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beech Botkin camera canopy dynamics canopy profile CNDO correlation crown class crown volume crown width developed diameter discussed distribution Ecol ecosystem equation factors figure forest canopy gradient analysis hobblebush I I I I I I I I QI increase indicates IOOOOIO I I IQ Q Jabowa model leaf biomass leaf weight leaves litter fall measured meter Mitchell and Chandler moisture nitrogen nitrogen availability northern hardwood nutrient nutrient availability nutrient content o.o o.o OOO OOO OOO paper birch permanent wilting point photosynthesis pin cherry PLOT STAND predict Q Q Q Q Q Q QQ Q ratios red oak regressions relationships resource gradients response sampling seedlings shade tolerance simulation soil species striped maple successional sugar maple Table technique tissue trees understory values variables white ash Whittaker yellow birch