Coral Reef - Darwin's Paradox

Darwin's paradox Coral... seems to proliferate when ocean waters are warm, poor, clear and agitated, a fact which Darwin had already noted when he passed through Tahiti in 1842.

This constitutes a fundamental paradox, shown quantitatively by the apparent impossibility of balancing input and output of the nutritive elements which control the coral polyp metabolism.

Recent oceanographic research has brought to light the reality of this paradox by confirming that the oligotrophy of the ocean euphotic zone persists right up to the swell-battered reef crest. When you approach the reef edges and atolls from the quasidesert of the open sea, the near absence of living matter suddenly becomes a plethora of life, without transition. So why is there something rather than nothing, and more precisely, where do the necessary nutrients for the functioning of this extraordinary coral reef machine come from ? — Francis Rougerie

During his voyage on the Beagle, Darwin described tropical coral reefs as oases in the desert of the ocean. He reflected on the paradox that tropical coral reefs, which are among the richest and most diverse ecosystems on earth, flourish surrounded by tropical ocean waters that provide hardly any nutrients.

Coral reefs cover less than 0.1% of the surface of the world’s ocean, yet they support over one-quarter of all marine species. This diversity results in complex food webs, with large predator fish eating smaller forage fish that eat yet smaller zooplankton and so on. However, all food webs eventually depend on plants, which are the primary producers. Coral reefs' primary productivity is very high, typically producing 5–10 g·cm−2·day−1 biomass.

One reason for the unusual clarity of tropical waters is they are deficient in nutrients and drifting plankton. Further, the sun shines year round in the tropics, warming the surface layer, making it less dense than subsurface layers. The warmer water is separated from deeper, cooler water by a stable thermocline, where the temperature makes a rapid change. This keeps the warm surface waters floating above the cooler deeper waters. In most parts of the ocean, there is little exchange between these layers. Organisms that die in aquatic environments generally sink to the bottom, where they decompose, which releases nutrients in the form of nitrogen (N), phosphorus (P) and potassium (K). These nutrients are necessary for plant growth, but in the tropics, they do not directly return to the surface.

Plants form the base of the food chain, and need sunlight and nutrients to grow. In the ocean, these plants are mainly microscopic phytoplankton which drift in the water column. They need sunlight for photosynthesis, which powers carbon fixation, so they are found only relatively near the surface. But they also need nutrients. Phytoplankton rapidly use nutrients in the surface waters, and in the tropics, these nutrients are not usually replaced because of the thermocline.

Around coral reefs, lagoons fill in with material eroded from the reef and the island. They become havens for marine life, providing protection from waves and storms.

Most importantly, reefs recycle nutrients, which happens much less in the open ocean. In coral reefs and lagoons, producers include phytoplankton, as well as seaweed and coralline algae, especially small types called turf algae, which pass nutrients to corals. The phytoplankton are eaten by fish and crustaceans, who also pass nutrients along the food web. Recycling ensures fewer nutrients are needed overall to support the community.

Coral reefs support many symbiotic relationships. In particular, zooxanthellae provide energy to coral in the form of glucose, glycerol, and amino acids. Zooxanthellae can provide up to 90% of a coral’s energy requirements. In return, as an example of mutualism, the corals shelter the zooxanthellae, averaging one million for every cubic centimeter of coral, and provide a constant supply of the carbon dioxide they need for photosynthesis.

Corals also absorb nutrients, including inorganic nitrogen and phosphorus, directly from water. Many corals extend their tentacles at night to catch zooplankton that brush them when the water is agitated. Zooplankton provide the polyp with nitrogen, and the polyp shares some of the nitrogen with the zooxanthellae, which also require this element. The varying pigments in different species of zooxanthellae give them an overall brown or golden-brown appearance, and give brown corals their colors. Other pigments such as reds, blues, greens, etc. come from colored proteins made by the coral animals. Coral which loses a large fraction of its zooxanthellae becomes white (or sometimes pastel shades in corals that are richly pigmented with their own colorful proteins) and is said to be bleached, a condition which, unless corrected, can kill the coral.

Sponges are another key to explaining Darwin’s paradox. They live in crevices in the coral reefs. They are efficient filter feeders, and in the Red Sea they consume about 60% of the phytoplankton that drifts by. The sponges eventually excrete nutrients in a form the corals can use.

The roughness of coral surfaces is the key to coral survival in agitated waters. Normally, a boundary layer of still water surrounds a submerged object, which acts as a barrier. Waves breaking on the extremely rough edges of corals disrupt the boundary layer, allowing the corals access to passing nutrients. Turbulent water thereby promotes reef growth and branching. Without the nutritional gains brought by rough coral surfaces, even the most effective recycling would leave corals wanting in nutrients.

Cyanobacteria provide soluble nitrates for the reef via nitrogen fixation.

Coral reefs also often depend on surrounding habitats, such as seagrass meadows and mangrove forests, for nutrients. Seagrass and mangroves supply dead plants and animals which are rich in nitrogen and also serve to feed fish and animals from the reef by supplying wood and vegetation. Reefs, in turn, protect mangroves and seagrass from waves and produce sediment in which the mangroves and seagrass can root.