Light responses of a scleractinian coral (Plerogyra sinuosa)

During daytime Plerogyra sinuosa Dana displays globular expandable tentacles (bubbles) which foster the photosynthetic ability of the coral. Adaptational responses of this coral to different depths (5–25 m) and light conditions were investigated by photosynthetic pigment analysis, insitu measurements of oxygen production, transplantation and shading experiments. Pigment concentrations per unit tissue dry weight were variable, but unrelated to depth. Pigment concentrations per zooxanthellae cell remained constant and bubble size increased with depth. Light intensity at 25 m was 20 to 25% of the 5-m value, but daily integrated rates of photosynthesis were 65% of the 5-m rates, indicating a higher light utilization efficiency in deeper corals. Coral heads transplanted from 25 to 5 m died within 20 d if not protected against UV-radiation, but corals transplanted from 5 to 25 m acclimatized to the new light condition. Photosynthetic oxygen production and bubble size increased in shaded, sun-adapted corals within 60 min and decreased in sun-exposed, shade-adapted corals. The variable bubble size is interpreted as an adaptational mechanism to optimize light exposure of zooxanthellae.     

UV-absorbing substances in zooxanthellate and azooxanthellate clams

The effects of UV-A and UV-B radiation on photosynthesis of zooxanthellae within the siphonal mantle of the giant clam, Tridacna crocea, and in isolation were studied. While UV-B irradiation (2.4 W m⁻², 20 min) completely suppressed photosynthesis of the isolated zooxanthellae, it had little effect on their photosynthetic ability if they were irradiated while within the siphonal mantle of the host tissue. Chemical analysis of the siphonal mantle of T. crocea showed the presence of significant amounts of mycosporine-like amino acids (MAAs), which absorb UV-A and -B light. However, no MAA was detected in the isolated zooxanthellae. MAAs were concentrated in the siphonal mantle and kidney tissues in comparison with other tissues. In the siphonal mantle, MAA concentrations were the highest in the outermost surface layer where most of the zooxanthella cells resided. This indicates that the zooxanthellae are protected from UV radiation by a screen of concentrated MAAs in the host clam. Aside from T. crocea, significant amounts of MAAs were found not only in other zooxanthellate clams, such as T. derasa, Hippopus hippopus, Colculum cardissa and Fragum unedo, but also in a closely related azooxanthellate clam, Vasticardium subrugosum. On the other hand, no MAA was detected in any of the zooxanthellae from these zooxanthellate clams. No MAA was detected in the tissues of a deep-sea bivalve, Calyptogena soyoae. Although MAAs seem to block strong UV radiation in the shallow-water clam, they are probably not essential for the clam's life in the dark. MAAs in shallow-water clams may be derived from food and accumulated in their tissues, especially in the siphonal mantle and kidney. Received: 29 November 1996 / Accepted: 13 January 1997     

Two new UV-absorbing mycosporine-like amino acids from the sea anemoneAnthopleura elegantissima and the effects of zooxanthellae and spectral irradiance on chemical composition and content

Many tropical cnidarians living in shallow water contain a class of ultraviolet-A (UV-A, 320 to 400 nm) and ultraviolet-B (UV-B, 280 to 320 nm) absorbing compounds known as mycosporine-like amino acids (MAAs). These compounds may provide protection from the deleterious effects of solar UV radiation. Using a novel application of reverse-phase high performance liquid chromatography, we find that the temperate sea anemoneAnthopleura elegantissima (collected in 1988 from Bodega Bay, California, and in 1991 from Santa Barbara, California) contains four major MAAs: shinorine, porphyra-334, and two new compounds, mycosporine-taurine and mycosporine-2 glycine. Analysis of zooxanthellate (containing zooxanthellae) and naturally apozooxanthellate (lacking zooxanthellae) specimens acclimated in the presence and absence of UV for 28 d in the spring of 1988 suggests that this anemone, unlike some other anthozoans, does not regulate the concentration of its MAAs in response to UV radiation. The presence of similar concentration of MAAs in apozooxanthellate and zooxanthellate specimens indicates that symbiosis with algae is not required for these compounds to be present in the anemone. The total concentration of MAAs in the zooxanthellae is only about 12% of that in their host's tissues.     

