Litter N retention over winter for a low and a high phenolic species in the alpine tundra

Mycorrhizal fungus colonization of roots may modify plant metal acquisition and tolerance. In the present study, the contribution of the extraradical mycelium of an arbuscular mycorrhizal (AM) fungus, Glomus mosseae (BEG 107), to the uptake of metal cations (Cu, Zn, Cd and Ni) by cucumber (Cucumis sativus) plants was determined. The influence of the amount of P supplied to the hyphae on the acquisition and partitioning of metal cations in the mycorrhizal plants was also investigated. Pots with three compartments were used to separate root and root-free hyphal growing zones. The shoot concentration of Cd and Ni was decreased in mycorrhizal plants compared to non-mycorrhizal plants. In contrast, shoot Zn and Cu concentrations were increased in mycorrhizal plants. High P supply to hyphae resulted in decreased root Cu concentrations and shoot Cd and Ni concentrations in mycorrhizal plants. These results confirm that some elements required for plant growth (P, Zn, Cu) are taken up by mycorrhizal hyphae and are then transported to the plants. Conversely, Cd and Ni were transported in much smaller amounts by hyphae to the plant, so that arbuscular mycorrhizal fungus colonization could partly protect plants from toxic effects of these elements. Selective uptake and transport of plant essential elements over non-essential elements by AM hyphae, increased growth of mycorrhizal plants, and metal accumulation in the root may all contribute to the successful growth of mycorrhizal plants on metal-rich substrates. These effects are stimulated when hyphae can access sufficient P in soil.     

Predicted responses of arctic and alpine ecosystems to altered seasonality under climate change

Global climate change is already having significant impacts on arctic and alpine ecosystems, and ongoing increases in temperature and altered precipitation patterns will affect the strong seasonal patterns that characterize these temperature‐limited systems. The length of the potential growing season in these tundra environments is increasing due to warmer temperatures and earlier spring snow melt. Here, we compare current and projected climate and ecological data from 20 Northern Hemisphere sites to identify how seasonal changes in the physical environment due to climate change will alter the seasonality of arctic and alpine ecosystems. We find that although arctic and alpine ecosystems appear similar under historical climate conditions, climate change will lead to divergent responses, particularly in the spring and fall shoulder seasons. As seasonality changes in the Arctic, plants will advance the timing of spring phenological events, which could increase plant nutrient uptake, production, and ecosystem carbon (C) gain. In alpine regions, photoperiod will constrain spring plant phenology, limiting the extent to which the growing season can lengthen, especially if decreased water availability from earlier snow melt and warmer summer temperatures lead to earlier senescence. The result could be a shorter growing season with decreased production and increased nutrient loss. These contrasting alpine and arctic ecosystem responses will have cascading effects on ecosystems, affecting community structure, biotic interactions, and biogeochemistry.     

Litter effects of two co-occurring alpine species on plant growth, microbial activity and immobilization of nitrogen

We measured the litter chemistry of two co‐dominant alpine species, Acomastylis rossii, a forb characterized by a low growth rate and N uptake capacity, and Deschampsia caespitosa, a grass characterized by a high growth rate and N uptake capacity, and examined the effect litter of these two species had on the growth of Deschampsia phytometers in a greenhouse. We also examined the influence of litter from the two species on microbial respiration and immobilization of N, in two separate laboratory microcosm experiments and in the field. We hypothesized that Acomastylis litter would reduce plant growth more than Deschampsia litter, corresponding with either 1) suppression of microbial activity and thus a decrease in N mineralization, or 2) by stimulation of microbial biomass and increasing microbial immobilization of N. Relative to Deschampsia litter, Acomastylis litter had higher total water soluble organic carbon (DOC), and higher total phenolic concentration. Deschampsia litter had 30 times higher carbohydrate (primarily glucose and fructose) concentrations than Acomastylis litter. Soil respiration, microbial biomass N, and consumption of DOC and N were higher with the Acomastylis litter treatment than the Deschampsia litter treatment in experimental microcosms, and both respiration and microbial biomass N were higher in field soils under canopies dominated by Acomastylis relative to those dominated by Deschampsia. These results indicate that phenolics in Acomastylis are a C source for soil microorganisms, rather than an inhibitor of microbial activity. Deschampsia phytometers grew significantly less, had higher root: shoot biomass ratios, and acquired less nitrogen in the Acomastylis litter treatment relative to the control and Deschampsia litter treatments. The results of these experiments indicate that Acomastylis litter influences soil N cycling by increasing microbial activity and N immobilization, which may influence N supply to neighboring plants. This mechanism has the potential to influence competitive interactions between Acomastylis and its neighbors.     

