Nuclear magnetic resonance spectroscopy of living systems: applications in comparative physiology

The most attractive feature of nuclear magnetic resonance spectroscopy (MRS) is the noninvasive and nondestructive measurement of chemical compounds in intact tissues. MRS already has many applications in comparative physiology, usually based on observation of 31P, since levels of phosphorus compounds indicate tissue energy status and are changed during exercise, fatigue, recovery, hypometabolism, anesthesia, hypoxia, hypercapnia, and osmotic and acid stress. Nuclei other than 31P may also be monitored, such as 1H, 13C, 15N, 19F, or 23Na, and applied in biological research. Particularly, 13C-MRS is interesting because it allows the analysis of metabolic pathways in living systems. Applications of MRS in comparative physiology and biochemistry are comprehensively discussed in this review. The main focus is on anaerobic metabolism during hypoxia, ischemia, and exercise. Species as widely different as slime molds, nematodes, frogs, turtles, and ducks have been studied by 31P-MRS. It is not surprising that striking species differences do occur, but many similarities are also observed. Unique is the occurrence of six different phosphagens with different values of Gibbs free energy in polychete worms The presence of a particular phosphagen may be related to the average oxygen tension within the tissues. Phosphagens and their kinases are also discussed in relation to hypercapnia and acid stress. Other topics discussed in this paper are enzyme kinetics, anesthetics, development and growth, parasitism, and the detection of previously unknown compounds.