Effects of ambient levels of solar ultraviolet radiation on zooxanthellae and photosynthesis of the reef coral Montipora verrucosa

Paired flat plates of the hermatypic coral Montipora verrucosa from Kaneohe Bay, Oahu, Hawaii, were acclimated to photosynthetically active radiation (PAR) only and to full sunlight (PAR+UV) for several weeks in the summer of 1990. After the acclimation period, photosynthesis, both in PAR-only and PAR+UV as well as dark respiration were measured. Levels of the UV-absorbing compounds, S320, density of zooxanthellae, and chlorophyll a concentration were determined. Corals acclimated in PAR+UV had higher levels of the UV-protective compounds and lower areal zooxanthellae densities than corals acclimated in PAR-only. Chlorophyll a per unit volume of coral host and per algal cell did not differ between corals from the two acclimation treatments. Corals acclimated to PAR+UV displayed higher photosynthesis in full sunlight than corals acclimated to PAR-only, but when photosynthesis was measured in the light regime to which the corals had been acclimated, there were no differences in photosynthesis. Dark respiration was the same for corals from the two acclimation treatments regardless of the light quality immediately preceding the dark period.Contribution No. 902 HIMB     

Effects of irradiance and ultraviolet radiation on photoadaptation in the zooxanthellae of Aiptasia pallida: primary production,photoinhibition, and enzymic defenses against oxygen toxicity

Cnidarians which contain symbiotic algae are constantly faced with the challenges of a changing photic regime and a hyperoxic environment. Zooxanthellae (Symbiodinium sp.) from the sea anemone Aiptasia pallida (Verrill), collected and cultured at Bermuda Biological Station in 1986, exhibit a suite of compensatory responses to changes in irradiance, ultraviolet radiation (UV), and to the toxicity resulting from their interaction with photosynthetically produced oxygen. Superoxide dismutase (SOD) and catalase inactivate superoxide radicals (O₂ ^-) and hydrogen peroxide (H₂O₂), which are mediators of oxygen toxicity, show an increase in specific activity with irradiance and in response to UV, both in cultured zooxanthellae (CZ) and freshly isolated zooxanthellae (FIZ) from acclimated anemones. CZ and FIZ exposed to environmentally realistic UV levels show a 30 to 40% increase in SOD activities compared with zooxanthellae exposed to similar irradiances without UV. CZ consistently show higher activities of both SOD and catalase compared to FIZ. Both CZ and FIZ exhibit changes in chlorophyll content and in the relationship between photosynthesis and irradiance which suggest photoadaptive changes in CO₂-fixing enzymes, the photosynthetic-electron transport system, or in photosynthetic unit size (PSU). UV has a greater effect on the photosynthetic capacity (P _max) of FIZ when compared to CZ acclimated at an equivalent irradiance with or without a UV component. UV also enhances the photoinhibition observed at high irradiance in both CZ and FIZ. Differences in enzyme activity between CZ and FIZ suggest an important role for the host in the protection of zooxanthellae against the direct effects of environmentally realistic UV while the photosynthetic performance of zooxanthellae in situ may not be as well protected.     

Experimental ecology of the temperate scleractinian coral Astrangia danae

The combined effects of temperature, light and symbiont density on the metabolic rate and calcification of the temperate coral Astrangia danae were studied experimentally using colonies containing different concentrations of zooxanthellae. After acclimation to five temperatures between 6.5° and 27°C, and incubation at three light levels and in darkness, respiration and photosynthesis were measured and corrected for rates due to commensals alone. Calcification rates were regressed on zooxanthellae concentration and production in order to define “symbiotic” and “non-symbiotic” averages, and the enhancement of calcification by symbiotic interactions in the polyps. Respiration by the polyparium varied less with temperature between 11.5° and 23°C than that of the commensals, suggesting physiological acclimation by the coral tissue. In-vivo zooxanthellae photosynthesis increased linearly with temperature and was near its maximum at 400 μEin m^?2 s^?1, but the photosynthesis of the endolithic algae of the corallum varied little between 11.5° and 27°C. Calcification at any given temperature was near its maximum at 40 μEin m^?2 s^?1 in both symbiotic and non-symbiotic corals. CaCO₃ deposition increased linearly with temperature in non-symbiotic colonies and in symbiotic colonies incubated in the dark. In symbiotic colonies, calcification in the light increased above these basic rates as temperature rose above 15°C. Below 15°C, symbiotic interactions failed to stimulate calcification, apparently due both to a lowering of zooxanthellae photosynthesis and to a decrease in the enhancing effect of any given level of primary production.     

