Rate of tree carbon accumulation increases continuously with tree size.

Forests are major components of the global carbon cycle, providing substantial feedback to atmospheric greenhouse gas concentrations. Our ability to understand and predict changes in the forest carbon cycle--particularly net primary productivity and carbon storage--increasingly relies on models that represent biological processes across several scales of biological organization, from tree leaves to forest stands. Yet, despite advances in our understanding of productivity at the scales of leaves and stands, no consensus exists about the nature of productivity at the scale of the individual tree, in part because we lack a broad empirical assessment of whether rates of absolute tree mass growth (and thus carbon accumulation) decrease, remain constant, or increase as trees increase in size and age. Here we present a global analysis of 403 tropical and temperate tree species, showing that for most species mass growth rate increases continuously with tree size. Thus, large, old trees do not act simply as senescent carbon reservoirs but actively fix large amounts of carbon compared to smaller trees; at the extreme, a single big tree can add the same amount of carbon to the forest within a year as is contained in an entire mid-sized tree. The apparent paradoxes of individual tree growth increasing with tree size despite declining leaf-level and stand-level productivity can be explained, respectively, by increases in a tree's total leaf area that outpace declines in productivity per unit of leaf area and, among other factors, age-related reductions in population density. Our results resolve conflicting assumptions about the nature of tree growth, inform efforts to undertand and model forest carbon dynamics, and have additional implications for theories of resource allocation and plant senescence.

Chemistry and long-term decomposition of roots of Douglas-fir grown under elevated atmospheric carbon dioxide and warming conditions.

Elevated atmospheric CO(2) concentrations and warming may affect the quality of litters of forest plants and their subsequent decomposition in ecosystems, thereby potentially affecting the global carbon cycle. However, few data on root tissues are available to test this feedback to the atmosphere. In this study, we used fine (diameter < or = 2 mm) and small (2-10 mm) roots of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings that were grown for 4 yr in a 2 x 2 factorial experiment: ambient or elevated (+ 180 ppm) atmospheric CO(2) concentrations, and ambient or elevated (+3.8 degrees C) atmospheric temperature. Exposure to elevated CO(2) significantly increased water-soluble extractives concentration (%WSE), but had little effect on the concentration of N, cellulose, and lignin of roots. Elevated temperature had no effect on substrate quality except for increasing %WSE and decreasing the %lignin content of fine roots. No significant interaction was found between CO(2) and temperature treatments on substrate quality, except for %WSE of the fine roots. Short-term (< or = 9 mo) root decomposition in the field indicated that the roots from the ambient CO(2) and ambient temperature treatment had the slowest rate. However, over a longer period of incubation (9-36 mo) the influence of initial substrate quality on root decomposition diminished. Instead, the location of the field incubation sites exhibited significant control on decomposition. Roots at the warmer, low elevation site decomposed significantly faster than the ones at the cooler, high elevation site. This study indicates that short-term decomposition and long-term responses are not similar. It also suggests that increasing atmospheric CO(2) had little effect on the carbon storage of Douglas-fir old-growth forests of the Pacific Northwest.

Successional changes in live and dead wood carbon stores: implications for net ecosystem productivity.

If forests are to be used in CO2 mitigation projects, it is essential to understand and quantify the impacts of disturbance on net ecosystem productivity (NEP; i.e., the change in ecosystem carbon (C) storage with time). We examined the influence of live tree and coarse woody debris (CWD) on NEP during secondary succession based on data collected along a 500-year chronosequence on the Wind River Ranger District, Washington. We developed a simple statistical model of live and dead wood accumulation and decomposition to predict changes in the woody component of NEP, which we call NEP(w). The transition from negative to positive NEP(w), for a series of scenarios in which none to all wood was left after disturbance, occurred between 0 and 57 years after disturbance. The timing of this transition decreased as live-tree growth rates increased, and increased as CWD left after disturbance increased. Maximum and minimum NEP(w) for all scenarios were 3.9 and -14.1 Mg C ha-1 year-1, respectively. Maximum live and total wood C stores of 319 and 393 Mg C ha(-1), respectively, were reached approximately 200 years after disturbance. Decomposition rates (k) of CWD ranged between 0.013 and 0.043 year-1 for individual stands. Regenerating stands took 41 years to attain a mean live wood mass equivalent to the mean mass of CWD left behind after logging, 40 years to equal the mean CWD mass in 500-year-old forest, and more than 150 years to equal the mean total live and dead wood in an old-growth stand. At a rotation age of 80 years, regenerating stands stored approximately half the wood C of the remaining nearby old-growth forests (predominant age 500 years), indicating that conversion of old-growth forests to younger managed forests results in a significant net release of C to the atmosphere.

Trends in bole biomass accumulation, net primary production and tree mortality in Pseudotsuga menziesii forests of contrasting age.