Original Articles: Differential Influence of Plant Species on Soil Nitrogen Transformations Within Moist Meadow Alpine Tundra

Plant species can influence nitrogen (N) cycling indirectly through the feedbacks of litter quality and quantity on soil N transformation rates. The goal of this research was to focus on small-scale (within-community) variation in soil N cycling associated with two community dominants of the moist meadow alpine tundra. Within this community, the small-scale patchiness of the two most abundant species (Acomastylis rossii and Deschampsia caespitosa) provides natural variation in species cover within a relatively similar microclimate, thus enabling estimation of the effects of plant species on soil N transformation rates. Monthly rates of soil N transformations were dependent on small-scale variation in both soil microclimate and species cover. The relative importance of species cover compared with soil microclimate increased for months 2 and 3 of the 3-month growing season. Growing-season net N mineralization rates were over ten times greater and nitrification rates were four times greater in Deschampsia patches than in Acomastylis patches. Variability in litter quality [carbon:nitrogen (C:N) and phenolic:N], litter quantity (aboveground and fine-root production), and soil quality (C:N) was associated with three principal components. Variability between the species in litter quality and fine-root production explained 31% of the variation in net N mineralization rates and 36% of net nitrification rates. Site variability across the landscape in aboveground production and soil C:N explained 33% of the variation in net N mineralization rates and 21% of net nitrification rates. Within the moist meadow community, the high spatial variability in soil N transformation rates was associated with differences in the dominant species' litter quality and fine-root production. Deschampsia-dominated patches consistently had greater soil N transformation rates than did Acomastylis-dominated patches across the landscape, despite site variability in soil moisture, soil C:N, and aboveground production. Plant species appear to be an important control of soil N transformation in the alpine tundra, and consequently may influence plant community structure and ecosystem function.     

Inference of allelopathy is complicated by effects of activated carbon on plant growth

Allelopathy can play an important role in structuring plant communities, but allelopathic effects are often difficult to detect because many methods used to test for allelopathy can be confounded by experimental artifacts. The use of activated carbon, a technique for neutralizing allelopathic compounds, is now employed in tests for allelopathy; however, this technique also could produce large experimental artifacts. In three independent experiments, it was shown that adding activated carbon to potting media affected nutrient availability and plant growth. For most species tested, activated carbon increased plant biomass, even in the absence of the potentially allelopathic agent. The increased growth corresponded to increased plant nitrogen content, likely resulting from greater nitrogen availability. Activated carbon also affected nitrogen and other nutrient concentrations in soil media in the absence of plants. The observed effects of activated carbon on plant growth can confound its use to test for allelopathy. The detection of allelopathy relies on the difference between plant growth in medium with carbon and that in medium without carbon in the presence of the potentially allelopathic competitor; however, this difference may be biased if activated carbon alters soil nutrient availability and plant growth even in the absence of the focal allelopathic agent.     

