C4 grassland in South Africa with a 1.7 m Carl Morrow for scale.
Species with the C4 photosynthetic pathway are in the minority in terms of species, but fix a large amount of the world's carbon, not to mention world's calories that humans consume.
Species with the C4 photosynthetic pathway differ from C3 species in a number of ways. We know that the C4 photosynthetic pathway evolved, or at least radiated during times of declining atmospheric CO2 concentrations. In accordance, C4 species have higher photosynthetic rates at glacial CO2 concentrations (~200 ppm) than C3 species. Therefore, it is generally thought to be an evolutionary response to low CO2 concentrations. In conditions of high light, low CO2, and warm temperatures, the C4 pathway reduces photorespiration and generates greater photosynthetic rates over C3 species.
Yet, the C4 photosynthetic pathway also confers greater resource use efficiency. The C4 pathway comes with increased energetic costs, but also confers greater photosynthetic water use and nitrogen use efficiency. More carbon is fixed in C4 species per unit water and nitrogen allocated to photosynthesis as internal CO2 concentrations are lower, which drives the greater WUE, and less N is needed for the same amount of photosynthesis, which drives greater NUE.
Some of the characteristics of C4 are a bit mythological. For example, although C4’s can have higher photosynthetic nitrogen use efficiency, many C4’s have high tissue N concentrations and many C3’s have as low an N concentration as the lowest C4. Not everything about plants is destined from photosynthetic properties.
That said, is there selective advantage to being more efficient with resources? Efficiency always comes at a cost. This much we know. You have to be inefficient with one resource to be more efficient with another. Light use efficiency comes at the expense of N use efficiency. N use efficiency comes at the expense of water use efficiency. Efficiency also costs time.
So what is the benefit of being efficient? For C4’s, under what conditions is it beneficial to be more efficient with water or nitrogen than C3’s. In a competitive world, efficiency in and of itself benefits no one but your competitors. The less water or nitrogen you use, the more there is for another. The benefit only comes if efficiency allows one to reduce the availability of the limiting resource below the level needed to sustain a potential competitor. Or tolerate more stressful conditions. Do C4’s reduce water or nitrogen availability to lower levels than C3’s? No evidence of that. Do C4’s tolerate lower water or nitrogen availability than C4’s? No evidence of that, either.
We also know that C4’s span a wide range of water and nitrogen availability. NADP-me type C4’s increase with mean annual precipitation, not decrease. And C4’s like the grasses we use in many lawns and golf courses have high nutrient requirements, not low, having evolved in grazing lawns that have high nutrient availability. In all, there is no evidence that C4’s preferentially occupy low water or low nitrogen habitats.
The efficiency of C4 species is one of the great mysteries of evolution. Is it an interesting by-product of selection for carbon gain under certain conditions? Or is it indirectly linked to success in ways that are not obvious? Likely, until we better understand the fundamental question of “Who wins and why?” in the plant world, that aspect of C4’s will still be a mystery.