 2 ribosomal DNA (ITS2-rDNA) variation. Symbiont distribution was depth- and colour morph-dependent. Type B15 occurred predominantly on the deeper reef and in green and purple colonies, while type B7 was present in shallow environments and brown colonies. Different light microhabitats at fixed depths had no effect on symbiont presence. This ecological distribution suggests that symbiont presence is potentially driven by light spectral niches. A reciprocal depth transplantation experiment indicated steady symbiont populations under environment change. Functional parameters such as pigment composition, chlorophyll a fluorescence and cell densities were measured for 25 m and included in multivariate analyses. Most functional variation was explained by two photobiological assemblages that relate to either symbiont identity or light microhabitat, suggesting adaptation and acclimation, respectively. Type B15 occurs with lower cell densities and larger sizes, higher cellular pigment concentrations and higher peridinin to chlorophyll a ratio than type B7. Type B7 relates to a larger xanthophyll-pool size. These unambiguous differences between symbionts can explain their distributional patterns, with type B15 being potentially more adapted to darker or deeper environments than B7. Symbiont cell size may play a central role in the adaptation of coral holobionts to the deeper reef. The existence of functional differences between B-types shows that the clade classification does not necessarily correspond to functional identity. This study supports the use of ITS2 as an ecological and functionally meaningful marker in Symbiodinium.     

Enhancement of photosynthesis in Sorghum bicolor by ultraviolet radiation

We assessed the influence of ultraviolet radiation (UV) on net photosynthetic CO₂ assimilation rate (P_n) in Sorghum bicolor, with particular attention to examining whether UV can enhance P_n via direct absorption of UV and absorption of UV‐induced blue fluorescence by photosynthetic pigments. A polychromatic UV response spectrum of leaves was constructed by measuring P_n under different UV supplements using filters that had sharp transmission cut‐offs from 280 to 382 nm, against a background of non‐saturating visible light. When the abaxial surface was irradiated, P_n averaged 4.6% higher with the UV supplement that cut‐off UV at 311 nm, compared to lower and higher UV wavelength supplements. This former supplement differed from higher wavelength supplements by primarily providing more UV between 320 and 350 nm. To assess the possibility of direct absorption of UV by photosynthetic pigments, we measured the absorbance of extracted chlorophylls. Chlorophyll a had absorbance peaks at 340 and 389 nm that were 49 and 72% of that at the sorét peak. Chlorophyll b had absorbance peaks at 315 and 346 nm that were both 35% of that at the sorét peak. Since the epidermis transmits some UV, the strong UV absorbance of chlorophyll implies a potential role for irradiance beyond the bounds of the conventionally defined photosynthetically active radiation waveband (400–700 nm). To assess the role of absorption of UV‐induced blue fluorescence, we measured the UV‐induced fluorescence excitation and emission spectra of leaves. Abaxial excitation peaked at 328 nm, while emission peaked at 446 nm. In this analysis, we used our abaxial fluorescence excitation spectrum and the UV photosynthetic inhibition spectrum of Caldwell et al. (1986) to weight the UV irradiance with each cut‐off filter, thereby estimating the potential contribution of UV‐induced blue fluorescence to photosynthesis and the inhibitory effects of UV irradiance on photosynthesis, respectively. With a non‐saturating visible background, we estimate that the absorption of UV‐induced blue fluorescence and the direct absorption of UV by photosynthetic pigments maximally enhanced photosynthesis by about 1% each with the UV supplement that cut‐off UV at 311 nm. We suggest that a portion of the incident UV on the S. bicolor leaves was used to drive photosynthesis.     

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.