Do correlations exist between chromatophore arrangement and photosynthetic activity in seaweeds?

In Dictyota dichotoma, as in many other plants, the chromatophores which at low intensities occupy the cell walls perpendicular to the light beam move to the side walls parallel to the light beam if exposed to high light intensities. The aim of this investigation was to find out whether or not the changes from low- to high-intensity arrangement and vice versa function as an active control mechanism to regulate photosynthetic activity in D. dichotoma under the respective light condition. Four different experimental approaches were made: (a) In white and blue light experiments the changes of the transmittance and of the rate of photosynthetic oxygen production in high- and low-intensity arrangement were compared. (b) The kinetics of the depression and recovery of the PS-rates, as well as of the transmittance changes, were determined during high- and low-intensity movement, respectively. (c) Transmittance and PS-rates of thalli under illumination with polarized and unpolarized light of the same intensity (1,000 1x) were compared. (d) PS-rates of thalli after darkening as well as after preirradiation with weak and strong red light, conditions under which the chromatophores occupy the same position in the cells, were measured. In all these experiments the photosynthetic activity was strongly influenced by pre-illumination, but was independent of the respective chromatophore arrangement. This finding was confirmed by experiments with two other algae: (1) In the brown alga Alaria esculenta which does not display light-induced chromatophore displacements and concomitant transmittance changes, pre-irradiation with high light intensities decreases the PS-rates. (2) In the green alga Ulva lactuca, which shows circadian chloroplast movements, the PS-rates depend on the pre-irradiation only, irrespective of the chloroplast position. Thus we may conclude that in these organisms the function of chromatophore displacements is not the regulation of photosynthetic activity. Other ecological functions are discussed.     

Effects of endogenous and some exogenous factors on the activity of the juvenile banana prawn Penaeus merguiensis

In conditions of alternating light and dark, juvenile Penaeus merguiensis de Man are more active during the dark phase. The rhythm persists in continuous dim red light, but not in continuous bright white light. The period of circadian rhythm shown in continuous dim red light varies between individuals from 22.75 to 26.0 h, with a mean of 23.8 h. The response of prawns to an artificially produced tidal situation is mediated by the presence or absence of water flow. No endogenous component of this tidal rhythm was demonstrated. There is an irregular short-term rhythm (period 2 to 3 h). It is suggested that this allows starved prawns to conserve energy.     

Photosynthesis and chlorophyll a fluorescence rhythms of marine phytoplankton

Circadian rhythms in photosynthesis were defined in field populations of phytoplankton. Measurements of carbon-dioxide fixation rates demonstrated that a diurnal periodicity of photosynthesis in samples incubated under natural light-dark (LD) cycles also were observed to continue in similar samples which had been photoadapted to constant dim light (LL) for 48 h. These changes in photosynthetic rates preceded sunset and sunrise, had daily amplitudes that ranged from 1.5 to 2.0, appeared to be independent of light-intensity, and displayed maxima about midday, while rates of dark fixation of carbon dioxide and the photosynthetic pigment content per cell were constant over the circadian cycle. Similar rhythmicity also was detected in room-temperature (22°C) chlorophyll a fluorescence yield, in both the obsence and presence of the photosynthesis inhibitor DCMU [3-(3,4-dichlorophenyl)-1, 1-dimethylurea]. However, the magnitude and timing of the fluorescence rhythm maxima seem to depend on wavelengths monitored and, in part, on the measuring technique used. Also, the circadian changes in the fluorescence intensity were abolished at low temperature (-60°C), and the shape of the emission spectra of chlorophyll fluorescence of cells in LD and LL did not change over time. The significance of the fluorescence rhythms with regard to chlorophyll a determinations and photosynthetic rates is discussed. It was concluded that there was sufficient similarity between circadian rhythms of photosynthesis in natural phytoplankton populations and in laboratory cultures of dinoflagellates to suggest that the mechanism of regulation may be the same for both of them.     

