Solar energy capture and transformation in the sea

Solar energy ultimately drives all biogeochemical cycles and sustains planetary habitability. All life forms and processes on Earth, including human economic and social systems, exist within a complex network of energy flow. In the sea, microorganisms comprise most of the genetic and metabolic diversity, and are responsible for a majority of the system energy flow including solar energy capture, transformation, and dissipation. All of these processes involve conversion of low quality forms of energy into a smaller fraction of higher quality energy plus degraded heat, in accordance with the basic laws of thermodynamics. Energy flow is at the core of ecosystem analysis (Odum 1968). Sunlight is the most abundant form of energy for marine microorganisms, and biophysical/biochemical mechanisms for solar energy capture have evolved by natural selection during eons of Earth’s history (Brown and Ulgiati 2004; Nealson and Rye 2003). Marine ecosystems, especially the expansive subtropical gyres, have an enormous capacity for solar energy capture and transformation. Ecologists often use the term “carbon and energy flow” to describe solar energy capture, organic matter transformation, and heat dissipation through the food web via the coupled processes of photosynthesis and respiration. A number of different methods have been used to track the flow of carbon and associated bioelements (e.g., nitrogen, phosphorus, oxygen, and sulfur), but energy flow is rarely if ever measured in field studies. An untested assumption is that matter and energy flow are inextricably and quantitatively linked in space and time in the open sea. Howard T. Odum, largely in collaboration with his brother Eugene P. Odum, pioneered the discipline of systems ecology. He observed and studied a variety of aquatic ecosystems and was the first to characterize them as networks of energy circuits (e.g., Silver Springs, Florida; Odum 1956). Odum later developed an explicit energy circuit language and set of symbols that could be used to represent interactive energy capture, transformation, and dissipation in both natural and manmade systems (Odum 1983a). While some scientists have criticized “Odum’s conjectures” and his energy-centric approach to the study of ecosystems (e.g., Mansson and McGlade 1993), the debate centers on the formidable obstacles to a comprehensive, quantitative analysis and understanding of ecological energy flow rather than a challenge to its fundamental importance in ecosystem analysis. In a pioneering essay on the relationship of energy flow to evolution, Alfred Lotka concluded that natural selection will operate to preserve and expand those species “possessing superior energy-capturing and directing devices” (Lotka 1922). Consequently, he reasoned, as long as there is a residue of untapped available energy, “the total organic mass of the system, the rate of circulation of mass through the system, and the total energy flux” will be maximized. This reasoning has since become known as the maximum power principle (Odum and Pinkerton 1955; Odum 1983b), and has led to vigorous debate over the validity and implications of what some have termed the fourth law of thermodynamics (see Sciubba 2011 for a recent assessment). The Earth is an energetically open system where solar energy input is balanced by radiative heat loss. There are numerous connections among the hydrosphere, lithosphere, and atmosphere such that materials and energy can be easily exchanged. In a thought-provoking commentary, On certain unifying principles in ecology, Ramon Margalef concluded that the energy required to maintain an ecosystem is inversely proportional to Domain Editor-in-Chief Jody W. Deming, University of Washington