Impacts of solar ultraviolet-B radiation on terrestrial ecosystems of Tierra del Fuego (southern Argentina). An overview of recent progress.

The southern part of Tierra del Fuego, in the southernmost tip of South America, is covered by dense Nothofagus spp. forests and Sphagnum-dominated peat bogs, which are subjected to the influence of ozone depletion and to increased levels of solar ultraviolet-B radiation (UV-B). Over the last 5 years we have studied some of the biological impacts of solar UV-B on natural ecosystems of this region. We have addressed two general problems: (i) do the fluctuations in UV-B levels under the influence of the Antarctic ozone 'hole' have any measurable biological impact, and (ii) what are the long-term effects of solar (ambient) UV-B on the Tierra del Fuego ecosystems? In this paper, we provide an overview of the progress made during the first 4 years of the project. We highlight and discuss the following results: (1) ambient UV-B has subtle but significant inhibitory effects on the growth of herbaceous and graminoid species of this region (growth reduction < or = 12%), whereas no consistent inhibitory effects could be detected in woody perennials; (2) in the species investigated in greatest detail, Gunnera magellanica, the inhibitory effect of solar UV-B is accompanied by increased levels of DNA damage in leaf tissue, and the DNA damage density in the early spring is clearly correlated with the dose of weighted UV-B measured at ground level; (3) the herbaceous species investigated thus far show little or no acclimation responses to ambient UV-B such as increased sunscreen levels and DNA repair capacity; and (4) ambient UV-B has significant effects on heterotrophic organisms, included marked inhibitory effects on insect herbivory. The results from the experiments summarized in this review clearly indicate that UV-B influences several potentially important processes and ecological interactions in the terrestrial ecosystems of Tierra del Fuego.

Consequences of changing biodiversity.

Human alteration of the global environment has triggered the sixth major extinction event in the history of life and caused widespread changes in the global distribution of organisms. These changes in biodiversity alter ecosystem processes and change the resilience of ecosystems to environmental change. This has profound consequences for services that humans derive from ecosystems. The large ecological and societal consequences of changing biodiversity should be minimized to preserve options for future solutions to global environmental problems.

Global biodiversity scenarios for the year 2100.

Scenarios of changes in biodiversity for the year 2100 can now be developed based on scenarios of changes in atmospheric carbon dioxide, climate, vegetation, and land use and the known sensitivity of biodiversity to these changes. This study identified a ranking of the importance of drivers of change, a ranking of the biomes with respect to expected changes, and the major sources of uncertainties. For terrestrial ecosystems, land-use change probably will have the largest effect, followed by climate change, nitrogen deposition, biotic exchange, and elevated carbon dioxide concentration. For freshwater ecosystems, biotic exchange is much more important. Mediterranean climate and grassland ecosystems likely will experience the greatest proportional change in biodiversity because of the substantial influence of all drivers of biodiversity change. Northern temperate ecosystems are estimated to experience the least biodiversity change because major land-use change has already occurred. Plausible changes in biodiversity in other biomes depend on interactions among the causes of biodiversity change. These interactions represent one of the largest uncertainties in projections of future biodiversity change.

Ecosystem water fluxes for two grasslands in elevated CO2: a modeling analysis.