Diurnal and circadian rhythms in the turbidity of growing Skeletonema costatum cultures

Skeletonema costatum (Grev.) Cleve grown in batch culture at low light intensity under a 14 h light: 10 h dark photocycle showed exponential cell proliferation (1.1 doublings d^-1) without significant phasing of the cell division by the light: dark cycle. The growth in carbon concentration was, however, restricted to the light period. The turbidity of the culture closely followed the carbon oattern, and was not affected by the increase in the cell number during the dark period. It was found that a trustule suspension had only approximately 1% of the turbidity of the corresponding intact algae. Culture turbidity was therefore regarded as a biomass parameter similar to the carbon concentration, without direct correlation to the timing of the cell division. The short-time variations in the turbidity of growing algal cultures were further studied in a cage culture turbidostat. The growth rate (based on turbidity) increased rapidly during the first half of the light period, decreased slightly towards the evening and was zero throughout the dark period. When transformed to continuous light, the growth of the culture continued to show damped oscillations for up to 1 wk, but with a period of 26.7 h instead of 24 h. The same circadian rhythm was observed in chlorophyll content, and is thus possibly a reflection of a freely oscillating internal biological clock. The cage culture turbidostat was found to be a suitable device for studies of the photocycle related regulation of biosynthesis in S. costatum.     

Occurrence and biological role of light-induced chromatophore displacements in seaweeds

The occurrence of light-induced chromatophore displacements and concomitant transmittance changes in marine algae was investigated by microscope and photometrically with an automated recording microphotometer system; 16 brown, 6 green and 20 red algae were studied. In most of the brown algae, both phaeoplast displacements and transmittance changes were found. In some red algae which are frequently exposed to direct sun light during emergence at low tide, light-induced transmittance changes were measured, but they could not unequivocally be correlated with changes in the position of rhodoplasts. Among green algae, only Ulva lactuca shows chloroplast displacements which, however, follow circadian rhythms and are consequently not light-induced in the strict sense. The dose-response curves of light-induced chromatophore displacements were measured in the following Fucus and Laminaria species: F. spiralis, F. vesiculosus, F. serratus, L. digitata, L. saccharina and L. hyperborea. While in Fucus species correlations between light-induced transmittance changes and zonation of the intertidal area seem to exist, no significant differences have been found in the Laminaria species. The physiological role and ecological importance of light-induced chromatophore displacements for seaweeds living in the intertidal belt are discussed.     

Kinetics of light-intensity adaptation in a marine planktonic diatom

The marine planktonic diatom Thalassiosira weisflogii was grown in turbidostat culture under both continuous and 12 hL: 12 hD illumination regimes in order to study the kinetics of adaptation to growth-irradiance levels. In both illumination regimes adaptation to a higher growth-irradiance level was accompanied by an increase in cell division rates and a decrease in chlorophyll a cell^-1. The rates of adaptation for both processes, derived from first order kinetic analysis, equaled each other in each experiment. The results suggest that during the transition from low-to-high growth-irradiance levels chlorophyll a is diluted by cell division and is not actively degraded. Introduction of a light/dark cycle lowered the rate of adaptation. In transitions from high-to-low growth-irradiance levels there was a sharp drop in growth rates and a slow increase in chlorophyll a cell^-1 under both continuous and intermittent illumination. In the 12 hL:12hD cycle there was a circadian rhythm in chlorophyll a cell^-1, where cellular chlorophyll contents increased during the light cycle and decreased during the dark cycle. This circadian rhythm was distinctly different from light intensity adaptation. For kinetic analysis of light intensity adaptation in a 12 hL: 12 hD cycle, the circadian periodicity was separated from the light intensity response by subjecting the data to a Kaiser window optimization digital filter. Kinetic parameters for light-intensity adaptation were resolved from the filtered data. The kinetics of lightintensity adaptation of marine phytoplankton are discussed in relation to their spatial variations and time scales of mixing.This research was performed at Brookhaven National Laboratory under the auspices of the United States Department of Energy under Contract No. DE-AC02-76 CH00016     