Light absorption and utilization among hermatypic corals: a study in Jamaica,West Indies

The chlorophyll specific absorption coefficient ( _c) was measured for zooxanthellae from six hermatypic coral species obtained, where possible, from four depths (1, 10, 30, 50 m) on reef sites near Discovery Bay, Jamaica in February and March 1983. Measurements of photosynthetic rates versus irradiance, as well as cellular and areal chlorophyll a, were also performed on these colonies or sister colonies. Together the data were used to compare minimum quantum requirements (1/Φ m) among species and depths and to assess the importance of light utilization to the growth and depth distribution of these corals. Our data suggest that, although _c was found to decrease with depth, interspecific differences in _c do not occur for zooxanthellae from the corals investigated. Minimum quantum requirements (1/Φ m) decreased significantly with depth, thereby reflecting an increase in photosynthetic light utilization efficiency with decreasing irradiance. Interspecific differences in 1/Φ m determinations were suggested but not statistically conclusive. We conclude that interspecific differences in gross photosynthesis, and perhaps growth and depth distribution, are primarily attributable to differences in the light utilization capacity of the whole coral, as reflected by the product of _c ^′ and chlorophyll per unit surface area, and in-situ quantum efficiencies. This research was performed under the auspices of the US Department of Energy under Contract No. DE-AC02-76CH00016     

Effects of dissolved ammonium addition and host feeding with Artemia salina on photoacclimation of the hermatypic coral Stylophora pistillata

 Effects of nutrient treatments on photoacclimation of the hermatypic coral Stylophora pistillata (Esper) were studied. Studies on photoacclimation of colonies from different light regimes in the field were evaluated and used to design laboratory experiments. Coral colonies were collected in the Gulf of Eilat (Israel) from January to March 1993. Exterior branches of colonies from different depths (1 to 40 m) displayed different trends in production characteristics at reduced and very low levels of illumination. From 24 ± 3% to 12 ± 2% of incident surface photosynthetic active radiation (PAR_o), zooxanthella population density and chlorophyll a+c per 10⁶ zooxanthellae increased, a trend seen in the range of light levels optimal for coral growth (90 to 30% PAR_o). The P _max of CO₂ per 10⁶ zooxanthellae decreased, while P _max of CO₂ per 10³ polyps increased, indicating an increase in zooxanthella population density at low light levels. Proliferous zooxanthella frequency (PZF, a measure of zooxanthella division) declined significantly at light levels <18 ± 3% PAR_o. At the lowest levels of illumination (<5% PAR_o), zooxanthella population density decreased, as did the PZF; chl a+c per 10⁶ zooxanthellae was unchanged. In 28-d experiments, exterior coral branches from the upper surfaces of colonies from 3 m depth (65 ± 4% PAR_o) were incubated in aquaria under bright (80 to 90% PAR_o), reduced (20 to 30% PAR_o), and extremely low (2 to 4% PAR_o) light intensities. At each light intensity, the corals were maintained in three feeding treatments: sea water (SW); ammonium enriched SW (SW + N); SW with Artemia salina nauplii (SW + A). An increase in P _max of CO₂ per 10³ polyps was found in corals acclimated to reduced light (20 to 30% PAR_o) in nutrient-enriched SW, while in SW, where the increase in zooxanthella population density was smaller, it did not occur. Nutrient enrichments (SW + N at 2 to 4% PAR_o and SW + A at 20 to 30% PAR_o) increased zooxanthella population density, but had no effect on chl a+c per 10⁶ zooxanthellae. Acclimation for 14 d to reduced (10 to 20% PAR_o) and extremely low (1 to 3% PAR_o) light intensities shifted ¹⁴C photoassimilation into glycerol and other compounds (probably glycerides), rather than sugars. Both ammonium addition and feeding with Artemia salina nauplii resulted in an increase in photosynthetic assimilation of ¹⁴C into amino acids. We conclude that acclimation to reduced light consists of two processes: an increase in photosynthetic pigments and in zooxanthella population density. Both processes require nitrogen, the increase in zooxanthella population density needing more; this adaptation is therefore limited in nitrogen-poor sea water. Received: 19 June 1998 / Accepted: 13 June 2000     

