In a forest ecosystem at steady state, net carbon (C) assimilation by plants and C loss through soil and litter decomposition by heterotrophic organisms are balanced. However, a perturbation to the system, such as increased mean soil temperature, will lead to faster decay, enhancing CO 2 release from decomposers, and thus upsetting the balance. Recent in situ experiments have indicated that the stimulation of soil respiration following a step increase in annual average soil temperature declines over time. One possible explanation for this decline may be changes in substrate availability. This hypothesis is examined by using the ecosystem model G'DAY, which simulates C and nitrogen (N) dynamics in plants and soil.We applied the model to observations from a soil-warming experiment in a Norway spruce (Picea abies (L.) Karst.) stand by simulating a step increase of soil temperature. The model provided a good qualitative reproduction of the observed reduction of heterotrophic respiration (R h ) under sustained warming. The simulations showed how the combined effects of faster turnover and reduced substrate availability lead to a transient increase of R h . The simulated annual increase in R h from soil was 60% in the first year after perturbation but decreased to 30% after a decade.One conclusion from the analysis of the simulations is that R h can decrease even though the temperature response function for decomposition remains unchanged. G'DAY suggests that acclimation of R h to soil warming is partly an effect of substrate depletion of labile C pools during the first decade of warming as a result of accelerated rates of mineralization. The response is attributed mainly to changing levels of C in pools with short time constants, reflecting the importance of high-quality soil C fractions. Changes of the structure or physiology of the decomposer community were not invoked. Therefore, it becomes a question of definition whether the simulated dynamics of the declining response of CO 2 release to the warming should be named acclimation or seen as a natural part of the system dynamics.
Experimental evidence indicates that the stomatal conductance and nitrogen concentration ([N]) of foliage decline under CO2 enrichment, and that the percentage growth response to elevated CO2 is amplified under water limitation, but reduced under nitrogen limitation. We advance simple explanations for these responses based on an optimisation hypothesis applied to a simple model of the annual carbon–nitrogen–water economy of trees growing at a CO2-enrichment experiment at Oak Ridge, Tennessee, USA. The model is shown to have an optimum for leaf [N], stomatal conductance and leaf area index (LAI), where annual plant productivity is maximised. The optimisation is represented in terms of a trade-off between LAI and stomatal conductance, constrained by water supply, and between LAI and leaf [N], constrained by N supply. At elevated CO2 the optimum shifts to reduced stomatal conductance and leaf [N] and enhanced LAI. The model is applied to years with contrasting rainfall and N uptake. The predicted growth response to elevated CO2 is greatest in a dry, high-N year and is reduced in a wet, low-N year. The underlying physiological explanation for this contrast in the effects of water versus nitrogen limitation is that leaf photosynthesis is more sensitive to CO2 concentration ([CO2]) at lower stomatal conductance and is less sensitive to [CO2] at lower leaf [N].
Stable isotope natural abundance measurements integrate across several biogeochemical processes in ecosystem N and C dynamics. Here, we report trends in natural isotope abundance (d 13 C and d 15 N in plant and soil) along a climosequence of 33 Nothofagus forest stands located within Patagonia, Southern Argentina. We measured 28 different abiotic variables (both climatic variables and soil properties) to characterize environmental conditions at each of the 33 sites. Foliar d 13 C values ranged from À35.4& to À27.7&, and correlated positively with foliar d 15 N values, ranging from À3.7& to 5.2&. Soil d 13 C and d 15 N values reflected the isotopic trends of the foliar tissues and ranged from À29.8& to À25.3&, and À4.8& to 6.4&, respectively, with no significant differences between Nothofagus species (Nothofagus pumilio, Nothofagus antarctica, Nothofagus betuloides). Principal component analysis and multiple regressions suggested that mainly water availability variables (mean annual precipitation), but not soil properties, explained between 42% and 79% of the variations in foliar and soil d 13 C and d 15 N natural abundance, which declined with increased moisture supply. We conclude that a decline in water use efficiency at wetter sites promotes both the depletion of heavy C and N isotopes in soil and plant biomass. Soil d 13 C values were higher than those of the plant tissues and this difference increased as annual precipitation increased. No such differences were apparent when d 15 N values in soil and plant were compared, which indicates that climatic differences contributed more to the overall C balance than to the overall N balance in these forest ecosystems.