Moon orientation in sandhoppers: effects of lighting treatments on the persistence of orientation ability

The moon orientation rhythm persists in sandhoppers removed from environmental entrainments relative to moon periodicity. In order to investigate the underlying mechanism, the crustacean amphipods Talorchestia capensis and Talitrus saltator were collected at new moon and exposed in the laboratory to an artificial light during the day, but to a variety of lighting treatments during the night: (1) continuous dark, (2) dim light in phase with the natural moon, (3) continuous dim light. The animals were tested under the moon 9–21 days later by using a new type of arena. The ability to orientate in a direction perpendicular to the shore persisted in sandhoppers under treatments 1 and 2, but not under treatment 3. A disturbance due to a phototactic tendency in Talorchestia was also observed in animals captured on the day of the test when exposed to unnatural lighting, but not when they were kept under natural light. The present findings show that the timing mechanism allowing compensation for changes in the moon’s position also persists in animals that have been long removed from entraining factors. The dependence of this orientation ability on nocturnal lighting and the disturbing effect of sudden changes in light intensity support the idea that the phasal lighting of the moon, and perhaps sunrise and sunset act as resetting factors for the moon’s orientation rhythm.     

Effects of light intensity on aging of the dinoflagellate Gonyaulax polyedra

Gonyaulax polyedra Stein grown in increasingly nutrientlimited batch culture undergoes the following changes (collectively termed aging): there is a decline in the intracellular concentrations of carbon, nitrogen and photosynthetic pigments; nitrate reductase activity decreases; rates of respiration and photosynthesis fall; and cell division virtually ceases (accompanied in bright light by a decrease in the volume of individual cells). The effect of light intensity on these aging events was tested by growing cells in either bright or dim light. The bright light (330 E m^-2 s^-1) was enough to saturate photosynthesis and the dim light (80 E m^-2 s^-1) was low enough to induce significant shade adaptation of photosynthesis without lowering growth rate. At both light intensities, a decline in carbon and nitrogen content preceded or accompanied all other monitored changes, and the sequence of aging events was similar. However the onset of the decline in intracellular nutrients and photosynthetic rate in low-light cells was delayed by a least one cell division time (i.e., to twice the cell density) in comparison to cells under bright light. At both light levels, pigment-protein complexes of the photosynthetic apparatus began to break down after intracellular carbon and nitrogen had been depleted to a critically low level. The beginning of the drop in pigmentation signalled the end of log-phase growth. It is suggested that the greater pigmentation of low-light cells may represent a larger nutrient supply than found in bright-light cells and could increase the survival time of nutrient-stressed populations.     

Factors affecting the photosynthetic capacity of laboratory cultures of the diatom Phaeodactylum tricornutum

The light-saturated photosynthetic capacity of cultures of Phaeodactylum tricornutum Bohlin grown under different conditions has been measured. In batch cultures grown in a regime of alternating light and dark periods, the photosynthetic capacity reaches a maximum before the end of the exponential phase of growth, and declines thereafter. In cultures illuminated at 0.7 mW (milliwatt)/cm², there is a 75% falloff in photosynthetic capacity per cell over an 8 day period following the time of maximum photosynthetic capacity. At 1.75 mW/cm², the corresponding fall-off is 85% over a 4 day period. Cultures exposed to a prolonged period of darkness (up to 16 days at 18°C) maintain a high photosynthetic capacity. Incubation in darkness also protects the cells from the deleterious effects of high temperature (28°C) upon photosynthetic capacity. The various fluctuations of photosynthetic capacity occur without any accompanying major changes in the concentration of chlorophyll a. Evidence from estimations of total protein and of the gross pattern of photosynthetic assimilation under different conditions suggests that the changes in photosynthetic capacity are largely controlled by the enzymic component of the photosynthetic machinery. By carefully controlling the conditions of dark incubation, the photosynthetic capacity can be reduced to a very low level without significantly affecting chlorophyll a concentration. Since the effect on photosynthetic capacity is reversible, it is possible to study aspects of chloroplast development without the complication of an associated synthesis of chlorophyll.     

