Emerson enhancement effect and quantum yield of photosynthesis for marine macroalgae in simulated underwater light fields

The contribution of enhancement to the total photosynthesis of marine macroalgae in their natural habitats was estimated by comparing the photosynthesis measured by O₂-electrode in five broad-band light fields with that predicted (on the assumption that no enhancement was occurring) from the photosynthetic action spectrum of each plant and the spectral distribution of the light fields. The excess of measured values divided by calculated values provided a measure of enhancement. Although 37% enhancement was observed for red algae in unfiltered quartz-iodine light, and 18% for green and brown algae, substantially lower values were obtained for all species in more natural light fields. In those typical of shallow coastal waters, phycoerythrin-rich red algae exhibited 15 to 20% enhancement, but little enhancement (<5%) was detected in other algae. In a green light field, representing deep coastal water, there was no significant enhancement in any species, and only green and brown algae showed any enhancement (ca 8%) in broad-band blue light, similar to that in deep oceanic waters. Quantum yields of 0.09 to 0.10 O₂ molecules per absorbed photon were recorded in most light fields for green and brown algae with thin thalli, but yields decreased in the blue light field and in species with thicker thalli. All red algae had quantum yields of about 0.08 O₂ molecules per absorbed photon, except in the blue light field, in which quantum yields were reduced by 70%.     

Photosynthetic rates of benthic marine algae in relation to light intensity and seasonal variations

The effect of light intensity on rate of photosynthesis was measured at irregular intervals over a 12-month period for 22 benthic marine algae in the western Baltic Sea. In most species photosynthesis (mg O₂·g dry weight^-1·h^-1) was highest in spring and summer, corresponding to the seasonal growth pattern of the algae. In winter all the species showed adaptation of the light compensation point. Highest productivity was shown by algae which are: short-lived annual species rather than perennials, eulittoral rather than sublittoral, and which possess sheet-like or filamentous thalli rather than coarsely branched forms. These factors are clearly inter-related.This research was carried out while one of us (R.J.K.) was an Alexander von Humboldt fellow at the University of Kiel, and is part of the programme Sonderforschungsbereich 95, Wechselwirkung Meer-Meeresboden, Universität Kiel.     

Photosynthesis and amelioration of desiccation in the intertidal saccate alga Colpomenia peregrina

Rates of gross photosynthesis for the intertidal saccate alga Colpomenia peregrina (Sauv.) Hamel were determined under submersed and emersed conditions. Maximal photosynthetic rates were lower than for most seaweeds but comparable with other saccate members of the genus. By fitting the data to a hyperbolic tangent function, maximal photosynthetic rates were estimated to be 5.29 mmol CO₂ m^-2 h^-1 under submersed conditions and 2.06 mmol CO₂ m^-2 h^-1 under emersed conditions. I_k for submersed thalli was 69.1 E m^-2 s^-1, wherea for emersed thalli it was 149.0 E m^-2 s^-1, or 2.2 times higher. At low tide in the field and under saturating irradiance, carbon from seawater retained within the thallus cavity was assimilated at 0.9 mmol CO₂ m^-2 h^-1. In the laboratory under emersed conditions, carbon from this source was taken up at 0.6 mmol CO₂ m^-2 h^-1 at 20°C and at 0.34 mmol CO₂ m^-2 h^-1. Retained seawater also greatly reduced drying under desiccating conditions. Experimental thalli from which seawater had been removed lost thallus water continuously throughout the drying period (120 min). On the other hand, control, thalli lost water for the first 15 min, after which no further water loss occurred. At the termination of the experiment, control thalli had lost 7.2% of their water, whereas experimental thalli had lost 39.2%. Desiccation affected the emersed photosynthetic rate of experimental and control thalli. Emersed photosynthetic rates for thalli dried for 15 min were higher than for fullyhydrated thalli. However, emersed photosynthesis of thalli dried for longer than 15 min was lower than fully-hydrated rates and was directly related to percent water loss. Utilizing data from this study, a model was constructed to determine total photosynthetic production of C. peregrina over a single daylight period. From these calculations it was determined that emersed photosynthesis can increase daily photosynthetic production of C. peregrina by 50%.     