The need to combine data from CO2 field experiments with climate data remains urgent, particularly because each CO2 experiment cannot run for decades to centuries. Furthermore, predictions for a given biome need to take into account differences in productivity and leaf area index (LAI) independent of CO2-derived changes. In this study, we use long-term weather records and field data from the Jasper Ridge CO2 experiment in Palo Alto, California, to model the effects of CO2 and climate variability on ecosystem water fluxes. The sandstone and serpentine grasslands at Jasper Ridge provide a range of primary productivity and LAI, with the sandstone as the more productive system. Modeled soil water availability agreed well with published observations of time-domain reflectometry in the CO2 experiment. Simulated water fluxes based on 10-year weather data (January 1985-December 1994) showed that the sandstone grassland had a much greater proportion of water movement through plants than did the serpentine; transpiration accounted for approximately 30% of annual fluxes in the sandstone and only 10% in the serpentine. Although simulated physiological and biomass changes were similar in both grasslands, the consequences of elevated CO2 were greater for the sandstone water budget. Elevated CO2 increased soil drainage by 20% in the sandstone, despite an approximately one-fifth increase in plant biomass; in the serpentine, drainage increased by <10% and soil evaporation was unchanged for the same simulated biomass change. Phenological changes, simulated by a 15-day lengthening of the growing season, had minimal impacts on the water budget. Annual variation in the timing and amount of rainfall was important for water fluxes in both grasslands. Elevated CO2 increased sandstone drainage >50 mm in seven of ten years, but the relative increase in drainage varied from 10% to 300% depending on the year. Early-season transpiration in the sandstone decreased between 26% and 41%, with elevated CO2 resulting in a simulated water savings of 54-76 mm. Even in years when precipitation was similar (e.g., 505 and 479 mm in years 3 and 4), the effect of CO2 varied dramatically. The response of grassland water budgets to CO2 depends on the productivity and structure of the grassland, the amount and timing of rainfall, and CO2-induced changes in physiology. In systems with low LAI, large physiological changes may not necessarily alter total ecosystem water budgets dramatically.

Downward flux of water through roots (i.e. inverse hydraulic lift) in dry Kalahari sands.

Downward transport of water in roots, in the following termed "inverse hydraulic lift," has previously been shown with heat flux techniques. But water flow into deeper soil layers was demonstrated in this study for the first time when investigating several perennial grass species of the Kalahari Desert under field conditions. Deuterium labelling was used to show that water acquired by roots from moist sand in the upper profile was transported through the root system to roots deeper in the profile and released into the dry sand at these depths. Inverse hydraulic lift may serve as an important mechanism to facilitate root growth through the dry soil layers underlaying the upper profile where precipitation penetrates. This may allow roots to reach deep sources of moisture in water-limited ecosystems such as the Kalahari Desert.

Differential use of large summer rainfall events by shrubs and grasses: a manipulative experiment in the Patagonian steppe.

In the Patagonian steppe, years with above-average precipitation (wet years) are characterized by the occurrence of large rainfall events. The objective of this paper was to analyze the ability of shrubs and grasses to use these large events. Shrubs absorb water from the lower layers, grasses from the upper layers, intercepting water that would otherwise reach the layers exploited by shrubs. We hypothesized that both life-forms could use the large rainfalls and that the response of shrubs could be more affected by the presence of grasses than vice versa. We performed a field experiment using a factorial combination of water addition and life-form removal, and repeated it during the warm season of three successive years. The response variables were leaf growth, and soil and plant water potential. Grasses always responded to experimental large rainfall events, and their response was greater in dry than in wet years. Shrubs only used large rainfalls in the driest year, when the soil water potential in the deep layers was low. The presence or absence of one life-form did not modify the response of the other. The magnitude of the increase in soil water potential was much higher in dry than in humid years, suggesting an explanation for the differences among years in the magnitude of the response of shrubs and grasses. We propose that the generally reported poor response of deep-rooted shrubs to summer rainfalls could be because (1) the water is insufficient to reach deep soil layers, (2) the plants are in a dormant phenological status, and/or (3) deep soil layers have a high water potential. The two last situations may result in high deep-drainage losses, one of the most likely explanations for the elsewhere-reported low response of aboveground net primary production to precipitation during wet years.

Maximum rooting depth of vegetation types at the global scale.

