Effects of high-frequency light fluctuations on growth and photoacclimation of the red algaChondrus crispus

High-frequency fluctuations due to wavefocusing are prominent characteristics of light in shallow marine environments. Effects of high-frequency (0.01 to 1 Hz) fluctuating light on growth rates of the red algaChondrus crispus Stackh., collected from Crane Neck Point, New York, USA, during July 1988, were determined by comparing plants grown under fluctuating and constant light regimes with similar daily irradiances. At high daily irradiance, growth rates were higher under fluctuating light than under constant light. Fluctuating light effects were frequency-dependent; growth was enhanced by fluctuations at 0.1 and 1 Hz, but not by fluctuations at 0.01 Hz. At low daily irradiance, growth rates were not affected by fluctuating light at any frequency tested. Enhancement of growth was not due to effects of high-frequency light fluctuations on photoacclimation responses ofC. crispus. Plants grown under fluctuating light at high daily irradiance actually exhibited lower photosynthetic capacity and efficiency (determined under constant light) than plants grown under constant light. These differences were attributable to variation in the density of Photosystem II reaction centers, which was low in plants grown under fluctuating light. Maximum turnover rate of whole-chain electron transport and activity of ribulose-1,5-bisphosphate carboxylase were affected by total daily irradiance, but not by high-frequency light fluctuations. Enhancement of growth under fluctuating light was partly attributable to reduced rates of dark respiration compared to rates of plants grown under constant light. The results also provided indirect evidence that high-frequency light fluctuations may enhance instantaneous photosynthetic rates. This effect could increase daily carbon gain and, therefore, stimulate growth ofC. crispus under high-frequency light fluctuations.     

Comparative study of crustacean larval photoresponses

Ovigerous females of four brachyuran (Cancer gracilis, Lophopanopeus bellus bellus, Hemigrapsus oregonensis and Scyra acutiforns) and two anomuran (Pagurus beringanus and P. granosimanus) species, which live as adults in coastal areas, were collected near Friday Harbor, Washington, USA, in 1985, and spectral sensitivity and phototactic pattern of their larvae were measured. Responses were compared with previous measurements on estuarine species to determine whether responsiveness varies with adult habitat. Estuarine and coastal species have similar photoresponses. Spectral sensitivity of the test brachyran species had two maxima, one near 400 to 420 nm and another around 500 nm. The anomuran species were similar, but had an additional peak in the region of 580 to 620 nm. This sensitivity is adapted to daytime light conditions in the adult environment, and available spectra at the time of larval movement during diel vertical migration. Upon light adaptation and stimulation with a narrow light field, all six species showed positive phototaxis to high light intensities and a pronounced negative response to low intensities. Only the negative response will occur in natural underwater light conditions, and it is part of a predator-avoidance shadow-response which operates in areas of higher light intensity. The same phototactic pattern is observed in all species when darkadapted except H. oregonensis. If nocturnal vertical migration occurs, this negative response may be responsible for the descent at sunrise and depth maintenance during the day.     

Light availability and diurnal growth of a green macroalga (Caulerpa cupressoides) and a seagrass (Halophila decipiens)

The effects of daily light period on diurnal growth patterns of a green macroalga [Caulerpa cupressoides v.lycopodium f.elegans (J. Agardh) Weber-van Bosse] and a seagrass (Halophila decipiens Ostenfeld) were investigated in Salt River submarine canyon in the US Virgin Islands in summer 1984. The daily light period, in which quantum irradiance exceeded the light saturation point for photosynthesis of the macroalga and seagrass, was manipulated in situ using lamps and shades. Plant growth was measured every 6 h for 7 d under natural and experimental daily light periods.C. cupressoides grew at the same rate day and night.H. decipiens grew more during the day than at night, a pattern that persisted under continuous light and dark treatments, indicating endogenous control of diurnal growth. Growth vs daily light period curves indicate thatC. cupressoides grew faster thanH. decipiens in short daily light periods, consistent with the observation that the macroalga penetrates to deeper water than the seagrass in Salt River canyon. Overall growth (day + night) ofH. decipiens was unaffected in lengthened light periods and reduced in shortened light periods. Chlorophyll content ofC. cupressoides was not correlated with light availability, while that ofH. decipiens was positively correlated. The alga and seagrass had different diurnal growth patterns but similar overall growth responses to daily light periods. This study shows that diurnal growth patterns are probably under endogenous control, while overall growth is a response to in situ light conditions.Contribution^#193 from West Indies Laboratory and the National Undersea Research Program     