Although it is generally accepted that the rate of accumulation of biomass declines as forests age, little is known about the relative contributions to this decline of changes in net primary production (NPP) and tree mortality. We used 10-15 years of observations of permanent plots in three small watersheds in and near the H.J. Andrews Experimental Forest, Oregon, to examine these issues. The three watersheds are of similar elevation and potential productivity and support young (29 years at last measurement), mature (approximately 100 years) and old (approximately 400 years) forest dominated by Pseudotsuga menziesii (Mirb.) Franco and Tsuga heterophylla (Raf.) Sarg. Accumulation of tree bole biomass was greatest in the young stand, reaching approximately 7 Mg ha-1 year-1 in the last measurement interval. Bole biomass accumulation was relatively constant (approximately 4-5 Mg ha-1 year-1) in the mature stand, and there was no net accumulation of bole biomass in the old-forest stand. The NPP of boles increased with time in the young stand, from approximately 3 to approximately 7 Mg ha-1 year-1, but was nearly constant in the mature and old-forest stands, at approximately 6 and 3-4 Mg ha-1 year-1, respectively. Mortality increased slowly in the young stand (from < 0.1 to 0.3 Mg ha-1 year-1), but fluctuated between 1-2 and 2-6 Mg ha-1 year-1 in the mature and old-forest stands, respectively. Thus, declining biomass accumulation with stand age reflects, in approximately equal amounts, both decreasing NPP and increasing mortality.

Consistent land- and atmosphere-based U.S. carbon sink estimates.

For the period 1980-89, we estimate a carbon sink in the coterminous United States between 0.30 and 0.58 petagrams of carbon per year (petagrams of carbon = 10(15) grams of carbon). The net carbon flux from the atmosphere to the land was higher, 0.37 to 0.71 petagrams of carbon per year, because a net flux of 0.07 to 0.13 petagrams of carbon per year was exported by rivers and commerce and returned to the atmosphere elsewhere. These land-based estimates are larger than those from previous studies (0.08 to 0.35 petagrams of carbon per year) because of the inclusion of additional processes and revised estimates of some component fluxes. Although component estimates are uncertain, about one-half of the total is outside the forest sector. We also estimated the sink using atmospheric models and the atmospheric concentration of carbon dioxide (the tracer-transport inversion method). The range of results from the atmosphere-based inversions contains the land-based estimates. Atmosphere- and land-based estimates are thus consistent, within the large ranges of uncertainty for both methods. Atmosphere-based results for 1980-89 are similar to those for 1985-89 and 1990-94, indicating a relatively stable U.S. sink throughout the period.

Use of helper-free replication-defective simian immunodeficiency virus-based vectors to study macrophage and T tropism: evidence for distinct levels of restriction in primary macrophages and a T-cell line.

Cell tropism of human and simian immunodeficiency viruses (HIV and SIV, respectively) is governed in part by interactions between the viral envelope protein and the cellular receptors. However, there is evidence that envelope-host cell interactions also affect postentry steps in viral replication. We used a helper-free replication-defective SIV macaque (SIVmac)-based retroviral vector carrying the enhanced jellyfish green fluorescent protein inserted into the nef region (V1EGFP) to examine SIV tropism in a single cycle of infection. Vector stocks containing envelope proteins from three different SIVmac clones, namely, SIVmac239 (T-lymphocyte tropic [T-tropic]), SIVmac316 (macrophage tropic [M-tropic]), and SIVmac1A11 (dualtropic), were tested. SIVmac239 replicates efficiently in many human T-cell lines, but it does not efficiently infect primary rhesus macrophages. Conversely, SIVmac316 efficiently infects primary macrophages, but it does not replicate in Molt4-Clone8 (M4C8) T cells. SIVmac1A11 replicates efficiently in both cell types. When primary macrophages were infected with V1EGFP pseudotyped by SIVmac316 or SIVmac1A11 envelopes, the infection was substantially (ca. 200- to 300-fold) more efficient than for the SIVmac239 pseudotype. Thus, in primary macrophages, a major component of M versus T tropism involves relatively early events in the infection cycle. Quantitative PCR studies indicated that synthesis and transport of vector DNA into the nucleus were similar for macrophages infected with the clone 239 and 316 pseudotypes, suggesting that the restriction for SIVmac239 infection is after reverse transcription and nuclear import of viral DNA. When the same vector pseudotypes were used to infect M4C8 cells, they all showed approximately equivalent infectivities, even though replication-competent SIVmac316 does not continue to replicate in these cells. Therefore, in M4C8 cells, restriction involves a late step in the infection cycle (after proviral integration and expression). Thus, depending on the cell type infected, envelope-dependent cell interactions that govern SIV M and T tropism may involve different steps in infection.

Effects on carbon storage of conversion of old-growth forests to young forests.

Simulations of carbon storage suggest that conversion of old-growth forests to young fast-growing forests will not decrease atmospheric carbon dioxide (CO(2)) in general, as has been suggested recently. During simulated timber harvest, on-site carbon storage is reduced considerably and does not approach old-growth storage capacity for at least 200 years. Even when sequestration of carbon in wooden buildings is included in the models, timber harvest results in a net flux of CO(2) to the atmosphere. To offset this effect, the production of lumber and other long-term wood products, as well as the life-span of buildings, would have to increase markedly. Mass balance calculations indicate that the conversion of 5 x 10(9) to 1.8 x 10(9) megagrams of carbon to the atmosphere.