Seasonal effects of light,temperature, nutrient concentration and salinity on the physiology and growth of Caulerpa paspaloides (Chlorophyceae)

Caulerpa paspaloides (Bory) Greville were collected during the winter and summer (1978 to 1979) from the Florida Keys, USA. Thalli collected during the winter photosynthesized more efficiently at low light intensities (I_c<1, I_k=38 Exm^-2xs^-1) than did thalli collected in the summer (I_c=13, I_k=111 Exm^-2xs^-1). Summer thalli exhibited higher P_max values (2.20 mgO₂xg^-1 dry wtxh^-1) than winter thalli (1.70 mg O₂xg^-1 dry wtxh^-1). Rates of rhizome elongation and frond initiation were strongly inhibited by winter temperatures. The maximum lethal temperature for summer thalli was 37° to 38°C as measured by both growth and photosynthesis. No evidence of nitrogen or phosphorus limitation was found. Relatively minor reductions in salinity (3S) resulted in significant increases in rhizome apex motality. Results indicate that low winter temperatures are responsible for reduced winter growth rates previously reported for the Key Largo population. Increased photosynthetic efficiency at low light intensities and preferential maintenance of rhizome elongation over frond initiation appear to allow this tropical macroalga to optimize growth and survival during the winter.     

Long-distance transport of 14C-labelled assimilates in the Fucales: directionality,pathway and velocity

A directed translocation of ¹⁴C-labelled photoassimilates was shown to occur in the thalli of six species of brown seaweeds belonging to the Fucales. The apical growing regions of these algae act as strong sinks in the source-tosink relationship of the long-distance transport. Physical interruptions of the thalli showed that the midrib is the main pathway of the assimilates in Fucus vesiculosus and F. serratus. ¹⁴C-histoautoradiography has shown that translocation takes place through the filamentous cells of the medulla. The translocation velocity of the labelled assimilates, calculated from time source profiles of the tracer, was estimated to be 2 to 4 cm · h^-1. It is suggested that the sieve elements of the Fucales act in a way similar to the conducting sieve elements of the Laminariales.     

Reproduction,growth and photosynthesis of gametophytes of Laminaria saccharina grown in blue and red light

The induction of reproductive activity by blue light in female gametophytes of the brown alga Lamnaria saccharina has been investigated and related to other effects of red and blue light on these plants. Although germination of zoo-spores is slightly delayed in red light, female gametophytes grow at similar rates in blue and red light of the same quantum irradiance up to an age of 8 to 10 days, when plants in blue light begin to form eggs. The percentage of plants forming eggs is proportional to the total quanta of blue wavelengths received up to a saturating value of about 400 μE·cm^-2; a 50% response is induced by 200 μE·cm^-2. The action spectrum for the induction of fertility has a main peak at 430 to 450 nm, with two subsidiary peaks in the blue and one in the near-ultraviolet light, but the action spectrum for photosynthesis is quite different, with peaks in the red, green and blue regions. These results indicate that egg formation in female gametophytes occurs as a specific morphogenetic response to blue light, which is independent of photosynthesis and growth, and trial experiments with male gametophytes of L. saccharina and gametophytes of other species of Laminariales indicate that these plants react similarly.     

Preparative and analytical extraction of pigments from brown algae with dimethyl sulfoxide

Dimethyl sulfoxide (DMSO) rapidly extracts much of the chlorophyll c and fucoxanthin from the intact thalli of brown algae. Subsequent extraction by acetone rapidly removes most of the chlorophyll a and -carotene. An analytical procedure for these pigments has been developed, based upon this extraction sequence, followed by partition of the acetone extract between hexane and aqueous acetone. Fucoxanthin and chlorophyll c can be recovered from the DMSO extract and the aqueous acetone phase by extraction into ethyl acetate and subsequent evaporation. The DMSO-acetone extraction sequence is also effective with green algae and a few red algae, although the full scope of the method has not been investigated.     