Alteration of photosystem II activity by atrazine on Chlamydomonas reinhardtii synchronized and asynchronized cell cycle cultures

Inhibition of photosystem II (PSII) activity by atrazine was investigated in the green alga Chlamydomonas reinhardtii during different states of the cell cycle. The algal cultures were maintained under continuous light or under light/dark cycle (16/8?h) to obtain homogenized cell cycle distribution. The cycle state of algal population was determined by the DNA content using flow cytometry and defined as newly divided cells before the initiation of DNA replication (G₀/G₁) and cells at the end of the replication cycle with fully duplicated DNA content (G₂/M). Under different synchronized states of the cell population, the photosynthetic activity was investigated after treatment at 10, 100, and 1000?µmol?L^?1 atrazine exposed for 24?h by using fluorescence parameters related to PSII activity measured with a plant efficiency analyzer and pulse-amplitude modulated methods. In this study, we found that the atrazine effect was different depending on cell cycle phases and the period of illumination. Algal cells under light–dark cycle showed inhibition of the PSII electron transport leading to an increase of heat energy dissipation by the PSII reaction center. Algal cells grown under continuous light was shown to be more resistant to atrazine than the cells grown under light–dark cycle.     

Activity rhythms of two cirolanid isopods from an exposed microtidal sandy beach in Uruguay

Activity rhythms of two cirolanid isopods, Excirolana armata and Excirolana braziliensis, were studied based on both seasonal field observations and laboratory experiments, at an exposed microtidal sandy beach in Uruguay. The natural emergence patterns of both species were observed in the field for 1 year, twice in each season, and correlated to sea level, expected tidal cycles and diel cycles. Laboratory experiments were carried out in order to detect endogenous rhythms of activity and observe how emergence of both species was affected by changes in light and/or sediment thixotropy. We also compared behavioral strategies of sympatric species that occupy different beach levels. Sea level (and thus swash zone position) during field sampling did not follow expected tidal cycles for most sampling occasions. E. armata was observed in activity most of the time, but activity only correlated with sea level on three out of eight occasions, and only once was correlated to expected tidal cycle. Laboratory results showed that emergence under constant conditions was rare; changes in sediment thixotropy stimulated emergence, but the response was not cyclical; light had little effect on this response. On the other hand, E. braziliensis was fairly scarce in the water column, but swimming individuals were observed always during the night. They displayed an endogenous circadian activity pattern in the laboratory which augmented in response to changes in sediment thixotropy. The natural light/dark cycle modulated both spontaneous and response emergence by increasing day/night differences in activity. In this study E. armata, a midlittoral species more exposed to sea level variations, seemed to rely entirely on different physical and/or biological cues to trigger emergence at the appropriate time. E. braziliensis, found mostly in the upper intertidal zone, emerged in a circadian rhythm, which was stimulated by changes in sediment thixotropy and reinforced by light cycles. The results of this study led us to conclude that on microtidal, unpredictable beaches, local physical and biological factors can combine to determine different activity strategies in organisms from different intertidal levels. Received: 23 March 2000 / Accepted: 30 August 2000     

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.     