Mode of zooxanthella transmission does not affect zooxanthella diversity in acroporid corals

Two distinct modes of algal endosymbiont acquisition exist in corals, a direct transmission from the parental colony to the eggs and a larval or post-larval uptake from the environment. The former, maternal-transmission mode is expected to be a more closed system, while the latter is believed to be an open system. Here we test the hypothesis that the diversity of symbionts in closed systems is lower than that in open systems. We examine the identity and diversity of the algal endosymbionts (zooxanthellae) in 25 Montipora species sampled from Irian Jaya (Indonesia) and Magnetic Island (central Great Barrier Reef) and compare the results with those previously obtained from Acropora species, which belong to the same family. All Montipora colonies examined harbour clade C zooxanthellae, with two colonies harbouring both clade C and D zooxanthellae simultaneously. Two algal strains (named C· and D· in this study) present in Montipora have not been observed in Acropora, and may have co-evolved with Montipora. Symbiodinium C· shows approximately 5% sequence divergence from C strains observed in Acropora spp. and occurs in 76% of the colonies examined. Nevertheless, several other C strains commonly found in other corals occur in some of the Montipora colonies. Montipora species transmit their algal endosymbionts directly to the eggs, while Acropora species have to acquire zooxanthellae from the environment every generation. Contrary to our expectations, the diversity of zooxanthellae is similar for the two genera, indicating that the mode of symbiont transmission (i.e. maternal versus horizontal) does not affect symbiont diversity in acroporid corals.Communicated by M.S. Johnson, Crawley     

Experimental ecology of the temperate scleractinian coral Astrangia danae I. Partition of respiration,photosynthesis and calcification between host and symbionts

Calcification, photosynthesis and respiration of the scleractinian coral Astrangia danae were calculated from the changes in total alkalinity, pH, calculated total CO₂, and oxygen concentration produced by colonies incubated in glass jars. A correction for changes in ammonia, nitrate and nitrite was taken into account and the method evaluated. The fluxes of oxygen and CO₂ were highly correlated (r=0.99). The statistical error of alkalinity determinations was less than 10% of the changes observed in the slowest calcifying samples. Metabolism of polyparium alone was estimated by difference after removal of tissue and reincubation of bare corallum. Zooxanthellae concentration in the polyps was obtained from cell counts made on homogenates of polyp tissue. The calculated photosynthetic rate of the zooxanthellae in vivo was 25 mol O₂ (10⁸ cell)^-1 h^-1 at a light intensity of 120 Ein m^-2 s^-1. In corals having 0.5x10⁹ zooxanthellae/dm² of colony area up to 8% of the total photosynthesis was attributed to the corallum microcosm. Polyp respiration, photosynthesis, and CaCO₃ uptake rates were all much higher than rates previously reported from A. danae, apparently because in these experiments the organisms were better fed. This increased photosynthesis in turn enhanced calcification still further. The symbiosis therefore appears to provide a growth advantage even to fed corals, under the conditions of these experiments.     