Alkanes and alkenes in marine benthic algae

Saturated and olefinic hydrocarbons were determined in additional species of benthic marine algae from the Cape Cod (Massachusetts, USA) area (see: Youngblood et al., 1971). The distribution of homologous and isomeric olefins was studied in plants of different age and in morphologically different parts of the same specimen. With two minor exceptions, only normal alkanes and alkenes are present. The methylene-interrupted C₁₉- and C₂₁-polyolefins are particularly abundant; ¹-heneicosahexaene and the corresponding pentaene are common to all brown algae, while the corresponding ³-isomers occur in green algae. The hydrocarbon concentration, the alkene-to-alkane ratio and the polyolefin content are highest in young plants or in rapidly growing tissues of older plants. This suggests a deeper involvement in cell biochemistry of straight-chain hydrocarbons than previously considered. The biosynthesis of the plant polyolefins remains to be explored; no immediately obvious precursors of the ¹-polyolefins were found among the algal fatty acids. The hydrocarbon composition of these benthic algae differs greatly from that of fossil fuels in its simplicity and predominately unsaturated nature. The separation of the isomers by gas chromatography and their structural elucidation by mass spectrometry, alone and in combination with hydrogenation and ozonolysis, are discussed.Contribution No. 3155 of the Woods Hole Oceanographic Institution.     

Nitrogen fixation by blue-green algae associated with the siphonous green seaweed Codium decorticatum: effects on ammonium uptake

Nitrogen fixation (acetylene reduction) at rates of up to 1.2 g N₂ g dry wt^-1 h^-1 was measured for the siphonous green seaweed Codium decorticatum. No nitrogenase activity was detected in C. isthmocladum. The nitrogenase activity was light sensitive and was inhibited by the addition of DCMU and triphenyl tetrazolium chloride. Additions of glucose did not stimulate nitrogen fixation. Blue-green algae (Calothrix sp., Anabaena sp., and Phormidium sp.) were implicated as the organisms responsible for the nitrogenase activity. They occurred in a reduced microzone within the C. decorticatum thallus where nitrogen fixation was optimized. Nitrogen fixation did not affect the kinetic constants for ammonium uptake in C. decorticatum (K_s=12.0 M, V_max=13.4 mol NH₃ g dry wt^-1 h^-1) determined using the perturbation method. Nevertheless, C. decorticatum thalli which fixed nitrogen had internal dissolved nitrogen concentrations which were over 1.4 times higher than in non-fixing thalli. This suggests that if C. decorticatum does derive part of its nitrogen requirement from the blue-green algae which it harbors, the transfer does not involve competition between this process and the uptake of ambient ammonium.     

Photosynthetic acclimation and biochemical responses ofGelidium sesquipedale cultured in chemostats under different qualities of light

