The seventh macronutrient: how sodium shortfall ramifies through populations, food webs and ecosystems.

Of the 25 elements required to build most organisms, sodium has a unique set of characteristics that ramify through terrestrial ecology. In plants, sodium is found in low concentrations and has little metabolic function; in plant consumers, particularly animals, sodium is essential to running costly Na-K ATPases. Here I synthesise a diverse literature from physiology, agronomy and ecology, towards identifying sodium's place as the '7th macronutrient', one whose shortfall targets two trophic levels - herbivores and detritivores. I propose that sodium also plays a central, though unheralded role in herbivore digestion, via its importance to maintaining microbiomes and denaturing tannins. I highlight how sodium availability is a key determinant of consumer abundance and the geography of herbivory and detritivory. And I propose a re-appraisal of the assumption that, because sodium is metabolically unimportant to most plants, it is of little use. Instead, I suggest that sodium's critical role in limiting herbivore performance makes it a commodity used by plants to manipulate their herbivores and mutualists, and by consumers like bison and elephants to generate grazing lawns: dependable sources of sodium.