The ontogeny of phototaxis by larvae of the crab Rhithropanopeus harrisii

Phototaxis by each zoeal stage of the crab Rhithropanopeus harrisii (Gould) was quantitatively measured by means of a microscope closed-circuit television system. The megalopa stage is indifferent to light stimulation and was thus not tested. The action spectrum for positive phototaxis is similar at each zoeal stage, having the most pronounced maximum at about 500 nm and smaller maxima at 400 and 280 nm. Responsiveness to various intensities of 500 nm light is also similar at each stage. After dark-adaptation strongest positive phototaxis occurs at intensities between 1.0 and 3.0x10^-4 W/m², with no negative phototaxis to lower intensities. After light-adaptation, the positive response occurs to higher intensities between 20 and 0.01 W/m², with a pronounced negative response to lower intensities. Generally, mean swimming speeds during positive phototaxis do not change with stimulation intensity, although during negative phototaxis mean speeds do vary with intensity. At the highest intensities which initiate a negative response, mean values are significantly greater than values for positive phototaxis. Based on the pattern of phototaxis at each developmental stage, a prediction of events during diurnal vertical migration is possible. The negative response after light-adaptation might also function as a shadow reflex.     

Diel vertical migration of the marine copepod<Emphasis Type="Italic"> Calanopia americana</Emphasis>. II. Proximate role of exogenous light cues and endogenous rhythms

The marine copepod Calanopia americana Dahl undergoes twilight diel vertical migration (DVM) in the Newport River estuary, North Carolina, USA, in synchrony with the light:dark cycle. Copepods ascend to the surface at sunset, descend to the bottom around midnight, and make a second ascent and descent before sunrise. Behavioral assays with C. americana in the laboratory during fall 2002/2003 and summer 2004 investigated aspects of three hypotheses for the proximate role of light in DVM: (1) preferendum hypothesis (absolute irradiance), (2) rate of change hypothesis (relative rates of irradiance change), and (3) endogenous rhythm hypothesis. Results suggest that C. americana responds to exogenous light cues consistent with its DVM pattern; changes in absolute irradiance evoked swimming responses that would result in an ascent at sunset and descent at sunrise, while relative rates of irradiance decrease at sunset (–0.0046 s^–1) evoked an ascent response, and relative rates of irradiance increase at sunrise (0.0042 s^–1) evoked a descent response. Furthermore, C. americana expressed an endogenous rhythm in vertical migration that was positively correlated with field observations of twilight DVM. Collectively, these results indicate that both exogenous light cues and endogenous rhythms play a proximate role in twilight DVM of C. americana, providing redundancy in the causes of its vertical migration.Communicated by J.P. Grassle, New Brunswick     

Representing the effects of diurnal variations in light on primary production on a seasonal time-scale

Rates of production by primary producers are strongly affected by light. Diurnal variations in irradiance produce characteristic diurnal patterns in primary production and respiration. In cases in which the processes of interest occur on a longer time-scale, it is not uncommon to ignore diurnal variations and use mean daily irradiance to force models of primary producers. This simplification reduces the computer processing time and input data requirements of models. However, this approach results in an error which may be significant if irradiance during part of the day is higher than that needed for growth at the maximum possible rate. Here, an alternative approach is presented and applied to a simulation of phytoplankton in a shallow lake. The model uses a semi-analytical calculation of mean daily growth rates that takes into account periods of supersaturation with respect to light to deliver results that conform closely to those of a model using hourly time-steps and irradiance forcing, but with a substantially smaller computational cost.     