The depth at which plants are able to grow roots has important implications for the whole ecosystem hydrological balance, as well as for carbon and nutrient cycling. Here we summarize what we know about the maximum rooting depth of species belonging to the major terrestrial biomes. We found 290 observations of maximum rooting depth in the literature which covered 253 woody and herbaceous species. Maximum rooting depth ranged from 0.3 m for some tundra species to 68 m for Boscia albitrunca in the central Kalahari; 194 species had roots at least 2 m deep, 50 species had roots at a depth of 5 m or more, and 22 species had roots as deep as 10 m or more. The average for the globe was 4.6±0.5 m. Maximum rooting depth by biome was 2.0±0.3 m for boreal forest. 2.1±0.2 m for cropland, 9.5±2.4 m for desert, 5.2±0.8 m for sclerophyllous shrubland and forest, 3.9±0.4 m for temperate coniferous forest, 2.9±0.2 m for temperate deciduous forest, 2.6±0.2 m for temperate grassland, 3.7±0.5 m for tropical deciduous forest, 7.3±2.8 m for tropical evergreen forest, 15.0±5.4 m for tropical grassland/savanna, and 0.5±0.1 m for tundra. Grouping all the species across biomes (except croplands) by three basic functional groups: trees, shrubs, and herbaceous plants, the maximum rooting depth was 7.0±1.2 m for trees, 5.1±0.8 m for shrubs, and 2.6±0.1 m for herbaceous plants. These data show that deep root habits are quite common in woody and herbaceous species across most of the terrestrial biomes, far deeper than the traditional view has held up to now. This finding has important implications for a better understanding of ecosystem function and its application in developing ecosystem models.

A global analysis of root distributions for terrestrial biomes.

Understanding and predicting ecosystem functioning (e.g., carbon and water fluxes) and the role of soils in carbon storage requires an accurate assessment of plant rooting distributions. Here, in a comprehensive literature synthesis, we analyze rooting patterns for terrestrial biomes and compare distributions for various plant functional groups. We compiled a database of 250 root studies, subdividing suitable results into 11 biomes, and fitted the depth coefficient ß to the data for each biome (Gale and Grigal 1987). ß is a simple numerical index of rooting distribution based on the asymptotic equation Y=1-ßd, where d = depth and Y = the proportion of roots from the surface to depth d. High values of ß correspond to a greater proportion of roots with depth. Tundra, boreal forest, and temperate grasslands showed the shallowest rooting profiles (ß=0.913, 0.943, and 0.943, respectively), with 80-90% of roots in the top 30 cm of soil; deserts and temperate coniferous forests showed the deepest profiles (ß=0.975 and 0.976, respectively) and had only 50% of their roots in the upper 30 cm. Standing root biomass varied by over an order of magnitude across biomes, from approximately 0.2 to 5 kg m-2. Tropical evergreen forests had the highest root biomass (5 kg m-2), but other forest biomes and sclerophyllous shrublands were of similar magnitude. Root biomass for croplands, deserts, tundra and grasslands was below 1.5 kg m-2. Root/shoot (R/S) ratios were highest for tundra, grasslands, and cold deserts (ranging from 4 to 7); forest ecosystems and croplands had the lowest R/S ratios (approximately 0.1 to 0.5). Comparing data across biomes for plant functional groups, grasses had 44% of their roots in the top 10 cm of soil. (ß=0.952), while shrubs had only 21% in the same depth increment (ß=0.978). The rooting distribution of all temperate and tropical trees was ß=0.970 with 26% of roots in the top 10 cm and 60% in the top 30 cm. Overall, the globally averaged root distribution for all ecosystems was ß=0.966 (r 2=0.89) with approximately 30%, 50%, and 75% of roots in the top 10 cm, 20 cm, and 40 cm, respectively. We discuss the merits and possible shortcomings of our analysis in the context of root biomass and root functioning.

Rooting depth, water availability, and vegetation cover along an aridity gradient in Patagonia.