The red algaGelidium sesquipedale (Clem.) Born. et Thur. has been cultured in chemostats to assess the effects of light quality and photon-fluence rate (PFR) on growth, photosynthesis and biochemical composition. Plants under blue and red light (BL and RL) showed higher growth rates than under white light (WL) of the same PFR (40 mol m^–2 s^–1). The light-saturated rate of photosynthesis was higher for algae grown under BL and RL than for algae grown under WL. When algae were transferred to WL of moderate PFR (100 mol m^–2 s^–1), the light-saturated rate of photosynthesis decreased, being higher in previously RL-grown algae than in previously BL- and WL-grown algae. The initial slope of photosynthesis-irradiance (PI) curves () was affected by PFR but not by light quality. Pigment content was little affected by light quality. Light-quality treatments also affected the biochemical composition of the alga; previous exposure to various light treatments activate or repress several metabolical pathways that are fully expressed in the subsequent phase of WL of moderate PFR. Thus, phycobiliproteins and soluble proteins increased for previously BL- and RL-grown algae, whereas insoluble carbohydrate concentration was reduced, indicating a change of the C-partitioning between carbon compounds and organic nitrogen compounds. Inorganic nitrogen metabolism was also affected by light: under WL of moderate PFR, NO₃ ^– was totally depleted from sea water, and maximal values of NO₃ ^– uptake were recorded. In addition, neither NO₂ ^– nor NH₄ ⁺ was released. However, when algae were transferred to a low PFR, there was a drastic reduction of NO₃ ^– uptake under WL, which only partially recovered over time. It was accompanied by the release of NO₂ ^–, but not NH₄ ⁺, to the culture medium. Under BL and RL, however, there was a transient enhancement of NO₃ ^– uptake that was followed by a net release of NO₂ ^– and NH₄ ^–. Growth rates were not correlated with PFR. This could be due to the the dynamics of internal carbon mobilization and accumulation in the algae. When algae were exposed to a moderate PFR of WL, carbon requirements for growth were satisfied by photosynthesis. Thus, there was a net accumulation of carbon in the tissue. In contrast, when algae were exposed to low PFRs of either WL, BL or RL, observed growth rates could not be maintained by photosynthesis and carbon was mobilized.     

Carbon acquisition strategies for marine macroalgae

A model system was developed to analyse differences in carbon acquisition strategies among macroalgae. During photosynthesis in a limited volume of seawater the capability of the algae to assimilate inorganic carbon as well as to change the alkalinity of the seawater was analysed. These properties were then related to the status of the carbonate equilibrium system of the seawater. The experimental system was assumed to simulate the conditions in the boundary layer during periods of low water exchange or high intensity irradiations. Fundamental differences were found between different algal classes, suggesting that capabilities to adapt to specific environmental conditions may be connected with dissimilarities in carbon acquisition strategies. In general, green algae were able to reach the highest pH (10.8 at 5°C), and thus to achieve the highest reduction in the level of inorganic carbon via a simple HCO₃ ^–/OH^– ion exchange process. For brown algae, pH increases due to carbon uptake never exceeded pH 9.7 (9.5 in a saltwater scale). In spite of this, members of the Fucaceae (littoral brown algae) were able to extract almost all of the dissolved inorganic carbon (DIC). This was achieved through a gradual decrease in the alkalinity of the enclosed water, so that the carbon assimilation could continue without any concomitant increase in pH. For red algae, the specific response was an increase in the level of inorganic carbon. Thus, for this algal class, no specific strategy for handling a shortage of inorganic carbon was documented. Within each algal class, differences in pH and DIC compensation points could be related to differences in the depths at which the algal species occurred. This paper also introduces a low cost and convenient method of analysing DIC in seawater.     

Seasonal photosynthetic characteristics of Ascoseira mirabilis (Ascoseirales,Phaeophyceae) from King George Island,Antarctica

Monthly variation in photosynthesis, dark respiration, chlorophyll a content and carbon: nitrogen (C:N) ratios in different lamina sections of adult plants of Ascoseira mirabilis Skottsberg from King George Island, Antarctica, was investigated between September 1993 and February 1994. Light saturated net photosynthesis (P _max) showed maximum values in September (12 to 25 mol O₂ g^-1 fr wt h^-1), and decreased towards the summer to values ranging between 2.0 and 5.0 mol O₂ g^-1. In the distal section, however, a second optimum occurred in December (25 mol O₂ g^-1 fr wt h^-1). Dark respiration rates were also highest in October and November and decreased strongly in December to February (6.0 and 1.0 mol O₂ g^-1 fr wt h^-1, respectively). Gross photosynthesis exhibited high values between September and December. Concomitant with the seasonal decrease of photosynthetic efficiency () from mean values of 1.2 mol O₂ g^-1 fr wt h^-1 (mol photons cm^-2 s^-1)^-1 in September to 0.3 mol O₂ g^-1 fr wt h^-1 (mol photons cm^-2 s^-1)^-1 in January, the initial light saturating point (I _k) gradually increased from 19 to 60 mol photons m^-2 s^-1. Likewise C:N ratios were low in spring (12 to 13) and increased in summer (20). In general, the photosynthetic parameters P _max, gross photosynthesis, and Chl a concentrations were significantly higher in the distal section of the thallus. In contrast, C:N ratios were lower in the distal section of the lamina. The results show that photosynthesis obviously strongly supports growth of the alga in late winter to spring, as it does in some morphologically related brown algae from temperate and polar regions. The question whether growth is additionally powered     