Effect of light intensity on activity and food-searching of larval herring,Clupea harengus: a laboratory study

Larvae of Clupea harengus were reared from spawning herring caught in March 1982 and 1983 in the Firth of Clyde, Scotland. An infra0red observation technique was used to record the behaviour of larval herring both in shallow dishes using a top view and in a tank 2 m deep using a side view. The amount of time larvae spent swimming, which was minimum in complete darkness, increased with increasing light intensity and as the larvae grew. Maximum swimming speeds of feeding larvae were recorded at light intensities between 10 and 100 lux. The presence of food organisms (Artemia sp., Brazilian strain) at light intensities below the feeding threshold (0.1 lux) caused an increase in the proportion of time spent active, but light intensities above the threshold had different effects, depending on developmental stage: larvae of 12 mm increased swimming speed, but 21 mm larvae decreased speed. In the 2 m deep tank in darkness, larvae displayed inactive periods wherein they sank head first, interspersed with periods of upward swimming. As light intensity increased, vertical swimming was replaced by horizontal swimming. These results are discussed with reference to food searching and vertical migration of larval herring in the sea.     

The concept of light intensity adaptation in marine phytoplankton: Some experiments with Phaeodactylum tricornutum

The historical background on adaptation of algae to various light intensities is analysed. It is argued that there is little evidence to suggest that previous growth at low light intensities enhances the ability of an alga to utilize these low light levels. Rather, the published evidence suggests that the most general response to growth at sub-optimal light intensities is a reduced ability to utilize saturating levels. The present experiments with Phaeodactylum tricornutum Bohlin have tested this concept of light intensity adaptation. Changing photosynthetic abilities during batch growth depended on the light intensity used for growth and these changes affected interpretations of the data. When measurements were made intensities appeared to photosynthesize (at all intensities) better than did those grown at higher light levels. When the changes during batch growth were taken into account, or when the alga was grown in turbidostat cultures, a different picture was obtained. Growth at reduced light intensities was accompanied by (a) increased chlorophyll content, (b) decreased rate of light-saturated photosynthesis expressed on a chlorophyll, cell number or cell protein basis, and (c) decreased activity of RuDP carboxylase. One result suggested that growth at a suboptimal light intensity did enhance the ability to utilize lower light levels. The light-saturation curve of cells grown in batch culture at 0.7 klux showed higher slopes at the low light intensities than did those grown at 12 klux. This was most marked when photosynthesis was expressed per cell, but was also apparent when it was put on a per chlorophyll basis.     

Variation in colony geometry modulates internal light levels in branching corals, <Emphasis Type="Italic">Acropora humilis</Emphasis> and <Emphasis Type="Italic">Stylophora pistillata</Emphasis>

Colonial photosynthetic marine organisms often exhibit morphological phenotypic plasticity. Where such plasticity leads to an improved balance between rates of photosynthesis and maintenance costs, it is likely to have adaptive significance. To explore whether such phenotypic plasticity leads to more favourable within-colony irradiance for reef-building branching corals, this relationship was investigated for two coral species Acropora humilis and Stylophora pistillata, along a depth gradient representing light habitats ranging from 500 to 25 μmol photons m⁻² s⁻¹, during 2006 at Heron Island, Great Barrier Reef (23.44°S, 151.91°E). In the present study changes in flow-modulated mass transfer co-varied with light as a function of depth. In low-light (deep) habitats, branch spacing (colony openness) in A. humilis and S. pistillata was 40–50% greater than for conspecifics in high-light environments. Also, branches of A. humilis in deep water were 40–60% shorter than in shallow water. Phenotypic changes in these two variables lead to steeper within-colony light attenuation resulting in 38% higher mean internal irradiance (at the tissue surface) in deep colonies compared to shallow colonies. The pattern of branch spacing was similar for S. pistillata, but this species displayed an alternate strategy with respect to branch length: shade adapted deep and cave colonies developed longer and thinner branches, allowing access to higher mass transfer and irradiance. Corals in cave habitats allowed 20% more irradiance compared to colonies found in the deep, and had a 47% greater proportion of irradiance compared to colonies in the shallow high-light environment. Such phenotypic regulation of internal light levels on branch surfaces partly explains the broad light niches of many branching coral species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Light response curve methodology and possible implications in the application of chlorophyll fluorescence to benthic diatoms