Modelling the photosynthetic production by sponges on Davies Reef,Great Barrier Reef

Sponge populations on Australia's Great Barrier Reef (GBR) may contain a mix of both phototrophic and heterotrophic species. The distribution of many of these sponges on reefs is assumed to be determined by light. A model was developed to investigate how the distribution of phototrophic sponges over depth is restricted by the availability of photosynthetically active radiation. Estimates of the balance between photosynthetic production and the total respiratory demand of entire sponge communities on Davies Reef (a middle-shelf reef of the Great Barrier Reef) are provided. These estimates are based upon published data for community composition and biomass, whilst photokinetic parameters have been determined for a variety of sponge species from oxygenexchange measurements. Phototrophic sponges on the fore-reef slope are predicted to exist at or above a state of net 24 h compensation (i.e., photosynthetic oxygen production by sponges balances or exceeds respiration over a 24 h period) to a depth of 30 m. It is proposed that phototrophic sponges are obligate phototrophs because the availability of light for photosynthesis corresponds with the lower depth limit of their distribution. Sponge communities (including both phototrophs and heterotrophs) from the fore-reef and lagoon exist close to a state of net 24 h compensation to a depth of 10 to 15 m. This balance shows diurnal variations, associated with the activity of phototrophs, such that instantaneous compensation of the community may occur to depths of 20 to 25 m when light is maximal.     

Pink-line syndrome, a physiological crisis in the scleractinian coral Porites lutea

Coral diseases are one of the major factors that alter coral cover and their diversity. We have earlier reported the “Pink-line syndrome” (PLS) in the scleractinian coral Porites lutea wherein a colored band appears between the dead and healthy tissue of a colony. About 20% of the P. lutea colonies were affected in Kavaratti of the Lakshadweep Islands in the Arabian Sea during April 1996 and the incidence increased fourfold within the next 4 years. Fungi were associated in both PLS-affected and healthy specimens, whereas the cyanobacterium Phormidium valderianum occurred exclusively in the PLS-affected specimens. There was an increased expression of a 29 kDa protein without any significant increase in total protein content in the PLS-affected colonies. A reduced number of zooxanthellae and an increase in zooxanthellae size, mitotic index, and chl a concentrations were some of the characteristics of the PLS-affected colonies. PLS induction experiments conducted using selected fungi and the cyanobacterium P. valderianum isolated from the affected colonies and abiotic factors, such as CO₂ enrichment and the effect of cyanobacterial photosynthesis inhibition, indicated that the CO₂ build-up around the host tissue caused the pink coloration. We hypothesize that these physiological changes disturb the mutualism between the zooxanthellae and the host. When the symbiosis is disturbed by the external CO₂, the host loses control over the zooxanthellae, causing their uncontrolled division. This process may lead to a break in photosynthate transfer to the host, thereby resulting in starvation and finally leading to partial mortality. We further hypothesize that these degenerative processes are triggered by the CO₂ produced by P. valderianum through its carbon concentration mechanism. In this context, any opportunistic cyanobacteria or other agents having potential to interfere with the physiology of the host or the symbiont can cause such a physiological disorder. The mechanism of PLS formation is an early warning to protect corals as the increasing atmospheric CO₂ could induce PLS-like physiological disorder in corals.     

Resource partitioning by reef corals as determined from stable isotope composition

The pattern of resource partitioning vs depth by corals collected in February 1983 from Jamaica and the Red Sea was determined from their stable carbon isotope composition. Observations were made on isolated zooxanthellae and corresponding algae-free animal tissue from eight species at four depths over a 50 m bathymetric range. Zooxanthellae ¹³C was high in shallow water and became lower as depth increased. This trend correlated significantly with the anual integrated photosynthetic rate. The trend is interpreted according to a depletion-diffusion hypothesis; in shallow water, at high rates of photosynthesis, metabolic CO₂ is nearly depleted and the supply of CO₂ from seawater bicarbonate is limited by diffusion. Since most of the available CO₂ is fixed, isotope fractionation is minimal. In deeper water, at lower rates of photosynthesis, metabolic CO₂ is ample, and isotope fractionation is greater. Animal tissue ¹³C was slightly lower than corresponding zooxanthellae values in shallow water. As depth increased the difference between zooxanthellae and animal tissue ¹³C increased and the latter approached the ¹³C of oceanic particulate organic carbon. These data suggest that carbon is translocated at all depths and that deep-water corals draw significantly on allocthonous sources of carbon.Contribution No. 436 of the Discovery Bay Marine Laboratory of the University of the West Indies     

The effects of antifoulant-paint-contaminated sediments on coral recruits and branchlets

