Summary
1.We evaluated the potential for three species of deciduous-forest herbs to exploit seasonal periods of direct irradiance. In particular, we investigated the importance of photosynthetic acclimation as a mechanism for shade-tolerant herbs to utilize direct light reaching the forest floor before canopy expansion in the spring and after canopy leaf drop in the autumn. 2. We measured the photosynthetic and growth characteristics of three co-occurring herbs of a northern hardwood forest: the spring ephemeral Allium tricoccum Ait., the summer-green Viola pubescens Ait., and the semi-evergreen Tiarella cordifolia L. 3. Leaf CO 2 exchange, leaf mass per area, and leaf biochemistry differed among species and seasonally within species to match the changing light environment below the forest canopy. From spring to summer, as irradiance dropped with the expansion of the overstorey canopy, Viola leaves exhibited reduction of both photosynthetic capacity and light compensation point. Weaker acclimation of less magnitude occurred in Tiarella leaves over the spring-summer light transition; this was followed by further acclimation to the stronger autumn irradiance. 4. Viola 's greater range of photosynthetic acclimation was associated with shifts in allocation between Rubisco and chlorophyll, as well as changes in total leaf nitrogen (N) concentration and leaf mass per area (LMA). In contrast, Tiarella 's narrow range of acclimation was associated solely with changes in allocation to Rubisco versus chlorophyll, with no changes in total leaf N or LMA. 5. Seasonal changes in leaf chemistry and structure in Viola suggest a stepwise ontogeny whereby individual leaves are able to function as 'sun leaves' for 3-5 weeks in the spring, and then as 'shade leaves' for up to 3 months in the summer. 6. Whole-plant biomass accumulation showed that all three species accumulated most of their annual biomass increment during periods of direct irradiance. These results demonstrate the importance of brief seasonal periods of strong irradiance to the growth of deciduous forest herbs, even shade-tolerant, summer and evergreen species.
We used a 72-year chronosequence to study the loss and recovery of ecosystem C pools following stand-replacing wildfire in Michigan, USA, jack pine (Pinus banksiana Lamb.) forests. We quantified the amount of C stored in aboveground plant biomass, standing dead timber, downed dead wood, surface organic soil, and mineral soil in 11 jack pine stands that had burned between 1 and 72 years previously. Total ecosystem C ranged from a low of 59 Mg C·ha1 in the 4-year-old stand to 110 Mg C·ha1 in the 72-year-old stand. Changes in total ecosystem C across the chronosequence conformed to theoretical predictions, in which C stocks declined initially as decomposition of dead wood and forest-floor C exceeded production by regenerating vegetation, and then increased asymptotically with the development of a new stand of jack pine. This pattern was well described by the following "gamma" function: total ecosystem C (Mg·ha1) = 112.2 39.6 × age0.351 × exp(0.053 × age01.039); mean-corrected R2 = 0.976. Using the first derivative of this parameterized gamma function, we estimated that jack pine stands function as a weak source of C to the atmosphere for only ca. 6 years following wildfire, and reach a maximum net ecosystem productivity of 1.6 Mg C·ha1·year1 by year 16. We attribute the rapid transition from carbon source to carbon sink in these ecosystems to two factors: (i) stand-replacing wildfires in these xeric forests leave behind little respirable substrate in surface organic horizons, and (ii) jack pine is able to rapidly reestablish following wildfires via serotinous cones. Jack pine stands remained net sinks for C across the chronosequence; however, net ecosystem productivity had declined to 0.12 C ha1·year1 by year 72. Carbon sequestration by mature jack pine ecosystems was driven primarily by continued growth of overstory jack pine, not by accumulation of detrital C.
We investigated the changes in soil processes following wildfire in Michigan jack pine (Pinus banksiana) forests using a chronosequence of 11 wildfire-regenerated stands spanning 72 years. The objective of this study was to characterize patterns of soil nutrients, soil respiration and N mineralization with stand development, as well as to determine the mechanisms driving those patterns. We measured in situ N mineralization and soil respiration monthly during the 2002 growing season and used multiple regression analysis to determine the important factors controlling these processes. Growing-season soil respiration rates ranged from a low of 156 g C/m2 in the 7-year-old stand to a high of 254 g C/m2 in the 22-year-old stand, but exhibited no clear pattern with stand age. In general, soil respiration rates peaked during the months of July and August when soil temperatures were highest. We used a modified gamma function to model a temporal trend in total N mineralization (total N mineralization=1.853-0.276xagexe-0.814xage; R2=0.381; P=0.002). Total N mineralization decreased from 2.8 g N/m2 in the 1-year-old stand to a minimum value of 0.5 g N/m2 in the 14-year-old stand, and then increased to about 1.5 g N/m2 in mature stands. Changes in total N mineralization were driven by a transient spike in N turnover in the mineral soil immediately after wildfire, followed by a gradual accrual of a slow-cycling pool of N in surface organic horizons as stands matured. Thus, in Michigan jack pine forests, the accumulation of surface organic matter appears to regulate N availability following stand-replacing wildfire.
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