Effects of non-circadian light/dark cycles on the growth and moulting of Palaemon elegans reared in the laboratory

The growth and moulting of Palaemon elegans Rathke has been compared under a circadian (12 h light: 12 h dark) and two non-circadian (8 h light:8 h dark and random light:dark) light/dark cycles. In prawns reared individually from hatching to the late juvenile phase, growth, measured as increase in total length, was significantly retarded in the non-circadian regimes, during zoeal, post-larval and early juvenile development. This effect was primarily the result of reduced increments at moulting in the non-circadian regimes. Growth of prawns reared from hatching to the post-larval phase, measured as wet and dry weights, was similarly reduced in a non-circadian regime. These effects support the hypothesis that the integrity of an animal's physiology is partially dependent on maintaining, through the action of daily environmental cycles, correct timing relationships between its oscillating sub-systems.     

The role of the queen in circadian rhythms of honeybees (Apis mellifera L.)

Summary Colonies and smaller social groups of honeybees (Apis mellifera carnica L.) show distinct free-running circadian rhythms similar to that of individual organisms. The workers of a colony synchronize their individual rhythms to one overall group rhythm. Caste plays an important role in this synchronization process. Queens were introduced into worker groups which were entrained to a phase-shifted light/dark cycle. The introduction of the queen caused a shift in the free-running phase under constant dark conditions. Single introduced workers had no effect on the free-running rhythms. This indicates that the queen plays an important role in the synchronization of circadian rhythms of honeybee colonies.     

Carbon and nitrogen metabolism by Monochrysis lutheri: Measurement of growth-rate-dependent respiration rates

Dark respiration rates were measured and carbon-excretion rates calculated for a nitrate-limited population of the marine chrysophyte Monochrysis lutheri grown in continuous culture at 20°C on a 12 h light-12 h dark cycle of illumination and over a series of 4 growth rates. A significant (P<0.05) positive correlation was found between dark respiration rate and growth rate. From a simple linear fit to the data, the respiration rate at maximum growth rate was estimated to be roughly 10.5% of the maximum gross-carbon-production rate, and more than three times higher than the extrapolated respiration rate at zero net-growth rate. Carbon-excretion rates showed no significant correlation with growth rate, and averaged less than 5% of the maximum gross-carbon-production rate. Mean cell nitrogen to carbon ratios were correlated in a virtually linear manner (r=0.994) with growth rate, and at a given growth rate were consistently higher than nitrogen to carbon ratios for the same species grown on continuous light. A comparison of carbon and nitrogen quotas as a function of growth rate for M. lutheri and other species suggests that the increase of cellular nitrogen at high growth rates under nitrate-limited growth conditions may be associated with the storage of cellular protein or amino acids rather than the presence of an inorganic nitrogen reservoir. The maximum nitrate uptake rate per cell during the day changed very little over the range of growth rates studied, and was comparable to the maximum uptake rate found for cells grown on continuous light. However, the cell nitrogen quota increased steadily with growth rate, causing a reduction in the maximum specific-uptake rate of nitrate during the day at high growth rates. The dark nitrate-uptake capacity of the population was clearly exceeded by the supply rate at the two higher growth rates, leading to a buildup of nitrate during the night which amounted to as much as 21% of the particulate nitrogen in the growth chamber by morning.Hawaii Institute of Marine Biology Contribution No. 478.     

Action spectra and spectral quantum yield of photosynthesis in marine macroalgae with thin and thick thalli

Net photosynthesis at 10mol photons m^-2 s^-1 in each of 24 wavelengths was measured in absolute units by an O₂-electrode and corrected for dark respiration to construct action spectra for gross photosynthesis in nine species of algae, which included plants with thin and thick thalli from each of four major pigment groups. The photosynthesis of green and brown algae with thin thalli decreased in green light, but species with thick thalli from these two groups had action spectra which were almost flat, and matched the optical blackness of the thalli but did not reflect the pigment differences between the species. Among the red algae, on the other hand, there was little difference between the action spectra for thin and thick algae. Only wavelengths absorbed by the phycobilin pigments were effective in photosynthesis, even in species (e.g. Chondrus, Phyllophora) which absorbed all visible wavelengths strongly. Maximal quantum yields of 0.10 to 0.12 O₂ molecules per absorbed photon were recorded for thin green and brown algae, but thicker algae in these two groups had lower values. Red algae exhibited maximal values close to 0.10 O₂ molecules per absorbed photon, irrespective of thallus thickness or phycocyanin content, but the quantum yields of phycoerythrin-rich species in the 600 to 650 nm waveband were lower than those of phycocyanin-rich species.     

