Plant roots acidify soil by releasing protons and organic acids so as to chemically weather soil minerals. Decaying remains of dead plants on soil may also form organic acids which contribute to soil acidification. Acidification from leaf litter on the O-horizon is more pronounced under coniferous trees such as pine, spruce and fir, which return fewer base cations to the soil, rather than under deciduous trees; however, soil pH differences attributed to vegetation often preexisted that vegetation, and help select for species which tolerate them. Calcium accumulation in existing biomass also strongly affects soil pH - a factor which can vary from species to species.