Susceptibility to tebufenozide and methoxyfenozide of beet armyworm [Spodoptera exigua (Hübner)] from the southern United States and Thailand was determined through exposure of first and third instars to dipped cotton leaves. Among the field populations evaluated, tebufenozide LC50 values for first and third instars, respectively, ranged from 0.377 to 4.41 and 4.37-46.6 microg (AI) /ml of solution. Methoxyfenozide LC50 values for first and third instars of field populations ranged from 0.058 to 0.487 and 0.601-3.83 microg (AI)/ml of solution. A Thailand field strain exhibiting reduced susceptibility to both compounds was subjected to intense laboratory selection for three nonconsecutive generations. At the LC50 and LC90, selected Thailand strains were 45-68 times and 150-1,500 times less susceptible to tebufenozide and 340-320 times and 120-67 times less susceptible to methoxyfenozide as first and third instars, respectively, when compared with the laboratory reference strain. Among the U.S. field populations evaluated, ones from Belle Glade, FL, and Florence, SC, were generally the most susceptible and ones from Maricopa and Parker, AZ, were the least susceptible. Selection of the Thailand field strain with tebufenozide reduced susceptibility to both compounds, and selection of Thailand strains previously pressured with either compound further reduced susceptibility to both, suggesting at least some commonality of resistance mechanism. Characterization of this resistance will provide information that will be helpful for pro-active management of resistance for this valuable group of insecticides.
Susceptibility to spinosad (Success 1 /Tracer 1 ) of beet armyworm (Spodoptera exigua) from the southern USA and Southeast Asia was determined through exposure of second-and third-instar larvae to dipped cotton leaves. LC 50 estimates of susceptibility of second-and third-instar larvae of ®eld populations ranged from 0.279 to 6.14 and 0.589 to 14.0 mg spinosad litre À1 , respectively. A Thailand population was 22-and 24-fold less susceptible than the six other US ®eld populations evaluated, and 85-and 58-fold less susceptible than a reference laboratory population, respectively. From these results, we initiated experiments to test the hypothesis that the Thailand population was resistant to spinosad. F 1 crosses between the resistant Thailand population and a susceptible reference strain yielded individuals that were 22-fold less sensitive to spinosad than the susceptible parent. This same resistant strain exhibited signi®cantly greater survivorship on plants treated with spinosad in the ®eld. Lastly, selection of an Arizona population resulted in a signi®cant reduction in susceptibility to spinosad, further substantiating the hypothesis of a genetic basis for resistance to spinosad. These ®ndings indicate a vulnerability of this new insecticide to resistance development in beet armyworm and should serve as a warning against excessive use of it.
CO2-enrichment experiments consistently show that rooting depth increases when trees are grown at elevated CO2 (eCO2), leading in some experiments to increased capture of available soil nitrogen (N) from deeper soil. However, the link between N uptake and root distributions remains poorly represented in forest ecosystem and global land-surface models. Here, this link is modeled and analyzed using a new optimization hypothesis (MaxNup) for root foraging in relation to the spatial variability of soil N, according to which a given total root mass is distributed vertically in order to maximize annual N uptake. MaxNup leads to analytical predictions for the optimal vertical profile of root biomass, maximum rooting depth, and N-uptake fraction (i.e., the proportion of plant-available soil N taken up annually by roots). We use these predictions to gain new insight into the behavior of the N-uptake fraction in trees growing at the Oak Ridge National Laboratory free-air CO2-enrichment experiment. We also compare MaxNup with empirical equations previously fitted to root-distribution data from all the world's plant biomes, and find that the empirical equations underestimate the capacity of root systems to take up N.
[1] The response of plant ecosystems to environmental change will determine whether the terrestrial biosphere will remain a substantial carbon sink or become a source during the next century. We use two ecosystem models, the Generic Decomposition And Yield model (G'DAY) and the daily time step version of the Century model (DAYCENT), to simulate net ecosystem productivity (NEP) for three contrasting ecosystems (shortgrass steppe in Colorado, tallgrass prairie in Kansas, and Norway spruce in Sweden) with varying degrees of water, temperature, and nutrient limitation, to determine responses to gradual increases in atmospheric CO 2 concentration ([CO 2 ]), temperature, and nitrogen input over 100 years. Using G'DAY, under rising [CO 2 ], there is evidence of C sink ''saturation,'' defined here as positive NEP reaching an upper limit and then declining toward zero, at all three sites (due largely to increased N immobilization in soil organic matter) but a positive C sink is sustained throughout the 100 years. DAYCENT also predicts a sustained C sink at all three sites under rising [CO 2 ], with evidence of C sink saturation for the Colorado grassland and the C sink levels off after 80 years for the Kansas grassland. Warming reduces soil C and the C sink in both grassland ecosystems but increases the C sink in the forest. Warming increases decomposition and soil N mineralization, which stimulates net primary productivity (NPP) at all sites except when inducing water limitation. At the forest site some of the enhanced N release is allocated to a woody biomass pool with a low N:C ratio so that warming enhances NEP without increased N input at the forest site, but not at the grassland sites. Responses to combinations of treatments are generally additive for DAYCENT but more interactive for G'DAY, especially under combined rising [CO 2 ] and warming at the strongly waterand N-limited shortgrass steppe. Increasing N input alleviates C sink saturation and enhances NEP, NPP, and soil C at all sites. At the water-limited grassland sites the effect of rising [CO 2 ] on growth is greatest during the drier seasons. Key sensitivities in the simulations of NEP are identified and include NPP sensitivity to gradual increase in [CO 2 ], N immobilization as a long-term feedback, and the presence or not of plant biomass pools with low N:C ratio.
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