Above-and belowground biomass distribution, isotopic composition of soil and xylem water, and carbon isotope ratios were studied along an aridity gradient in Patagonia (44-45°S). Sites, ranging from those with Nothofagus forest with high annual rainfall (770 mm) to Nothofagus scrub (520 mm), Festuca (290 mm) and Stipa (160 mm) grasslands and into desert vegetation (125 mm), were chosen to test whether rooting depth compensates for low rainfall. Along this gradient, both mean above-and belowground biomass and leaf area index decreased, but average carbon isotope ratios of sun leaves remained constant (at-27‰), indicating no major differences in the ratio of assimilation to stomatal conductance at the time of leaf growth. The depth of the soil horizon that contained 90% of the root biomass was similar for forests and grasslands (about 0.80-0.50 m), but was shallower in the desert (0.30 m). In all habitats, roots reached water-saturated soils or ground water at 2-3 m depth. The depth profile of oxygen and hydrogen isotope ratios of soil water corresponded inversely to volumetric soil water contents and showed distinct patterns throughout the soil profile due to evaporation, water uptake and rainfall events of the past year. The isotope ratios of soil water indicated that high soil moisture at 2-3 m soil depth had originated from rainy periods earlier in the season or even from past rainy seasons. Hydrogen and oxygen isotope ratios of xylem water revealed that all plants used water from recent rain events in the topsoil and not from water-saturated soils at greater depth. However, this study cannot explain the vegetation zonation along the transect on the basis of water supply to the existing plant cover. Although water was accessible to roots in deeper soil layers in all habitats, as demonstrated by high soil moisture, earlier rain events were not fully utilized by the current plant cover during summer drought. The role of seedling establishment in determining species composition and vegetation type, and the indirect effect of seedling establishment on the use of water by fully developed plant cover, are discussed in relation to climate change and vegetation modelling.

CO2 alters water use, carbon gain, and yield for the dominant species in a natural grassland.

Global atmospheric CO2 is increasing at a rate of 1.5-2 ppm per year and is predicted to double by the end of the next century. Understanding how terrestrial ecosystems will respond in this changing environment is an important goal of current research. Here we present results from a field study of elevated CO2 in a California annual grassland. Elevated CO2 led to lower leaf-level stomatal conductance and transpiration (approximately 50%) and higher mid-day leaf water potentials (30-35%) in the most abundant species of the grassland, Avena barbata Brot. Higher CO2 concentrations also resulted in greater midday photosynthetic rates (70% on average). The effects of CO2 on stomatal conductance and leaf water potential decreased towards the end of the growing season, when Avena began to show signs of senescence. Water-use efficiency was approximately doubled in elevated CO2, as estimated by instantaneous gas-exchange measurements and seasonal carbon isotope discrimination. Increases in CO2 and photosynthesis resulted in more seeds per plant (30%) and taller and heavier plants (27% and 41%, respectively). Elevated CO2 also reduced seed N concentrations (9%).

Long-Term Forage Production of North American Shortgrass Steppe.

We evaluated the relationship between annual forage production and annual and seasonal precipitation and temperature at a shortgrass steppe site in north-central Colorado using a long-term data set (52 yr). We also constructed a relationship between forage production and aboveground net primary production (ANPP). Precipitation fluctuated randomly, but temperature had clear warming and cooling trends including a 17-yr warming trend from 1974 to 1990. Forage production was significantly related to both annual and seasonal precipitation but not temperature. Precipitation events between 15 and 30 mm accounted for most of the variability in production because they accounted for most of the variability in precipitation and because they wetted the soil layers that have the largest effect on production. Forage production amplified variability in annual precipitation. Production showed time lags of several years in responding to increases in precipitation. Change in vegetation structure has a characteristic response time, which contrains production responses in wet years. Constraint caused by vegetation structure is the reason why regional ANPP-precipitation models have a steeper slope than long-term models and point out a weakness of exchanging space for time in predicting production patterns.

Effect of animal husbandry on herbivore-carrying capacity at a regional scale.

