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P is an extensive variable and is the population's “Wallace pressure”—its total energy flux and output in joules per second.
M is an extensive variable, and is the total mass of chemical components the population holds at any time; R is an intensive variable and is the population's average energy content expressed in “darwins” or joules per “biomole” (where the biomole is a unit of population size).
(Corollary: prodigious savant are always possible; and even “the walls of rude minds are scrawled all over with facts, with thoughts” Ralph Waldo Emerson).
(Corollary: “Bernard of Chartres used to say that we are like dwarfs on the shoulders of giants, so that we can see more than them, and things at a greater distance, not by virtue of any sharpness of sight on our part, or any physical distinction, but because we are carried high and raised up by their giant size” John of Salisbury, Metalogicon, 1159).
S is an intensive variable and is the population's (specific) work rate or its dynamical specific energy, but expressed as watts delivered per kilogrammes of mass maintained per “biomole”.