Living Organisms Exist in a Dynamic Steady State, Never at Equilibrium with Their Surroundings

The molecules and ions contained within a living organism differ in kind and in concentration from those in the organism’s surroundings. A Paramecium in a pond,  a shark in the ocean, an erythrocyte in the human bloodstream, an apple tree in an orchard—all are different in composition from their surroundings and, once they have reached maturity, all (to a first approximation) maintain a constant composition in the face of constantly changing surroundings. Although the characteristic composition of an organism changes little through time, the population of molecules within the organism is far from static. Small molecules, macromolecules, and supramolecular complexes are continuously synthesized and then broken down in chemical reactions that involve a constant flux of mass and energy through the system. The hemoglobin molecules carrying oxygen from your lungs to your brain at this moment were synthesized within the past month; by next month they will have been degraded and entirely replaced by new hemoglobin molecules. The glucose you ingested with your most recent meal is now circulating in your bloodstream; before the day is over these particular glucose molecules will have been converted into something else—carbon dioxide or fat, perhaps—and will have been replaced with a fresh supply of glucose, so that your blood glucose concentration is more or less constant over the whole day. The amounts of hemoglobin and glucose in the blood remain nearly constant because the rate of synthesis or intake of each just balances the rate of its breakdown, consumption, or conversion into some other product. The constancy of concentration is the result of a dynamic steady state, a steady state that is far from equilibrium. Maintaining this steady state requires the constant investment of energy; when the cell can no longer generate energy, it dies and begins to decay toward equilibrium with its surroundings. We consider below exactly what is meant by  “steady state” and “equilibrium.”