A laboratory system for examining the influence of light on diel activity of stream invertebrates

I describe a laboratory system for investigating the role of light as a proximate cue for diel changes in locomotor activity and vertical location on the substrate of stream macro-invertebrates. The system consisted of computer-controlled halogen lamps positioned over a laboratory stream in which video-recordings were made of Stenonema modestum mayfly nymphs located on the undersides of unglazed tile substrates. Locomotor activity of study organisms in response to light changes were quantified during computer-programmed and reproducible light/dark (LD) cycles. The system provided the flexibility to simulate a variety of light environments so that the separate influences of light intensity and light change on diel activities of individuals and populations could be examined, which is difficult under natural light conditions. As a group, nymphs responded similarly to simulated twilight (light decrease from 7.9 × 102 to 6.9 × 10−2 μW cm−2 at a constant –1.9 × 10−3 s−1 rate of relative light change) and to natural twilight, suggesting that proposed mechanisms of light control of diel activities in nature can be adequately tested in the simulated environment. However, locomotor activity and vertical movements among individual mayflies were highly variable under controlled conditions, suggesting that physiological differences influence their responses to environmental conditions

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The use of an artificial light system to assess the influence of relative light change on diel activity cycles of nymphs of the mayfly, Stenonema modestum, in the presence and absence of predators

A mechanism by which light controls diel changes in locomotor activity and surface location of mayfly nymphs (Stenonema modestum Banks), named the Stimulus-based Timing and Activity-Rate (STAR) Model, was tested. Nymph movements were video-recorded in time-lapse from underneath unglazed artificial substrates in a laboratory stream. Light/dark cycles were simulated using computer-controlled halogen lamps. Light increases and decreases were generated to maintain constant rates of relative light change throughout simulated twilight periods. Nymph locomotor activity and position on the substrate were measured in response to rate of light change. Experiments tested whether adaptation light intensity (10\sp{-4} or 10\sp{-6} W cm\sp{-2}), time of day (AM or PM), length of the period of light change, or predators, altered nymph responses to light change. Timing of both heightened nocturnal locomotor activity and leaving the substrate were significantly correlated (R\sp2 =.93; p $\u3c$ 0.001 and R\sp2 =.71; p $\u3c$ 0.004, respectively) with rate of relative light decrease. Rate of change in light was a correlated with the difference between daytime and nighttime locomotor activity (R\sp2 =.38, p $\u3c$ 0.02). The onset of nocturnal locomotor activity was advanced when nymphs were adapted to a low daytime light intensity. Lowered daytime light did not change the time mayflies left the undersides of the substrate. There was no difference in the locomotor activity response between AM and PM experiments, but significantly greater numbers of nymphs left the substrate undersides during simulated twilight in the PM experiments (p $\u3c$ 0.009, F\sb{1.14} = 9.3. The difference between daytime and nighttime locomotor activity diminished during shortened periods of light decrease. When the time intervals over which light was reduced became smaller than the latency period of the response, there was no nocturnal increase in locomotor activity. Nymphs left the substrate undersides regardless of the length of time over which light was reduced. Locomotor activity was greater in the presence of fish odor (Notropis comutus and Rhinichthys cataractae) than in water not containing predators. Locomotor activity was reduced during the daytime in the presence of Paragnetina media stoneflies. Synergistic effects between fish and stoneflies resulted in differences in the timing and locomotor activity of both stoneflies and mayflies

Interactions between carbon and nitrogen dynamics in estimating net primary productivity for potential vegetation in North America

We use the terrestrial ecosystem model (TEM), a process-based model, to investigate how interactions between carbon (C) and nitrogen (N) dynamics affect predictions of net primary productivity (NPP) for potential vegetation in North America. Data on pool sizes and fluxes of C and N from intensively studied field sites are used to calibrate the model for each of 17 non-wetland vegetation types. We use information on climate, soils, and vegetation to make estimates for each of 11,299 non-wetland, 0.5° latitude × 0.5° longitude, grid cells in North America. The potential annual NPP and net N mineralization (NETNMIN) of North America are estimated to be 7.032 × 1015 g C yr−1 and 104.6 × 1012 g N yr−1, respectively. Both NPP and NETNMIN increase along gradients of increasing temperature and moisture in northern and temperate regions of the continent, respectively. Nitrogen limitation of productivity is weak in tropical forests, increasingly stronger in temperate and boreal forests, and very strong in tundra ecosystems. The degree to which productivity is limited by the availability of N also varies within ecosystems. Thus spatial resolution in estimating exchanges of C between the atmosphere and the terrestrial biosphere is improved by modeling the linkage between C and N dynamics. We also perform a factorial experiment with TEM on temperate mixed forest in North America to evaluate the importance of considering interactions between C and N dynamics in the response of NPP to an elevated temperature of 2°C. With the C cycle uncoupled from the N cycle, NPP decreases primarily because of higher plant respiration. However, with the C and N cycles coupled, NPP increases because productivity that is due to increased N availability more than offsets the higher costs of plant respiration. Thus, to investigate how global change will affect biosphere-atmosphere interactions, process-based models need to consider linkages between the C and N cycles