The temperature responses of soil respiration in deserts: a seven desert synthesis

The temperature response of soil respiration in deserts is not well quantified. We evaluated the response of respiration to temperatures spanning 67°C from seven deserts across North America and Greenland. Deserts have similar respiration rates in dry soil at 20°C, and as expected, respiration rates are greater under wet conditions, rivaling rates observed for more mesic systems. However, deserts differ in their respiration rates under wet soil at 20°C and in the strength of the effect of current and antecedent soil moisture on the sensitivity and magnitude of respiration. Respiration increases with temperature below 30°C but declines for temperatures exceeding 35°C. Hot deserts have lower temperature sensitivity than cold deserts, and insensitive or negative temperature sensitivities were predicted under certain moisture conditions that differed among deserts. These results have implications for large-scale modeling efforts because we highlight the unique behavior of desert soil respiration relative to other systems. These behaviors include variable temperature responses and the importance of antecedent moisture conditions for soil respiration.     

Complex carbon cycle responses to multi‐level warming and supplemental summer rain in the high Arctic

The Arctic has experienced rapid warming and, although there are uncertainties, increases in precipitation are projected to accompany future warming. Climate changes are expected to affect magnitudes of gross ecosystem photosynthesis (GEP), ecosystem respiration (ER) and the net ecosystem exchange of CO₂ (NEE). Furthermore, ecosystem responses to climate change are likely to be characterized by nonlinearities, thresholds and interactions among system components and the driving variables. These complex interactions increase the difficulty of predicting responses to climate change and necessitate the use of manipulative experiments. In 2003, we established a long‐term, multi‐level and multi‐factor climate change experiment in a polar semidesert in northwest Greenland. Two levels of heating (30 and 60 W m^?2) were applied and the higher level was combined with supplemental summer rain. We made plot‐level measurements of CO₂ exchange, plant community composition, foliar nitrogen concentrations, leaf δ¹³C and NDVI to examine responses to our treatments at ecosystem‐ and leaf‐levels. We confronted simple models of GEP and ER with our data to test hypotheses regarding key drivers of CO₂ exchange and to estimate growing season CO₂‐C budgets. Low‐level warming increased the magnitude of the ecosystem C sink. Meanwhile, high‐level warming made the ecosystem a source of C to the atmosphere. When high‐level warming was combined with increased summer rain, the ecosystem became a C sink of magnitude similar to that observed under low‐level warming. Competition among our ER models revealed the importance of soil moisture as a driving variable, likely through its effects on microbial activity and nutrient cycling. Measurements of community composition and proxies for leaf‐level physiology suggest GEP responses largely reflect changes in leaf area of Salix arctica, rather than changes in leaf‐level physiology. Our findings indicate that the sign and magnitude of the future High Arctic C budget may depend upon changes in summer rain.     

Greater deciduous shrub abundance extends tundra peak season and increases modeled net CO2 uptake

Satellite studies of the terrestrial Arctic report increased summer greening and longer overall growing and peak seasons since the 1980s, which increases productivity and the period of carbon uptake. These trends are attributed to increasing air temperatures and reduced snow cover duration in spring and fall. Concurrently, deciduous shrubs are becoming increasingly abundant in tundra landscapes, which may also impact canopy phenology and productivity. Our aim was to determine the influence of greater deciduous shrub abundance on tundra canopy phenology and subsequent impacts on net ecosystem carbon exchange (NEE) during the growing and peak seasons in the arctic foothills region of Alaska. We compared deciduous shrub‐dominated and evergreen/graminoid‐dominated community‐level canopy phenology throughout the growing season using the normalized difference vegetation index (NDVI). We used a tundra plant‐community‐specific leaf area index (LAI) model to estimate LAI throughout the green season and a tundra‐specific NEE model to estimate the impact of greater deciduous shrub abundance and associated shifts in both leaf area and canopy phenology on tundra carbon flux. We found that deciduous shrub canopies reached the onset of peak greenness 13 days earlier and the onset of senescence 3 days earlier compared to evergreen/graminoid canopies, resulting in a 10‐day extension of the peak season. The combined effect of the longer peak season and greater leaf area of deciduous shrub canopies almost tripled the modeled net carbon uptake of deciduous shrub communities compared to evergreen/graminoid communities, while the longer peak season alone resulted in 84% greater carbon uptake in deciduous shrub communities. These results suggest that greater deciduous shrub abundance increases carbon uptake not only due to greater leaf area, but also due to an extension of the period of peak greenness, which extends the period of maximum carbon uptake.     