Saturated and unsaturated hydrocarbons in marine benthic algae

Saturated and olefinic hydrocarbons were determined in 24 species of green, brown and red benthic marine algae from the Cape Cod area (Massachusetts, USA). Among the saturated hydrocarbons, n-pentadecane predominates in the brown and n-heptadecane in the red algae. A C₁₇ alkyleyclopropane has been identified tentatively in Ulvalactuca and Enteromorpha compressa, two species of green algae. Mono-and diolefinic C₁₅ and C₁₇ hydrocarbons are common. The structures of several new C₁₇, C₁₉ and C₂₁ mono-to hexaolefins have been elucidated by gas chromatography, mass spectrometry and ozonolysis. In fruiting Ascophyllum nodosum, the polyunsaturated hydrocarbons carbons occur exclusively in the reproductive structures. The rest of the plant contains n-alkanes from C₁₅ to C₂₁. A link between the reproductive chemistry of benthic and planktonic algae and their olefin content is suggested. An intriguing speculation is based on Paffenhöfer's (1970) observation that the sex ratio of laboratory reared Calanus helgolandicus depends upon the species of algae fed to the nauplii. The percentage of males produced correlates with our analyses of heneicosahexaene in the algal food. Our analyses of the hydrocarbons in benthic marine algae from coastal environments should aid studies of the coastal food web and should enable us to distinguish between hydrocarbon pollutants and the natural hydrocarbon background in inshore waters.Contribution No. 2582 of the Woods Hole Oceanographic Institution; and Contribution No. 227, Systematics-Ecology Program.     

Capability of dynamic photoinhibition in Arctic macroalgae is related to their depth distribution

The capability of several macroalgal species to protect photosynthesis from excessive irradiance by dynamic photoinhibition was investigated relative to their depth distribution in summer 1995 in the Kongsfjord (79°N; 12°E, Ny Ålesund, Spitsbergen, Norway). Photoinhibition of photosynthesis was induced by exposure of algae from different water depths to a high photon fluence rate of 500?μmol?m^?2?s^?1 for 2?h. Changes in optimal quantum yield (F _v/F _m) were measured during the inhibition phase. Recovery of photosynthesis was subsequently induced by dim white light (10?μmol?m^?2?s^?1) and observed as changes in the variable fluorescence. With a newly developed mathematical model different parameters of the response kinetics of inhibition and recovery were calculated and related to the depth distribution of each algal species. It is shown that two components with slow and fast reaction kinetics, respectively, are involved in photoinhibition and recovery of photosynthesis. Their possible molecular bases are discussed. The half-life time (τ) of the inhibition and recovery phases, i.e. the time necessary to reach half maximal response, is clearly related to the depth distribution of the investigated species. Algae collected close to the water surface show a fast reaction of both photoinhibition and recovery and, hence, have a low τ. With increasing depth the reactions become slower and τ increases. τ was highest in deep water algae. Further analysis of the reaction kinetics in Laminaria saccharina shows that the relative proportion of the two kinetics involved change with the collection depth. In contrast, a significant difference in the reaction rates of both kinetics was not observed.     

