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The Effect of Excess Algae on Coral Mortality
By Laura Miotke
The death and bleaching of corals have been examined throughout recent years with a number of different culprits in mind. Global warming has been charged with dissolving higher amounts of CO2 into the ocean, increasing its acidity and causing coral bleaching and mortality. On a more local level, scientist for decades now, have observed that corals die in regions of high nutrient disposal and overfishing. But what about these actually kill the coral? In these local areas of high coral death, observers have witnessed a shift from a coral and algae balanced environment to a predominantly fleshy algal lawn. Fleshy algae are considered to be the combination of seaweeds (or macroalgae) and turf algae. These are usually limited in growth by the grazing fish population and the low nutrient environments surrounding the coral reefs. Thus, it seems intuitive that the removal of herbivores or the excess of nutrients (such as phosphorous and nitrogen) would cause the overgrowth of the fleshy algae. In a normally balanced reef, the role of the algae is that of primary producer. Through photosynthesis the plants fix the dissolved carbon dioxide, playing a role in the aquatic pH, as well as the calcium carbonate availability. As said before the growth of the algae is limited by a lack of nitrogen and phosphorus, but since CO2 and light are in excess they hardly ever lack usable carbon. Thus, in low nutrient environments they are producing more photosynthate than they can use to grow, and end up releasing most of it in the form of dissolved organic carbon (DOC). This DOC, which is constantly produced in healthy reef environments, is used by the billions of microorganisms as an energy and carbon source. This abundance and diversity of microorganisms is residing in the water but also on the coral themselves, on the algae and pretty much everything else in the ecosystem. The coral produces a mucosal layer to inhibit bacteria from entering through its cellular layer. This layer also provides a habitat for an entire community of bacteria to live. This community has a complex symbiosis that benefits drastically from any excess DOC in the area. Thus if algae increases (due to overfishing or nutrient dumping), more DOC is produced and the bacteria living on the corals grow rapidly.
Table 1. Amino acids, chlorophyll a, and tissue nitrogen in Gracilaria edulis after 3 days field incubation in One Tree Island ENCORE experimental patch reefs, phase 2 (high nutrient loading period).
Figure 1. Montastraea annularis. Percentage of coral mortality and bleaching (mean ± SD, given above) from nitrogen (N, includes ammonium chloride and calcium nitrate), potassium phosphate (P) and organic carbon (C) additions after 30 d. Asterisks (**) indicate significant differences between treatments and controls (p < 0.05, Mann-Whitney U-test). Carbon treatments included a naturally occurring carbon source, particulate organic matter (POM).
In 2001, the ENCORE project tested the effect of nutrient addition on various aspects of a small portion of the Great Barrier Reef. In one section they tested these conditions on the filamentous macroalgal species, Gracilaria edulis (Rhodophyta). G. edulis, collected in Moreton Bay, Queensland, was incubated in the conditioned waters in a perforated plastic container (that allowed water exchange) for 3 days. The pigment, tissue nutrient and amino acid content were measured. As seen in Table 1, the amount of Citrulline, a 3N amino acid used for nitrogen storage, was greatly increased in the +N and +N+P conditions. The total amino acids, tissue nitrogen and pigment chlorophyll also increased from the control, especially in the +N +P condition. This indicated that the G. edulis algae was adapting to the excess nutrients and increasing their uptake. (Koop Encore) In the next experiment, Kuntz and Kline (et. al) showed that while the added nutrients were increasing the growth of the algae they were not directly killing the coral. They created a specialized system called the Aquatic Automated Dosing and Maintenance System (AADAMS) which administered seawater with various treatments to corresponding chambers containing coral nubbins. The coral nubbins used were Montastraea annularis from the Bocas del Toro, Republic of Panama. The nubbins were treated with, nitrate, ammonia, 5 DOC sources (D-glucose, lactose, galactose, starch, arabinose) and untreated seawater controls. The nubbins were monitored every day for 30 days and mortality was measure as the relative percentage of polyps that died (defined as the complete loss of tissue revealing bare skeleton). Bleaching was also observed. As seen in Figure 1, only the conditions treated with DOC (carbon sources) showed a significantly higher mortality than the controls. This indicates that the added nutrients do not directly kill the coral nubbins, while DOC does. In the same experiment, they also measured microbial growth in response to the same additions (nitrogen, phosphorous and DOC). Four nubbins (of the same species) were placed in an aquaria and treated with one treatment. At 1, 2, 