Chlorophyll a fluorescence has been increasingly applied to benthic microalgae, especially diatoms, for measurements of electron transport rate (ETR) and construction of rapid light response curves (RLCs) for the determination of photophysiological parameters [mainly the maximum relative ETR (rETR_max), the light saturation coefficient (E _k) and the maximum light use coefficient (α)]. Various problems with the estimation of ETR from the microphytobenthos have been identified, especially in situ. This study further examined the effects of light history of the cells and light dose accumulation during RLCs on the fluorescence measurements of ETR using the benthic diatom Navicula phyllepta. RLCs failed to saturate when using incremental increases in irradiance, however, curves with decreasing irradiance did saturate. Patterns indicating photoacclimation in response to light histories were observed, with higher rETR_max and E _k, and lower α, at high light compared to low light. However, these differences could be negated by increasing the RLC irradiance duration from 30 to 60 s. It is suggested that problems arose as a result of rapid fluorescence variations due to ubiquinone (Q_A) oxidation and non-photochemical chlorophyll fluorescence quenching (NPQ) which depended upon the light history of the cells and the RLCs accumulated light dose. Also, RLCs with irradiance duration of 10 s were shown to have a high level of error possibly specific to the fluorimeter programming. It is suggested that RLCs, using a Diving-PAM fluorimeter on benthic diatoms, should be run using decreasing irradiance steps of 30 s duration.     

Light-shade adaptation by the oceanic dinoflagellates Pyrocystis noctiluca and P. fusiformis

Species-specific rates of photosynthetic carbon uptake (P), chlorophyll a content and P versus irradiance (P-I), have been measured for cells of Pyrocystis noctiluca and P. fusiformis isolated from natural populations collected in the euphotic zone within and below the surface mixed layer in the Sargasso Sea. These same measurements and the assay for ribulose bis-phosphate carboxylase (RuBP-Case), have been made for cultures of P. noctiluca in a 12 h L: 12 h D photoperiod at 9 different constant or at changing light intensities. In nature chl a cell^-1 was constant throughout the euphotic zone. The photosynthetic capacity (P_max), of cells captured below the surface mixed layer was lower by a factor of 10 compared with cells collected from the surface mixed layer. The P_max for P. noctiluca collected and incubated within the surface mixed layer was the same as for cell cultures grown under high light, non nutrient-limiting conditions, suggesting that photosynthesis in the natural system was not nutrient limited. In laboratory cultures under constant low light intensities, chl a cell^-1 increased by a factor of 5 while both P_max and RuBPCase activity decreased by a factor of ca 4 compared with high light intensities. In changing light intensities both P_max and RuBPCase activities were decreased by factors of 4 during low light intervals while chl a cell^-1 approached a constant intermediate value. The change in chl a cell^-1 in response to prolonged exposure to constant low light intensities was first order with a rate constant of 0.33 d^-1. For all irradiance conditions in culture, the P-I dependence could be described by the simple Michaelis-Menten formula. The ratio of P_max to K_I, (the light intensity where P=P_max/2) was a constant with a Coefficient of Variation of 12%: The constancy of this ratio, the parallel changes in RuBPCase activity with P_max and the constant chl a cell^-1 in the Sargasso Sea imply that for P. noctiluca and presumably P. fusiformis in nature, a dark enzymatic step rather than changes in photosynthetic pigment concentrations may regulate the photosynthetic capacity in the changing photic environment.Contribution no. 1141 from McCollum-Pratt Institute and Department of Biology, The Johns Hopkins University. Supported by DOE contract no. EY 76S20 3278, NSF no. OCE 76-02571 and ONR no. N300014-81-C-0062     