The 184-m cargo ship "Bunga Teratai Satu" ran aground on Sudbury Reef, within the Great Barrier Reef Marine Park, on 2 November 2000. Although no cargo or fuel was lost, the ship remained aground for 12 days and a large quantity of antifoulant paint containing tributyltin (TBT), zinc, and copper was scraped from the hull during the grounding and subsequent refloating operation. This resulted in extensive contamination of the reef sediments for up to 250 m surrounding the grounding site. Two laboratory-based experiments assessed the impact of contaminated sediments on the survival of both newly settled corals of Acropora microphthalma and branchlets of A. formosa. Newly settled corals exposed to sediments containing 8.0 mg kg^–1 TBT, 72 mg kg^–1 Cu, and 92 mg kg^–1 Zn or greater suffered significantly higher mortality after 72 h, compared to control or low-concentration treatments. Coral recruits exposed to 40 mg kg^–1 TBT (Sn), 306 mg kg^–1 Cu, and 403 mg kg^–1 Zn were all killed within 38 h. Branchlets from adult corals exposed to sediments with a high concentration of contaminants (TBT 160 mg kg^–1, Cu 1,180 mg kg^–1, and Zn 1,570 mg kg^–1) suffered significant mortality (38%), whereas branchlets placed in treatments with lower levels of contaminants suffered no mortality. Visual bleaching of the branchlets was observed at high contaminant levels, but an overall reduction in the symbiotic zooxanthellae populations was not observed in surviving corals. The photosynthetic yields of light-adapted zooxanthellae remained constant in live branchlets, indicating that the TBT-contaminated sediment may be more toxic to the host than the symbiont. Our results show that antifoulant contamination at ship-grounding sites has the potential to cause major mortality of resident coral communities and can have a negative impact on the recovery of adult populations.Communicated by P.W. Sammarco, Chauvin     

Effects of feeding frequency and symbiosis with zooxanthellae on nitrogen metabolism and respiration of the coral Astrangia danae

Colonies of the temperate coral Astrangia danae occur naturally with and without zooxanthellae. Basal nitrogen excretion rates of nonsymbiotic colonies increased with increasing feeding frequency [average excretion rate was 635 ng-at N (mg-at tissue-N)^-1 h^-1]. Reduced excretion rates of symbiotic colonies were attributed to N uptake by the zooxanthellae. Nitrogen uptake rates of the zooxanthellae averaged 8 ng-at N (10⁶ cells)^-1 h^-1 in the dark and 21 ng-at N (10⁶ cells)^-1 h^-1 at 200 Ein m^-2 s^-1. At these rates the zooxanthellae could provide 54% of the daily basal N requirement of the coral if all of the recycled N was translocated. Basal respiration rates were 172 nmol O₂ cm^-2 h^-1 for starved colonies and 447 nmol O₂ cm^-2 h^-1 for colonies fed three times per week. There were no significant differences between respiration rates of symbiotic and nonsymbiotic colonies. N excretion and respiration rates of fed (symbiotic and nonsymbiotic) colonies increased greatly soon after feeding. N absorption efficiencies decreased with increasing feeding frequency. A N mass balance, constructed for hypothetical situations of nonsymbiotic and symbiotic (3×10⁶ zooxanthellae cm^-2) colonies, starved and fed 15 g-at N cm^-2wk^-1, showed that the presence of symbionts could double the N growth rate of feeding colonies, and reduce the turnover-time of starved ones, but could not provide all of the N requirements of starved colonies. Rates of secondary production, estimated from rates of photosynthesis and respiration were similar to those estimated for reef corals.     

Tolerance of estuarine benthic diatoms to high concentrations of ammonia,nitrite ion,nitrate ion and orthophosphate

A carbon-14 assimilation method was used to determine action spectra and photosynthesis versus irradiance (P versus I) curves of natural populations of phytoplankton and zooxanthellae from a coral reef fringing Lizard Island in the Australian Barrier Reef. The action spectra were related to the phytoplankton species composition. The curves showed shade adaptation in phytoplankton from deeper waters and in the zooxanthellae. Rates of photosynthesis of zooxanthellae were shown to be highly but variably dependent on their host organisms. Photosynthetic production by zooxanthellae was about 0.9 gC m^-2 day^-1, which is about three times higher than phytoplankton production in the waters close to the reef.     