Influence of light on algal symbionts of the deep water coral Leptoseris fragilis

Photoadaptations of zooxanthellae living within the deep water coral Leptoseris fragilis taken from the Gulf of Aqaba (Red Sea) were studied. Specimens-collected in summer 1988 between 110 and 120 m depth —were transplanted to 70 and 160 m. At each depth individuals were exposed in their natural growth position (oral side facing the surface) or in a reverse growth position (oral side facing the bottom). After 1 yr of exposure the corals were collected and the zooxanthellae were isolated. As a function of the availability of light with depth and growth position several algal parameters showed changes which are related to photoadaptations. The relatively low density of zooxanthellae of 0.15x10⁶ cellsxcm^-2 at a natural growth depth of 116 m decreased to 0.0034x10⁶ cellsxcm^-2 (2%) at 160 m in specimens growing with a natural orientation. In corals with a downward-facing oral surface at the same depth (160 m) only degenerated algae could be observed. With respect to depth dependence the volume of the algae decreased from 728 m³ at 116 m to 406 m³ at a depth of 160 m and the content of pigments increased. The augmentation of peridinin per cell was low (two times at 160 m compared to 116 m). Chlorophyll a and in particular chlorophyll c ₂ concentrations per cell were enhanced. Compared to natural amounts at 116 m, chl a was five times and chl c ₂ eight times higher at 160 m. At all depths the chl c ₂ content per cell was higher than for chl a. The formation of chl a/chl c ₂ complexes as light harvestor is discussed. Light harvesting, with chl c ₂ prevailing may be explained as a special type of chromatic adaptation of L. fragilis in a double sense: (1) in the habitat light short wavelengths predominate. This light can be directly absorbed with pigments such as chl a and chl c ₂. (2) Host pigments absorb visible violet light and transform these wavelengths, less suitable for photosynthesis, into longer ones by means of autofluorescence. The emitted longer wavelengths fit the absorption maxima of the algal pigments. Thus the host supports photosynthesis of his symbionts. Corals exposed at 160 m depth with a downward facing oral surface were alive after 1 yr and the host wavelength transforming pigment system was still present, but zooxanthellae were absent or degenerated. The light field at 160 m seems therefore to be critical: the combined photoadaptations of host and symbionts, allowing photosynthesis under barren light conditions, seem to be exhausted. In L. fragilis the photoadaptive strategies of host and symbionts cooperate harmoniously. In addition, the adaptations are interlocked with the particular light situation of the habitat with respect to light quantity and quality. The cooperation of physical and organismic parameters examplifies how evolution and, in particular, coevolution has led to optimal fitness.     

Light harvesting in the green alga Ostreobium sp., a coral symbiont adapted to extreme shade

Ostreobium sp. (Chlorophyta: Siphonales) can be found as green bands within the skeletal material of a number of stony corals in the Indo-Pacific and Caribbean regions. Many of these corals also contain symbiotic dinoflagellates in the overlaying coral polyps that effectively screen out all the typical photosynthetically active radiation from the algae in the green bands below. Ostreobium sp., nevertheless, grows photosynthetically. Its action spectrum and absorption spectrum have been shown to extend much further into the near infra-red compared to other green algae. In the present study, carried out in 1987, fluorescence excitation and emission spectra were measured in Ostreobium sp. and compared to spectra obtained from the green alga Ulva sp. and the brown alga Endarachne sp. Xanthophylls, probably siphonein and an unidentified xanthophyll probably related to siphonaxanthin, are photosynthetically active in Ostreobium sp., and can sensitize Photosystem II fluorescence at 688 nm and Photosystem I (PS I) fluorescence at 718 nm. The fluorescence emission spectra of Ostreobium sp. measured at 25° C and 77 K were not remarkably different from those of the green alga Ulva sp. Absorbance changes induced by light were measured in Ostreobium sp. from 670 to 750 nm and were like those normally seen in green plants except that, in addition to the minimum expected for the reaction-center chlorophyll of PS I (P700) at 703 nm, another minimum was seen at 730 nm. It is possible that this spectrumreflects the functioning of a reaction center of Photosystem I that has adapted to function in light highly enriched in far-red wavelengths.CIW-DPB Publication No. 1021     

