Tree species effects on ecosystem water-use efficiency in a high-elevation, subalpine forest.

Ecosystem water-use efficiency (eWUE; the ratio of net ecosystem productivity to evapotranspiration rate) is a complex landscape-scale parameter controlled by both physical and biological processes occurring in soil and plants. Leaf WUE (lWUE; the ratio of leaf CO(2) assimilation rate to transpiration rate) is controlled at short time scales principally by leaf stomatal dynamics and this control varies among plant species. Little is known about how leaf-scale variation in lWUE influences landscape-scale variation in eWUE. We analyzed approximately seven thousand 30-min averaged eddy covariance observations distributed across 9 years in order to assess eWUE in two neighboring forest communities. Mean eWUE was 19% lower for the community in which Engelmann spruce and subalpine fir were dominant, compared to the community in which lodgepole pine was dominant. Of that 19% difference, 8% was attributed to residual bias in the analysis that favored periods with slightly drier winds for the spruce-fir community. In an effort to explain the remaining 11% difference, we assessed patterns in lWUE using C isotope ratios. When we focused on bulk tissue from older needles we detected significant differences in lWUE among tree species and between upper and lower canopy needles. However, when these differences were scaled to reflect vertical and horizontal leaf area distributions within the two communities, they provided no power to explain differences in eWUE that we observed in the eddy covariance data. When we focused only on bulk needle tissue of current-year needles for 3 of the 9 years, we also observed differences in lWUE among species and in needles from upper and lower parts of the canopy. When these differences in lWUE were scaled to reflect leaf area distributions within the two communities, we were able to explain 6.3% of the differences in eWUE in 1 year (2006), but there was no power to explain differences in the other 2 years (2003 and 2007). When we examined sugars extracted from needles at 3 different times during the growing season of 2007, we could explain 3.8-6.0% of the differences in eWUE between the two communities, but the difference in eWUE obtained from the eddy covariance record, and averaged over the growing season for this single year, was 32%. Thus, overall, after accounting for species effects on lWUE, we could explain little of the difference in eWUE between the two forest communities observed in the eddy covariance record. It is likely that water and C fluxes from soil, understory plants, and non-needle tissues, account for most of the differences observed in the eddy covariance data. For those cases where we could explain some of the difference in eWUE on the basis of species effects, we partitioned the scaled patterns in lWUE into two components: a component that is independent of canopy leaf area distribution, and therefore only dependent on species-specific differences in needle physiology; and a component that is independent of species differences in needle physiology, and only dependent on species-specific influences on canopy leaf area distribution. Only the component that is dependent on species influences on canopy leaf area distribution, and independent of inherent species differences in needle physiology, had potential to explain differences in eWUE between the two communities. Thus, when tree species effects are important, canopy structure, rather than species-specific needle physiology, has more potential to explain patterns in eWUE.

The effects of tree rhizodeposition on soil exoenzyme activity, dissolved organic carbon, and nutrient availability in a subalpine forest ecosystem.

Previous studies have found that root carbon inputs to the soil can stimulate the mineralization of existing soil carbon (C) pools. It is still uncertain, however, whether this "primed" C is derived from elevated rates of soil organic matter (SOM) decomposition, greater C release from microbial pools, or both. The goal of this research was to determine how the activities of the microbial exoenzymes that control SOM decomposition are affected by root C inputs. This was done by manipulating rhizodeposition with tree girdling in a coniferous subalpine forest in the Rocky Mountains of Colorado, USA, and following changes in the activities of nine exoenzymes involved in decomposition, as well as soil dissolved organic C, dissolved organic and inorganic nitrogen (N), and microbial biomass C and N. We found that rhizodeposition is high in the spring, when the soils are still snow-covered, and that there are large ephemeral populations of microorganisms dependent upon this C. Microbial N acquisition from peptide degradation increased with increases in microbial biomass when rhizodeposition was highest. However, our data indicate that the breakdown of cellulose, lignin, chitin, and organic phosphorus are not affected by springtime increases in soil microbial biomass associated with increases in rhizodeposition. We conclude that the priming of soil C mineralization by rhizodeposition is due to growth of the microbial biomass and an increase in the breakdown of N-rich proteins, but not due to increases in the degradation of plant litter constituents such as cellulose and lignin.

Climatic influences on net ecosystem CO2 exchange during the transition from wintertime carbon source to springtime carbon sink in a high-elevation, subalpine forest.

The transition between wintertime net carbon loss and springtime net carbon assimilation has an important role in controlling the annual rate of carbon uptake in coniferous forest ecosystems. We studied the contributions of springtime carbon assimilation to the total annual rate of carbon uptake and the processes involved in the winter-to-spring transition across a range of scales from ecosystem CO2 fluxes to chloroplast photochemistry in a coniferous, subalpine forest. We observed numerous initiations and reversals in the recovery of photosynthetic CO2 uptake during the initial phase of springtime recovery in response to the passage of alternating warm- and cold-weather systems. Full recovery of ecosystem carbon uptake, whereby the 24-h cumulative sum of NEE (NEEdaily) was consistently negative, did not occur until 3-4 weeks after the first signs of photosynthetic recovery. A key event that preceded full recovery was the occurrence of isothermality in the vertical profile of snow temperature across the snow pack; thus, providing consistent daytime percolation of melted snow water through the snow pack. Interannual variation in the cumulative annual NEE (NEEannual) was mostly explained by variation in NEE during the snow-melt period (NEEsnow-melt), not variation in NEE during the snow-free part of the growing season (NEEsnow-free). NEEsnow-melt was highest in those years when the snow melt occurred later in the spring, leading us to conclude that in this ecosystem, years with earlier springs are characterized by lower rates of NEEannual, a conclusion that contrasts with those from past studies in deciduous forest ecosystems. Using studies on isolated branches we showed that the recovery of photosynthesis occurred through a series of coordinated physiological and biochemical events. Increasing air temperatures initiated recovery through the upregulation of PSII electron transport caused in part by disengagement of thermal energy dissipation by the carotenoid, zeaxanthin. The availability of liquid water permitted a slightly slower recovery phase involving increased stomatal conductance. The most rate-limiting step in the recovery process was an increase in the capacity for the needles to use intercellular CO2, presumably due to slow recovery of Rubisco activity. Interspecific differences were observed in the timing of photosynthetic recovery for the dominant tree species. The results of our study provide (1) a context for springtime CO2 uptake within the broader perspective of the annual carbon budget in this subalpine forest, and (2) a mechanistic explanation across a range of scales for the coupling between springtime climate and the carbon cycle of high-elevation coniferous forest ecosystems.