Adaptation of solitary corals and their zooxanthellae to low light and UV radiation

Adaptation of solitary corals, Fungia repanda and F. echinata, and their zooxanthellae to low light and ultraviolet light B (UV-B) was studied with respect to changes in their protein contents, photosynthetic pigment contents and the photosynthesis-irradiance (P-I) curves. The corals were collected from 1 to 50 m depths in the Republic of Belau (Paulau) in 1990 and 1991. The chlorophyll a content in a unit surface area of the coral did not change significantly with the depth of the habitat, whereas cellular chlorophyll a in the algae increased with the depth. Zooxanthellae density and protein content in a unit surface area of Fungia spp. decreased with the depth. Photosynthetic parameters normalized by a unit surface area of the Fungia spp., maximum gross photosynthetic rate (P _gmax area^-1) and dark respiration rate (R area^-1), were negatively correlated with the depth, while initial slope of the P-I curve () did not show significant correlation with the depth. Compensation light intensity (Ic) decreased with the depth. In isolated zooxanthellae, P _max chl a ^-1, and R chl a ^-1 decreased with the depth, while _{chl a} was constant. P _gmax cell^-1 and R cell^-1 did not change significantly but _cell increased with the depth. Ic decreased with the depth as in the intact corals. Reduction of protein content in a unit area of the coral from deeper habitat implies decrease of host animal tissues. Reduction of Ic can be explained by decrease of R area^-1, which may be due to the diminution of animal tissues. The photoadaptational response to low light intensity of intact Fungia spp. was found to be a combination of the photoadaptation of symbiotic algae and the decrease of host animal tissue. In order to study their adaptation to ultraviolet (UV) radiation, P-I curves of Fungia spp. and isolated zooxanthellae were analyzed before and after UV-B irradiation. 1 h UV-B irradiation showed no effect on the photosynthetic rate of the shallow water (1 m) corals, while it inhibited the photosynthesis of the deep water (30 m) corals and zooxanthellae isolated from both shallow and deep water corals. These results indicate that the host, Fungia spp., in shallow water have protective mechanism for intense UV-B in their habitat. These photoadaptational mechanisms seem to allow the Fungia spp. to have wide vertical distribution where light intensity spans more than two orders of magnitude.     

Effect of solar ultra-violet radiation on the kelp Ecklonia radiata

During spring and summer, 1982–1986, experiments were carried out near Marmion Reef, Western Australia. In summer, nearly 30% of the surface solar ultraviolet radiation (280 to 400 nm) penetrates offshore waters to 5 m depth. Experimental removal of the mature Ecklonia radiata kelp canopy in summer results in tissue damage, photopigment destruction, reduced growth, and low survivorship of subcanopy kelp sporophytes. These effects do not occur with canopy removal in winter. Laboratory experiments revealed that the UV component of radiation, rather than intense photosynthetically active radiation, was responsible for the inhibition of growth and photodamage. UV radiation probably affects survival of the settlement stages of E. radiata sporophytes, thus excluding them from otherwise suitable substrata in shallow waters. UV radiation is implicated in the reduction of canopy productivity in summer.     

Depth-dependent photoadaption by zooxanthellae of the reef coral Montastrea annularis

Zooxanthellae living in colonies of the Caribbean reef coral Montastrea annularis photoadapt to depth-dependent attenuation of submarine light. Studies carried out at Discovery Bay, Jamaica, show that in shallow-living coral colonies, the zooxanthellae appear photoadapted to function at high light intensities, and do poorly if transplanted to low light intensities; in contrast, zooxanthellae in deeper-living coral colonies can be damaged by high light intensities. The adaptation to decreasing light intensity and changing spectral quality appears to be accomplished by increasing the size of the photosynthetic unit (PSU), as opposed to increasing the number of PSU's per cell. Whole cell absorption increases with depth, partially offsetting the loss of light energy due to depth-dependent attenuation. Calculations of photosynthetically usable radiation, the light an alga is capable of absorbing in its own submarine habitat, suggest that the algae at different depths are optimizing rather than maximizing their ability to harvest submarine light energy.     

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.