Potential net primary productivity in South America: application of a global model

We use a mechanistically based ecosystem simulation model to describe and analyze the spatial and temporal patterns of terrestrial net primary productivity (NPP) in South America. The Terrestrial Ecosystem Model (TEM) is designed to predict major carbon and nitrogen fluxes and pool sizes in terrestrial ecosystems at continental to global scales. Information from intensively studies field sites is used in combination with continental—scale information on climate, soils, and vegetation to estimate NPP in each of 5888 non—wetland, 0.5° latitude °0.5° longitude grid cells in South America, at monthly time steps. Preliminary analyses are presented for the scenario of natural vegetation throughout the continent, as a prelude to evaluating human impacts on terrestrial NPP. The potential annual NPP of South America is estimated to be 12.5 Pg/yr of carbon (26.3 Pg/yr of organic matter) in a non—wetland area of 17.0 ° 106 km2. More than 50% of this production occurs in the tropical and subtropical evergreen forest region. Six independent model runs, each based on an independently derived set of model parameters, generated mean annual NPP estimates for the tropical evergreen forest region ranging from 900 to 1510 g°m—2°yr—1 of carbon, with an overall mean of 1170 g°m—2°yr—1. Coefficients of variation in estimated annual NPP averaged 20% for any specific location in the evergreen forests, which is probably within the confidence limits of extant NPP measurements. Predicted rates of mean annual NPP in other types of vegetation ranged from 95 g°m—2°yr—1 in arid shrublands to 930 g°m@?yr—1 in savannas, and were within the ranges measured in empirical studies. The spatial distribution of predicted NPP was directly compared with estimates made using the Miami mode of Lieth (1975). Overall, TEM predictions were °10% lower than those of the Miami model, but the two models agreed closely on the spatial patterns of NPP in south America. Unlike previous models, however, TEM estimates NPP monthly, allowing for the evaluation of seasonal phenomena. This is an important step toward integration of ecosystem models with remotely sensed information, global climate models, and atmospheric transport models, all of which are evaluated at comparable spatial and temporal scales. Seasonal patterns of NPP in South America are correlated with moisture availability in most vegetation types, but are strongly influenced by seasonal differences in cloudiness in the tropical evergreen forests. On an annual basis, moisture availability was the factor that was correlated most strongly with annual NPP in South America, but differences were again observed among vegetation types. These results allow for the investigation and analysis of climatic controls over NPP at continental scales, within and among vegetation types, and within years. Further model validation is needed. Nevertheless, the ability to investigate NPP—environment interactions with a high spatial and temporal resolution at continental scales should prove useful if not essential for rigorous analysis of the potential effects of global climate changes on terrestrial ecosystems

Analyzing the discharge regime of a large tropical river through remote sensing, ground-based climatic data, and modeling

This study demonstrates the potential for applying passive microwave satellite sensor data to infer the discharge dynamics of large river systems using the main stem Amazon as a test case. The methodology combines (1) interpolated ground-based meteorological station data, (2) horizontally and vertically polarized temperature differences (HVPTD) from the 37-GHz scanning multichannel microwave radiometer (SMMR) aboard the Nimbus 7 satellite, and (3) a calibrated water balance/water transport model (WBM/WTM). Monthly HVPTD values at 0.25° (latitude by longitude) resolution were resampled spatially and temporally to produce an enhanced HVPTD time series at 0.5° resolution for the period May 1979 through February 1985. Enhanced HVPTD values were regressed against monthly discharge derived from the WBM/WTM for each of 40 grid cells along the main stem over a calibration period from May 1979 to February 1983 to provide a spatially contiguous estimate of time-varying discharge. HVPTD-estimated flows generated for a validation period from March 1983 to February 1985 were found to be in good agreement with both observed arid modeled discharges over a 1400-km section of the main stem Amazon. This span of river is bounded downstream by a region of tidal influence and upstream by low sensor response associated with dense forest canopy. Both the WBM/WTM and HVPTD-derived flow rates reflect the significant impact of the 1982–1983 El Niño-;Southern Oscillation (ENSO) event on water balances within the drainage basin