Tree Species Traits Influence Soil Physical,Chemical, and Biological Properties in High Elevation Forests

Background

Previous studies have shown that plants often have species-specific effects on soil properties. In high elevation forests in the Southern Rocky Mountains, North America, areas that are dominated by a single tree species are often adjacent to areas dominated by another tree species. Here, we assessed soil properties beneath adjacent stands of trembling aspen, lodgepole pine, and Engelmann spruce, which are dominant tree species in this region and are distributed widely in North America. We hypothesized that soil properties would differ among stands dominated by different tree species and expected that aspen stands would have higher soil temperatures due to their open structure, which, combined with higher quality litter, would result in increased soil respiration rates, nitrogen availability, and microbial biomass, and differences in soil faunal community composition.

Methodology/Principal Findings

We assessed soil physical, chemical, and biological properties at four sites where stands of aspen, pine, and spruce occurred in close proximity to one-another in the San Juan Mountains, Colorado. Leaf litter quality differed among the tree species, with the highest nitrogen (N) concentration and lowest lignin∶N in aspen litter. Nitrogen concentration was similar in pine and spruce litter, but lignin∶N was highest in pine litter. Soil temperature and moisture were highest in aspen stands, which, in combination with higher litter quality, probably contributed to faster soil respiration rates from stands of aspen. Soil carbon and N content, ammonium concentration, and microbial biomass did not differ among tree species, but nitrate concentration was highest in aspen soil and lowest in spruce soil. In addition, soil fungal, bacterial, and nematode community composition and rotifer, collembolan, and mesostigmatid mite abundance differed among the tree species, while the total abundance of nematodes, tardigrades, oribatid mites, and prostigmatid mites did not.

Conclusions/Significance

Although some soil characteristics were unaffected by tree species identity, our results clearly demonstrate that these dominant tree species are associated with soils that differ in several physical, chemical, and biotic properties. Ongoing environmental changes in this region, e.g. changes in fire regime, frequency of insect outbreaks, changes in precipitation patterns and snowpack, and land-use change, may alter the relative abundance of these tree species over coming decades, which in turn will likely alter the soils.     

Species effects on resource supply rates: do they influence competitive interactions?

We examined two ways in which species effects on nitrogen supply rates could influence species competitive interactions and lead to coexistence between species with very different patterns of resource use. First, through differential effects on the rates of resource cycling, species may modify resources in divergent directions to better tolerate neighbors under self-modified conditions (tolerance modification). Second, plant-induced shifts in resource supply could enhance the degree to which species can suppress individuals (suppression modification). We addressed these hypothesized mechanisms in the non-successional moist meadow alpine tundra, which is codominated by Acomastylis rossii, a slow-growing roseaceous forb with high nutrient retention that is associated with slow rates of nitrogen supply, and Deschampsia caespitosa, a graminoid with rapid growth potential that is associated with fast rates of nitrogen (N) supply. Neighbors inhibited Acomastylis transplants (the species with high N retention) less than Deschampsia transplants (the species with rapid potential growth) in both neighborhood types, and hence neighborhood type did not influence species tolerance to neighbors. Likewise, Acomastylis neighborhoods inhibited transplant growth more than Deschampsia neighborhoods regardless of transplant species identity. When N supply rates were enhanced through repeated N additions in the two neighborhood types or in the presence of each species’ litter, the inhibitory effects associated with Acomastylis were offset to a greater degree than those associated with Deschampsia, as predicted by the suppression modification. These effects appeared to be density or size-dependent, with few effects observed at low density. Our results suggest that species effects on supply rates may influence competitive interactions, particularly if these effects are complemented by other sources of temporal or spatial variation such as pulses in resource availability. This revised version was published online in June 2006 with corrections to the Cover Date.