Seasonal growth and biomass of the subtropical seagrassHalodule wrightii in relation to continuous measurements of underwater irradiance

Continuous year-round measurements of photosynthetically active radiation (PAR) were collected in relation to leaf elongation and plant biomass in the shoal-grass,Halodule wrightii Aschers., within three different estuarine systems on the south Texas coast (Laguna Madre: May 1989 to September 1993; Corpus Christi Bay: February 1990 to September 1993; San Antonio Bay; May 1990 to April 1991). Large differences in water transparency at all three sites masked seasonal variations in surface insolation as reflected in average diffuse attenuation coefficient (k) values ranging from 0.7 to 2.9 and differences in the maximum depth penetration ofH. wrightii, which varied from 0.6 to about 1.3 m. The continuous presence of a chrysophyte (brown tide) algal bloom in Laguna Madre since 1990 led to significant decreases in spring leaf elongation rates and a nearly 50% decline in below-ground biomass, which was reflected in root:shoot ratio (RSR) values that declined from 5.4 in 1989 to 2.3 in 1992. Increased turbidity and lower light levels in San Antonio Bay also corresponded with diminished plant biomass and the subsequent loss of plants; at both locations, the annual quantum flux ranged from 2200 to 2400 mol m^-2yr^-1, or about 18% of surface irradiance (SI). In contrast,H. wrightii populations growing at ca.1.2 m depths and characterized by high RSR values (4.0) were exposed to 5100 to 5700 mol m^-2yr^-1, or about 41 to 46% SI. Under these conditions, plants were exposed to daily saturating levels of PAR (H _sat) of 3 to 8 h during the spring/summer period of maximum growth, compared to an average of 2 h in Laguna Madre (after 1990) and San Antonio Bay based on field-derived measurements of photosynthetic parameters. Leaf elongation inH. wrightii exhibited a clear circannual rhythm at all sites, regardless of underwater light levels and therefore was not a sensitive indicator of light stress. Instead, chronic long-term reductions in underwater PAR were most strongly reflected in total plant biomass. The higher light demand (18% SI) forH. wrightii in relation to many other seagrasses (11% SI; Duarte 1991) may be related to its higher photosynthetic light requirement, but may also reflect the different methods used to evaluate the minimum light requirements of seagrasses. In estuarine and coastal waters, which are characterized by large and unpredictable variations in water transparency, continuous measurements of in situ PAR are invaluable in assessing the growth and photosynthetic response of seagrasses to variations in underwater irradiance.The University of Texas at Austin, Marine Science Institute Contribution No. 913     

Influence of temperature on the growth rate of Skeletonema costatum in response to variations in daily light intensity

Daily light intensities (I _o) can vary 10-fold during the winter-spring and late-summer diatom blooms in New England, USA, coastal waters. Laboratory cultures and natural populations incubated in dialysis sacs were examined to determine the time course of growth rate in Skeletonema costatum (Greville) Cleve in response to variations in daily light intensity during two bloom periods in Narragansett Bay, Rhode Island, USA. Log-phase cultures of S. costatum require 2 d to attain maximum growth rates at 2°C following transfer to saturating intensities. At 20°C, only 1 d is required. As temperature increases, Detonula confervacea (Cleve) Gran, Thalassiosira nordenskiöldii Cleve and Ditylum brightwellii (West) Grunow also exhibit rapid increases in mean daily division rates (K) following transfer to saturating light intensities. Thalassiosira pseudonana Hustedt, however, did not alter the time required to achieve maximum K as temperature varied. Natural populations of S. costatum did not show a well-defined relationship between K and light. Throughout a winterspring bloom, K was limited by low temperatures and exhibited no clear response to variations in I _o. A change in K in response to variation in I _o may occur on a daily basis during the summer, when temperatures are near 20°C; this has yet to be verified for in situ populations.     