Temporal adaptations in visual systems of deep-sea crustaceans

The temporal characteristics of the visual systems of deep-sea crustaceans were examined at varying light levels. Experimental organisms were collected off Hawaii and southern California in 1991 and 1992, and continually maintained in the dark. At the University of California Santa Barbara Marine Laboratory, the temporal components of both visual interneuron activity and electroretinograms (ERGs) of adult Gnathophausia ingens were tested at threshold and with background light levels up to 6 log units above threshold. To compare these responses with those of mid-water organisms inhabiting shallower depths, the temporal characteristics of the ERGs of Oplophorus spinosus and juvenile G. ingens were assessed. Adults of both species showed little or no change in the time components of their responses when light-adapted, except for the response latency. There was also no evidence of a circadian rhythm in visual sensitivity. O. spinosus, a vertical migrator, and juvenile G. ingens, which inhabit a shallower depth than the adults, exhibited more rapid responses than adult G. ingens. These varying responses are considered adaptations to the differences in light levels at the characteristic depths of the organisms.     

DNA synthesis and the cell cycle in the noxious red-tide dinoflagellateGymnodinium nagasakiense

Gymnodinium nagasakiense is a noxious red tide dinoflagellate often associated with damage to fisheries in Japan. DNA synthesis and the cell cycle in this organism were investigated from 1989 to 1990 by determining relative DNA contents of individual cells using an epifluorescence microscopy-based microfluorometry system. The nuclei were stained with the DNA-specific fluorochrome 4-6-diamidino-2-phenylindole (DAPI). Because photosynthetic pigment interferes with the fluorescence from the DAPI-DNA complex, the pigment was eliminated by methanol treatment as a first step in quantitative microfluorometry. Nuclear DNA contents, cell size distribution, cell density, and frequency of paired cells were determined every 2h for 24h using cells grown on a 12h light:dark cycle. DNA synthesis and cell division were tightly phased to a particular period of the light:dark cycle. DNA synthesis (S phase) occurred from 10:00 to 22:00 hrs and was followed by cytokinesis. The presence of such a distinct S phase strongly suggests thatG. nagasakiense has a typical eukaryotic cell cycle. This type of cell cycle makes it possible to estimate speciesspecific in situ growth rate based on the diel pattern of DNA synthesis.     

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.     

Calcium exchange and the measurement of calcification rates in the calcareous coralline red alga Amphiroa foliacea

⁴⁵Ca washout data of living thalli of Amphiroa foliacea Lamouroux in the light and dark show that there are three kinetically distinct Ca²⁺-exchanging compartments with approximate half-times (t 1/2) of 300, 20 and 3 min. The two slower compartments appear to be exchange from organic Ca²⁺-binding components of the cell wall, while the fast compartment probably represents exchange on the CaCO₃ crystal surface. Killed and decalcified thalli have a fourth compartment, with a t 1/2 of 20 to 35 min (other compartment half-times are 300, 40, 3 min), which has been identified as the greatly increased intercellular space produced during drying and decalcification. The ⁴⁵Ca and ¹⁴C uptake data show that a large proportion of the label initially taken up is into compartments other than the CaCO₃. As a result of this uptake, binding, and exchange of radioisotope, significant errors occur during the measurement of calcification rates, unless a kinetic analysis is carried out. Using such a technique, CaCO₃ calcification rates of A. folicea were measured with ⁴⁵Ca or ¹⁴C as tracers. Light stimulates calcification by up to 2.6 times, depending upon the age of the plant. Young segments have a markedly higher rate of calcification and photosythesis than do the older segments.