Effect of phosphate and ammonium levels on photosynthetic and respiratory responses of the red alga Gracilaria verrucosa

Gracilaria verrucosa (Hudson) Papenfuss exposed to nutrient enriched media (0.1 mM PO₄; 1.0 mM NH ₄ ⁺ ) by pulse feeding 2 h every third day for a period of 5 wk at 20°C and 25–30 salinity showed significantly higher rates of photosynthesis regardless of photon flux density correlated with increased pigment levels. Algae in nonenriched media showed significantly higher levels of soluble carbohydrates and decreased levels of phycoerythrin and chlorophyll a. Photosynthetic and respiratory responses to temperature 15°, 25°, 30°C and salinity (15, 25, 30 S) combinations indicate broad tolerances by both nutrient enriched and non-nutrient enriched algae. Photosynthetic and respiratory rates were highest at the high temperatures. Pulse-fed algae had significantly higher photosynthetic rates than non-nutrient enriched plants at all temperature and salinity combinations. Non-nutrient enriched algae had significantly higher respiratory rates than nutrient enriched algae at only 30°C and 15. The respiratory rates of both nutrient enriched and non-nutrient algae decreased under combinations of higher temperatures and salinities. G. verrucosa, grown without nutrients, has lower tolerances to environmental stresses.     

Photosynthetic characteristics of Prochloron sp./ascidian symbioses

The prokaryotic green alga Prochloron sp. (Prochlorophyta) is found in symbiotic association with colonial didemnid ascidians that inhabit warm tropical waters in a broad range of light environments. We sought to determine the light-adaptation features of this alga in relation to the natural light environments in which the symbioses are found, and to characterize the temperature sensitivity of photosynthesis and respiration of Prochloron sp. in order to assess its physiological role in the productivity and distribution of the symbiosis. Colonies of the host ascidian Lissoclinum patella were collected from exposed and shaded habitats in a shallow lagoon in Palau, West Caroline Islands, during February and March, 1983. Some colonies from the two light habitats were maintained under conditions of high light (2 200 E m^–2 s^–1) and low light (400 E m^–2 s^–1) in running seawater tanks. The environments were characterized in terms of daily light quantum fluxes, daily periods of light-saturated photosynthesis (H_sat), and photon flux density levels. Prochloron sp. cells were isolated from the hosts and examined for their photosynthesis vs irradiance relationships, respiration, pigment content and photosynthetic unit features. In addition, daily P:R ratios, photosynthetic quotients, carbon balances and photosynthetic carbon release were also characterized. It was found that Prochloron sp. cells from low-light colonies possessed lower chlorophyll a/b ratios, larger photosynthetic units sizes based on both reaction I and reaction II, similar numbers of reaction center I and reaction center II per cell, lower respiration levels, and lower P_max values than cells from high-light colonies. Cells isolated from low-light colonies showed photoinhibition of P_max at photon flux densities above 800 E m^–2 s^–1. However, because the host tissue attenuates about 60 to 80% of the incident irradiance, it is unlikely that these cells are normally photoinhibited in hospite. Collectively, the light-adaptation features of Prochloron sp. were more similar to those of eukaryotic algae and vascular plant chloroplasts than to those of cyanobacteria, and the responses were more sensitive to the daily flux of photosynthetic quantum than to photon flux density per se. Calculation of daily minimum carbon balances indicated that, though high-light cells had daily P:R ratios of 1.0 compared to 4.6 for low-light cells, the cells from the two different light environments showed nearly identical daily carbon gains. Cells isolated from high-light colonies released between 15 and 20% of their photosynthetically-fixed carbon, levels sufficient to be important in the nutrition of the host. Q₁₀ responses of photosynthesis and respiration in Prochloron sp. cells exposed briefly (15–45 min) to temperatures between 15° and 45°C revealed a discontinuity in the photosynthetic response at the ambient growth temperatures. The photosynthetic rates were found to be more than twice as sensitive to temperatures below ambient (Q₁₀=3.47) than to temperatures above ambient (Q₁₀=1.47). The Q₁₀ for respiration was constant (Q₁₀=1.66) over the temperature range examined. It appears that the photosynthetic temperature sensitivity of Prochloron sp. may restrict its distribution to warmer tropical waters. The ecological implications of these findings are discussed in relation to published data on other symbiotic systems and free-living algae.     