Growth and competition of the marine diatoms Phaeodactylum tricornutum and Thalassiosira pseudonana. II. Light limitation

The marine diatoms Phaeodactylum tricornutum (Bohlin) and Thalassiosira pseudonana (Hasle and Heimdal) were grown under both continous illumination and a 14 h light: 10 h dark cycle at light intensities ranging from 1.53×10^-4 to 2.95×10^-1 ly min^-1. Under both photoperiods, T. pseudonana exhibited higher division rates than P. tricornutum at high light intensities, but the reverse was true at all light intensities <3×10^-3 ly min^-1. Comparison of these results with available data on light-limited growth of other planktonic algae suggests that P. tricornutum may be unusually efficient at maintaining its cell division rate at low light intensity. This efficiency may contribute substantially to its success in turbid, nutrient-enriched mass algal culture systems, the only environments in which it is known to attain great numbers.Contribution No. 4086 from the Woods Hole Oceanographic Institution.     

Light intensity and the feeding behaviour of herring,Clupea harengus

An infra-red sensitive video-recording technique was used to study the effects of darkness and light intensities from 0.0001 to 270 photopic lx on the feeding behaviour of herring (Clupea harengus L.). When offered natural zooplankton, consisting of a mixture ofCalanus finmarchicus, Euchaeta norvegica, Oithona similis, Balanus sp. nauplii, and crustacean nauplii as prey, the fish fed by biting (snapping) at light intensities above a threshold of 0.001 lx and were size-selective, taking the larger organisms first. When fed on pure cultures of CaliforniaArtemia sp. nauplii (San Francisco Bay brand), the threshold light intensity was 0.01 lx. Swimming speed increased with increasing light intensity when the fish were actively feeding by biting. When the fish were filter-feeding on high densities ofArtemia sp. nauplii in the light, they continued to school and swimming speed was not related to light intensity.     

Diel variation in chlorophyll a,carbohydrate and protein content of the marine diatom Skeletonema costatum

Skeletonema costatum (Greville) Cleve isolated from Narragansett Bay, USA, was incubated at 3 light intensities (ca. 0.008, 0.040 and 0.075 ly min^-1) under a 12 h light: 12 h dark (12L:12D) photoperiod at 2°, 10° and 20°C. Cellular chlorophyll a increased at intensities less than ca. 0.040 ly min^-1; increases occured within one photoperiod at temperatures above 10°C. Cellular carbohydrate increased with light intensity at all temperatures; increases during the photophase were due to net production of the dilute acid-soluble fraction. Cellular protein increased during the photoperiod at 10° and 20°C; there was little difference in cellular protein among all cultures after one photoperiod. The rate at which cellular chlorophyll a increased in response to a decrease in light suggests that diel variation in cellular chlorophyll a is temperature-dependent in S. costatum. Protein: carbohydrate ratios ranged from ca. 0.5 to 2.0 over a diel cycle; ratios increased at lower intensities and higher temperatures. The diel range in protein:carbohydrate ratios equals that in cultures developing nitrogen deficiency; thus, use of this ratio as an index to phytoplankton physiological state must account for diel light effects.     

Influence of light and temperature on the growth rate of estuarine benthic diatoms in culture

Four species of estuarine benthic diatoms: Amphiprora c. f. paludosa W. Smith, Nitzschia c. f. dissipata (Kützing) Grunow, Navicula arenaria Donkin, and Nitzschia sigma (Kützing) W. Smith were grown in unialgal cultures. The growth rates of the diatoms were determined as the rate of increase of the chlorophyll a content of the cultures. The diatoms were cultured at different combinations of temperture, daylength, and quantum irradiance. The highest growth rates of Navicula arenaria occurred at 16° to 20°C; the other 3 species had their optimum at 25°C or higher. The small-celled species had higher growth rates at their optimum temperature, but at lower temperatures the growth rates of all 4 species became very similar. The minimum daily quantum irradiance that could effect light-saturated growth at 12° and 20°C ranged from 2.5 to 5.0 E.m^-2.day^-1. At 12°C, two species had their highest growth rates under an 8 h daily photoperiod. At 20°C, the three species tested all had highest growth rates under 16 h daily photoperiod. The growth response of the benthic diatoms is comparable to that of several cultures of planktonic diatoms, as described in the literature. The influence of temperature and quantum irradiance on the diatoms in the present investigation was comparable to the influence of temperature and light intensity on the ¹⁴C-fixation of marine benthic diatoms (Colijn and van Buurt, 1975).     