4, and 26 hours samples of the coral surface mucopolysaccharide layer (SML) were taken. The microbial production was measured with [3H-methyl]-thymidine incorporation. Only the carbon+ treatment elicited an increased Based on the results of these three studies, along with many other follow-ups growth in the microbial mucosal layer (Figure 2). This would indicate that the DOC does not kill the coral directly but increased the growth of the microbes and slightly variant additions, give a very plausible explanation for the localized local to the coral. causes of coral death. The progression happens as follows: The next piece fell into place with the experiments of Smith et. al, who tested the effects of algae on coral. With the hypothesis in mind that DOC, excreted Nutrients are dumped into the ocean, which increases P and N, which cause the from algae, was the cause of coral death and microbial growth, Smith created a two-chamber set-up that allowed water exchange but separated the coral fleshy algae to grow (or the herbivores who usually keep the algae down are and the algae (see Figure 4). A 0.02 lm Anodisc filter was used which prevented the movement of viruses and bacteria but overfished). The algae then produced an increased level of DOC, which not dissolved compounds. When the coral species Pocillopora verrucosa was placed across the filter from the green algae Dictyosphaeria cavernosa for 36 hours there showed a marked decrease in photosynthetic efficiency (thus indicating unhealthy coral tissue) than in the control (with no algae across the filter) increases the growth of microbes living in the coral mucosal layer. The (Figure 3). This indicates that the algae caused the coral to die without direct contact, but through a permeable membrane. Furthermore, when treated with microbes use all the O2 and the coral suffocates. an antibiotic (Ampicillan) all the coral species lived, even those in a chamber with algae. This supports the conclusions of Kuntz and Kline that it is the Resources: microbial species, stimulated by DOC, were killing the coral. Furthermore, it is the microbial species native to the corals that is causing the death, not the Kline, D.I., et al. (2006) Role of elevated organic carbon levels and microbial activity in coral mortality. Marine microbes on the algae. Ecology Progress Series 314: 119-125 Smith took this conclusion even further in the second part of the experiment. A Clark-type oxygen microprobe was moved up and down relative to the coral surface to find the range of oxygen maxima. Then a reading of the dissolved oxygen was taken and averaged over 0.5 seconds. Aerated seawater was used Koop, K., et al. (2001) ENCORE: The effect of nutrient enrichment on coral reefs. Synthesis of results and for the 100% dissolved oxygen calibration point. Seawater amended with 0.1 M sodium ascorbate and 0.1 M sodium hydroxide was used as the 0% conclusions. Marine Pollution Bulletin 42: 91-120 dissolved oxygen calibration point. On the control coral the oxygen reading was higher than that of the normal aerated seawater. This is due to the oxygen Smith, J.E., et al. (2006) Indirect effects of algae on coral: Algae-mediated, microbe-induced coral mortality. production of the photosynthesizing zooxanthellae that live symbiotically in the coral. In the coral samples across from the algae the microprobe readings showed drastically hypoxic conditions along the surface of the coral (Figure 4). This result indicates that the increase in microbes on the surface of the coral Ecology Letters 9: 834-845 causes its death by suffocation.
Figure 2. Montastraea annularis. Microbial production as measured by thymidine incorporation in coral surface mucopolysaccharide layer (SML) when exposed to nitrogen, phosphorus and carbon treatments after 1, 2, 4 and 26 h of exposure (±SD). Asterisks (**) indicate that the microbial production rate was significantly greater than the controls (p < 0.001, MannWhitney U-test)
Figure 3. Results of quantum yield of photochemistry for PS II (Fv/Fm) as measured with pulse amplitude modulation fluorometry (PAM). Measurements are of dark-adapted yield (DAY) following saturating pulses for both coral and algae. (a) DAY for Dictyosphaeria cavernosa and Pocillopora verrucosa following 36 h in experimental chambers (n=5); (b) DAY for coral and algae (Control=coral or algae grown alone, with algae=coral and algae grown together in chambers, +AMP=with ampicillin, -AMP =seawater alone) following 36 h in experimental chambers (n=5 for experimental and 3 for controls); (c) DAY for both coral and algae pooled for all species combinations assessed after 48 h in experimental containers (n=20). Figure 4. Photographs showing a coral algal pair in experimental chambers and coral controls from experiment 1. The graphs show oxygen microprobe data taken along transects across the respective organismal surfaces. The x-axis are in units of time and do not correspond to distance along the organism¶s surface. 100% oxygen saturation represents the background water (red), any addition above this indicates input via photosynthesis and any decrease indicates oxygen consumption.
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