Equilibrium responses of global net primary production and carbon storage to doubled atmospheric carbon dioxide: sensitivity to changes in vegetation nitrogen concentration

We ran the terrestrial ecosystem model (TEM) for the globe at 0.5° resolution for atmospheric CO2 concentrations of 340 and 680 parts per million by volume (ppmv) to evaluate global and regional responses of net primary production (NPP) and carbon storage to elevated CO2 for their sensitivity to changes in vegetation nitrogen concentration. At 340 ppmv, TEM estimated global NPP of 49.0 1015 g (Pg) C yr−1 and global total carbon storage of 1701.8 Pg C; the estimate of total carbon storage does not include the carbon content of inert soil organic matter. For the reference simulation in which doubled atmospheric CO2 was accompanied with no change in vegetation nitrogen concentration, global NPP increased 4.1 Pg C yr−1 (8.3%), and global total carbon storage increased 114.2 Pg C. To examine sensitivity in the global responses of NPP and carbon storage to decreases in the nitrogen concentration of vegetation, we compared doubled CO2 responses of the reference TEM to simulations in which the vegetation nitrogen concentration was reduced without influencing decomposition dynamics (“lower N” simulations) and to simulations in which reductions in vegetation nitrogen concentration influence decomposition dynamics (“lower N+D” simulations). We conducted three lower N simulations and three lower N+D simulations in which we reduced the nitrogen concentration of vegetation by 7.5, 15.0, and 22.5%. In the lower N simulations, the response of global NPP to doubled atmospheric CO2 increased approximately 2 Pg C yr−1 for each incremental 7.5% reduction in vegetation nitrogen concentration, and vegetation carbon increased approximately an additional 40 Pg C, and soil carbon increased an additional 30 Pg C, for a total carbon storage increase of approximately 70 Pg C. In the lower N+D simulations, the responses of NPP and vegetation carbon storage were relatively insensitive to differences in the reduction of nitrogen concentration, but soil carbon storage showed a large change. The insensitivity of NPP in the N+D simulations occurred because potential enhancements in NPP associated with reduced vegetation nitrogen concentration were approximately offset by lower nitrogen availability associated with the decomposition dynamics of reduced litter nitrogen concentration. For each 7.5% reduction in vegetation nitrogen concentration, soil carbon increased approximately an additional 60 Pg C, while vegetation carbon storage increased by only approximately 5 Pg C. As the reduction in vegetation nitrogen concentration gets greater in the lower N+D simulations, more of the additional carbon storage tends to become concentrated in the north temperate-boreal region in comparison to the tropics. Other studies with TEM show that elevated CO2 more than offsets the effects of climate change to cause increased carbon storage. The results of this study indicate that carbon storage would be enhanced by the influence of changes in plant nitrogen concentration on carbon assimilation and decomposition rates. Thus changes in vegetation nitrogen concentration may have important implications for the ability of the terrestrial biosphere to mitigate increases in the atmospheric concentration of CO2 and climate changes associated with the increases

Direct observations of the activity responses of mayfly nymphs to relative light change and light intensity

Studies were conducted in a laboratory stream to examine changes in locomotor activity and vertical movement on tile substrates of Stenonema modestum nymphs (Ephemeroptera: Heptagenidae) in response to relative light decrease and acclimation (adaptation) light intensity. Evening twilight, a period during which activity and location changes of mayfly nymphs including S. modestum occur in nature, was simulated by reducing light at constant rates of relative light decrease using a computer-controlled lighting system. Nymphs were kept at one of two light levels prior to the light reduction to test for changes in their response to light caused by visual adaptation to ambient light intensity. The timing of both increased locomotor activity and changes in vertical location on the substrate of S. modestum was strongly correlated with the strength of the relative decrease in light intensity (light stimulus). Timing of both activities took place earlier for nymphs adapted for 60 min at a constant, but reduced light intensity at the water\u27s surface of 8 µW/cm2 than for those adapted for 60 min at a constant noon-time light intensity of 800 µW/cm2 Individual nymph responses to light change were contingent upon their initial activity during the light-adaptation periods. Relative light change was the controlling factor initiating both the photokinetic (the non-directional locomotor activity on the lower substrate surfaces) and phototactic (light-directed vertical movements between the lower and upper substrate surfaces) activity changes in these stream invertebrates