Effects of O2, pH and DIC on photosynthetic net-O2 evolution by marine macroalgae

Net photosynthetic O₂ evolution by five marine macroalgae:Ulva lactuca L.,Enteromorpha sp.,Ceramium strictum Harvey,Fucus serratus L., andF. vesiculosus L., collected from Danish waters in the summer of 1983 was followed at increasing O₂ and with pH either fixed close to pH 7, 8 or 9, or drifting upwards during photosynthesis in a closed chamber to determine the effects of changing O₂, pH and DIC (dissolved inorganic carbon) on photosynthesis. Increasing O₂, increasing pH and decreasing DIC together limited O₂ evolution. Raising the O₂ concentration with pH and DIC held constant resulted in less inhibition of net-O₂ evolution than when all three factors acted together. The O₂ inhibition of photosynthesis was similar to the reported O₂ inhibition of ribulose 1,5-bisphosphate carboxylase isolated from lower and higher plants. Net-O₂ evolution as a function of the molar ratio of O₂ to HCO^– ₃ + CO₂ in solution provided a general, linear relationship (r ² = 0.72 to 0.84), predicting inhibition of photosynthesis based on O₂ pH and DIC changing together. Slopes of this relationship, representing competition between O₂ and carbon based on external concentrations, were similar for the five taxonomically different algae, suggesting that similar processes act to reduce net-O₂ evolution.     

Characterization of in vivo absorption features of chlorophyte,phaeophyte and rhodophyte algal species

Despite the plentiful diversity of macroalgae in coastal environments, few studies have examined the in vivo absorption features of common marine macrophytes. Here we report on results of a survey of 12 central California common intertidal and subtidal taxa, representing Chlorophyta, Phaeophyta and Rhodophyta carried out in the summer of 1988. Computer-assisted analyses were used to obtain fourth-derivative spectra from in vivo absorption spectra determined at room temperature to obtain spectral diagnostics for the different algal divisions and to provide a means to determine whether spectral features could be used to identify stress responses among these plants. Among the Chlorophyta, characteristic maxima for chlorophylls a and b were resolved in all species examined, and a spectral component attributable to siphonaxanthin-like carotenoid(s) was observed in two Ulva species and the coenocytic alga Derbesia marina. Representatives of the Phaeophyta were characterized by similar maxima for chlorophyll a and c, and for fucoxanthin. Among the Rhodophyta, maxima for chlorophyll a, as well as B- and R-type phycoerythrin were resolved. Differences in in vivo absorption features were detected for two tidal populations of Porphyra perforata and Mastocarpus papillatus. High-tidal thalli absorb less green light in regions characterized by phycoerythrin, but have enhanced carotenoid absorption compared with lowtidal thalli. Resolution of spectra by fourth-derivative analysis revealed significant differences in phycoerythrin and carotenoid contents. The spectral changes observed appear to reflect environmental and possibly populational characteristics of these algae. The spectral analyses described here provide robust, non-invasive means to characterize subtle responses of macroalgae to environment in ways not possible previously. Additional merits of these fourth-derivative analyses for use in environmental studies are discussed.This contribution is dedicated to the memory of Professor L. R. Blinks, a pioneer in photosynthesis research and algal physiology