Understanding freeze stress in biological tissues: Thermodynamics of interfacial water

A thermodynamic approach to distinguish forms of freeze energy that injure plants as the temperature decreases is developed. The pattern resulting from this analysis dictated the sequence of thermal requirements for water to exist as an independent state. Improvement of freezing tolerance in biological systems depends on identification of a specific form of stress, just as control of a disease depends on identification of the pathogen causing the disease. The forms of energy that stress hydrated systems as temperature decreases begin with disruption of biological function from chill injury that occurs above freezing. Initiation of non-equilibrium freezing with sufficient free energy to drive disruptive effects can occur in a supercooled system. As the temperature continues to decrease and freezing occurs in an equilibrium manner, adhesion at hydrated interfaces contributes to disruptive effects as protoplasts contract by freeze-dehydration. If protective systems are able to prevent injury from direct interactions with ice, passive effects of freeze-dehydration may cause injury at lower temperatures. The temperature range in which an injury occurs is an indicator of the form of energy causing stress. The form of energy is thus a primary guide for selection of a protective mechanism. An interatomic force model whose response to temperature change corresponds with the enthalpy pattern might help define freeze stress from a unique perspective.