Biomass and nitrogen allocation to leaf, root, stem, and reproduction was determined in a 35—field chronosequence that spans the first 60 yr of secondary succession on a Minnesota sand plain. Biomass (grams per square metre) in leaf and root increased during succession, but reproductive biomass declined, and that in stem remained constant. Because root biomass increased twice as rapidly as leaf biomass, the proportion of total biomass in root increased during succession, whereas that in leaf, reproduction, and stem declined. In an additional study, biomass allocation was determined on a species—by—species basis for 46 species common at different times during succession. This study showed a similar pattern of increasing proportional root allocation and declining proportional reproductive and stem allocation during succession. These changes were accompanied by an increase in total soil nitrogen and a decrease in light penetration to the soil surface during succession. Increasing root allocation and decreasing reproductive allocation suggest that succession on these nutrient—poor soils is the transient dynamics of colonization and competitive displacement, with later successional species being superior nitrogen competitors because of higher root allocation. Allocation trade—offs root, stem, leaf, and seed can lead to initial dominance by species with high seed and leaf allocation, presumably because of greater colonization and/or maximal growth rates. Thus, this succession differs markedly from successions on rich soils, for which stem allocation is increasingly important. These results contradict the resource ratio hypothesis as an explanation for the pattern of early succession on impoverished soils.
Soil C and N dynamics were studied in a sequence of old fields of increasing age to determine how these biogeochemical cycles change during secondary succession. In addition, three different late-successional forests were studied to represent possible "steady state" conditions. Surface soil samples collected from the fields and forests were analyzed for total C, H,O-soluble C, total N, potential net N mineralization, potential net nitrification, and microbial biomass. Aboveand belowground plant biomass was estimated within each of the old field sites.Temporal changes in soil organic C, total N and total plant biomass were best described by a gamma function [y = at* ecrd + f] whereas a simple exponential model [y = a(1 -em*') + c] provided the best fit to changes in H,O-soluble C, C:N ratio, microbial C, and microbial N. Potential N mineralization and nitrification linearly increased with field age; however, rates were variable among the fields. Microbial biomass was highly correlated to soil C and N pools and well correlated to the standing crop of plant biomass. In turn, plant biomass was highly correlated to pools and rates of N cycling. Patterns of C and N cycling within the old field sites were different from those in a northern hardwood forest and a xeric oak forest; however, nutrient dynamics within an oak savanna were similar to those found in a 60-yr old field. Results suggest that patterns in C and N cycling within the old-field chronosequence were predictable and highly correlated to the accrual of plant and microbial biomass.
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