All significant properties of the herbivore trophic level, including biomass, consumption and productivity, are significantly correlated with primary productivity across a broad range of terrestrial ecosystems. Here we show that livestock biomass in South American agricultural ecosystems across a 25-fold gradient of primary productivity exhibited a relationship with a slope essentially identical to unmanaged ecosystems, but with a substantially greater y-intercept. Therefore the biomass of herbivores supported per unit of primary productivity is about an order of magnitude greater in agricultural than in natural ecosystems, for a given level of primary production. We also present evidence of an increase in livestock body size with primary productivity, a pattern previously characterized in natural ecosystems. To our knowledge this is the first quantitative documentation at a regional scale of the impact of animal husbandry practices, such as herding, stock selection and veterinary care, on the biomass and size-structure of livestock herds compared with native herbivores.

Resource partitioning between shrubs and grasses in the Patagonian steppe.

Experiments were conducted in the Patagonian steppe in southern South America to test the following hypotheses: (a) grasses take up most of the water from the upper layers of the soil and utilize frequent and short-duration pulses of water availability; (b) shrubs, on the contrary, take up most of the water from the lower layers of the soil and utilize infrequent and long-duration pulses of water availability. Grasses and shrubs were removed selectively and the performance of plants and the availability of soil resources were monitored. Results supported the overall hypothesis that grasses and shrubs in the Patagonian steppe use mainly different resources. Removal of shrubs did not alter grass production but removal of grasses resulted in a small increase in shrub production which was mediated by an increase in deep soil water and in shrub leaf water potential. The efficiency of utilization of resources freed by grass removal was approximately 25%. Shrubs used water exclusively from lower soil layers. Grasses took up most of the water from upper layers but they were also capable of absorbing water from deep layers. This pattern of water partitioning along with the lack of response in leaf nitrogen to the removal treatments suggested that shrubs may be at a disadvantage to grasses with respect to nutrient capture and led to questions about the role of nutrient recirculation, leaching, and nitrogen fixation in the steppe.

Small rainfall events: An ecological role in semiarid regions.

Small precipitation events account for a large proportion of the precipitation received in semiarid regions, and their potential ecological importance has previously been ignored. We investigated the effect of a small rainfall event (5 mm) upon Bouteloua gracilis, the dominant grass species of the central and southern Great Plains of North America. An effect of a small event on leaf water potential and leaf conductance to water vapor was observed in less than 12 h and lasted for up to two days.The remarkable short response time of Bouteloua gracilis to a rainfall stimulus enables this species to utilize small events and, therefore, may influence its persistence as a dominant species in the steppe region.We proposed response times to be one of the major species characteristics determining capacity for utilizing different portions of the water resource of the region. We suggest that small precipitation events are ecologically significant and a qualitatively distinct resource for ecosystems in semiarid regions.

Water status of soil and vegetation in a shortgrass steppe.

In an attempt to describe some major relationships between soil and plant compartments in a shortgrass steppe, the process of water loss from the system and plant water relations throughout a drying cycle were studied. The water supply was manipulated and some soil and plant variables monitored throughout a drying cycle. Leaf conductance and leaf water potential of blue grama (Bouteloua gracilis) were measured periodically at predawn and noon. Soil water content and water potential of different layers were also monitored.Three different periods were distinguished in the water loss process throughout a drying cycle. These distinctions were made taking into account the relative contribution of different soil layers. Leaf conductance and water potential at noon slowly declined throughout the first 50 days of plant growth. After that, they rapidly decreased, reaching values of 0.29 mm s-1 and-5.0 MPa, respectively. The predawn leaf water potential remained unchanged around-0.5 MPa during the first 45 days, then rapidly decreased. This occurred when soil water of the wettest soil layer was near depletion.Predawn leaf water potentials were highly correlated with water potentials of the wettest layer. Leaf conductance and water potential at noon were correlated with effective soil water potential (soil water potential weighted by the root distribution in the profile). We concluded that root surface area limited the water flow through an important part of the day in this semiarid ecosystem. Axial root resistance did not appear important in determining the equilibrium status between leaves and the wettest soil layer.