Production dynamics of the eelgrass, <Emphasis Type="Italic">Zostera marina</Emphasis> in two bay systems on the south coast of the Korean peninsula

Production dynamics of eelgrass, Zostera marina was examined in two bay systems (Koje Bay and Kosung Bay) on the south coast of the Korean peninsula, where few seagrass studies have been conducted. Dramatically reduced eelgrass biomass and growth have been observed during summer period on the coast of Korea, and we hypothesized that the summer growth reduction is due to increased water temperature and/or reduced light and nutrient availabilities. Shoot density, biomass, morphological characteristics, leaf productivities, and tissue nutrient content of eelgrass were measured monthly from June 2001 to April 2003. Water column and sediment nutrient concentrations were also measured monthly, and water temperature and underwater irradiance were monitored continuously at seagrass canopy level. Eelgrass shoot density, biomass, and leaf productivities exhibited clear seasonal variations, which were strongly correlated with water temperature. Optimal water temperature for eelgrass growth in the present study sites was about 15–20°C during spring period, and eelgrass growths were inhibited at the water temperature above 20°C during summer. Daily maximum underwater photon flux density in the study sites was usually much higher than the light saturation point of Z. marina previously reported. Densities of each terminal, lateral, and reproductive shoot showed their unique seasonal peak. Seasonal trends of shoot densities suggest that new eelgrass shoots were created through formation of lateral shoots during spring and a part of the vegetative shoots was transformed into flowering shoots from March. Senescent reproductive shoots were detached around June, and contributed to reductions of shoot density and biomass during summer period. Ambient nutrient level appeared to provide an adequate reserve of nutrient for eelgrass growth throughout the experimental period. The relationships between eelgrass growth and water temperature suggested that rapid reductions of eelgrass biomass and growth during summer period on the south coast of the Korean peninsula were caused by high temperature inhibition effects on eelgrass growth during this season.     

Growth patterns of Codium fragile ssp. tomentosoides in response to temperature,irradiance, salinity,and nitrogen source

Seasonal patterns of growth, reproduction, and productivity of Codium fragile spp. tomentosoides (van Goor) Silva were monitored at 3 locations in Rhode Island. Maximal growth occurred during the summer and was more significantly correlated with temperature than any other factor measured in this study. Multiple correlation models suggested an interaction between temperature, irradiance, and available nitrogen. Maximal reproduction occurred in late summer and early fall. The maximal productivity, based on harvested quadrats, was 2. 10 g dry weight m^-2 day^-1. A large amount (up to 87.3%) of the annual production entered the detrital food chain during the winter by fragmentation of the thallus. Culture studies examined the effects of temperature (6° to 30°C), irradiance (7 to 140 E m^-2 sec^-1), daylength (8 h light: 16 h dark to 24 h light: O h dark) and salinity (6 to 48) on growth. Differentiated thalli grew over a broad range of experimental conditions, with maximal growth at 24°C, 24 to 30 S, a minimal irradiance of 28 E m^-2 sec^-1, and 16 h daylength. The effect of increasing daylength was due to increased total daily irradiance rather than to a true photoperiodic effect. Undifferentiated sporelings survived and grew in a narrower range of environmental conditions than thalli. c. fragile spp. tomentosoides grew equally well with nitrate, nitrite, ammonium, and urea as a nitrogen source. The addition of NaHCO₃ stimulated growth at levels of 2.4 to 4.8 mM, suggesting an inorganic carbon limitation in static cultures. This study supports the hypothesis that the in situ seasonal growth pattern of c. fragile spp. tomentosoides is primarily due to the interaction of temperature and irradiance.     

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.