Ambient Plots Research Articles

Opposing plant community responses to warming with and without herbivores

If controls over primary productivity and plant community composition are mainly environmental, as opposed to biological, then global change may result in large-scale alterations in ecosystem structure and function. This view appears to be favored among investigations of plant biomass and community responses to experimental and observed warming. In far northern and arctic ecosystems, such studies predict increasing dominance of woody shrubs with future warming and emphasize the carbon (C)-sequestration potential and consequent atmospheric feedback potential of such responses. In contrast to previous studies, we incorporated natural herbivory by muskoxen and caribou into a 5-year experimental investigation of arctic plant community response to warming. In accordance with other studies, warming increased total community biomass by promoting growth of deciduous shrubs (dwarf birch and gray willow). However, muskoxen and caribou reduced total community biomass response, and responses of birch and willow, to warming by 19%, 46%, and 11%, respectively. Furthermore, under warming alone, the plant community shifted after 5 years away from graminoid-dominated toward dwarf birch-dominated. In contrast, where herbivores grazed, plant community composition on warmed plots did not differ from that on ambient plots after 5 years. These results highlight the potentially important and overlooked influences of vertebrate herbivores on plant community response to warming and emphasize that conservation and management of large herbivores may be an important component of mitigating ecosystem response to climate change.

Long-term experimental warming alters nitrogen-cycling communities but site factors remain the primary drivers of community structure in high arctic tundra soils

Arctic air temperatures are expected to rise significantly over the next century. Experimental warming of arctic tundra has been shown to increase plant productivity and cause community shifts and may also alter microbial community structure. Hence, the objective of this study was to determine whether experimental warming caused shifts in soil microbial communities by measuring changes in the frequency, relative abundance and/or richness of nosZ and nifH genotypes. Five sites at a high arctic coastal lowland were subjected to a 13-year warming experiment using open-top chambers (OTCs). Sites differed by dominant plant community, soil parent material and/or moisture regimen. Six soil cores were collected from each of four replicate OTC and ambient plots at each site and subdivided into upper and lower samples. Differences in frequency and relative abundance of terminal restriction fragments were assessed graphically by two-way cluster analysis and tested statistically with permutational multivariate analysis of variance (ANOVA). Genotypic richness was compared using factorial ANOVA. The genotype frequency, relative abundance and genotype richness of both nosZ and nifH communities differed significantly by site, and by OTC treatment and/or depth at some sites. The site that showed the most pronounced treatment effect was a wet sedge meadow, where community structure and genotype richness of both nosZ and nifH were significantly affected by warming. Although warming was an important factor affecting these communities at some sites at this high arctic lowland, overall, site factors were the main determinants of community structure.

PHENOLOGICAL SEQUENCES REVEAL AGGREGATE LIFE HISTORY RESPONSE TO CLIMATIC WARMING

Climatic warming is associated with organisms breeding earlier in the season than is typical for their species. In some species, however, response to warming is more complex than a simple advance in the timing of all life history events preceding reproduction. Disparities in the extent to which different components of the reproductive phenology of organisms vary with climatic warming indicate that not all life history events are equally responsive to environmental variation. Here, we propose that our understanding of phenological response to climate change can be improved by considering entire sequences of events comprising the aggregate life histories of organisms preceding reproduction. We present results of a two-year warming experiment conducted on 33 individuals of three plant species inhabiting a low-arctic site. Analysis of phenological sequences of three key events for each species revealed how the aggregate life histories preceding reproduction responded to warming, and which individual events exerted the greatest influence on aggregate life history variation. For alpine chickweed (Cerastium alpinum), warming elicited a shortening of the duration of the emergence stage by 2.5 days on average, but the aggregate life history did not differ between warmed and ambient plots. For gray willow (Salix glauca), however, all phenological events monitored occurred earlier on warmed than on ambient plots, and warming reduced the aggregate life history of this species by 22 days on average. Similarly, in dwarf birch (Betula nana), warming advanced flower bud set, blooming, and fruit set and reduced the aggregate life history by 27 days on average. Our approach provides important insight into life history responses of many organisms to climate change and other forms of environmental variation. Such insight may be compromised by considering changes in individual phenological events in isolation.

Host location behavior of Cotesia plutellae Kurdjumov (Hymenoptera: Braconidae) in ambient and moderately elevated ozone in field conditions

In field O(3)-enrichment experiments increased herbivore densities have been reported, which could be due to negatively affected host location behavior of natural enemies. We addressed the impact of doubling background O(3) on the host location of the parasitoid Cotesia plutellae by conducting 24-h trials in an open-air O(3)-fumigation system during two consecutive years. Two circles (radii 1.40 and 4.00 m) of Plutella xylostella-infested potted cabbage plants were placed in the O(3) and ambient plots. Female wasps were released into each plot from the center, and observed 5 times over a 24-h period to assess their host location capability. Thereafter, plants were kept in laboratory conditions until larvae pupation to determine parasitism rates. No significant differences were detected between ambient and O(3)-enriched environments either in the number of wasps found in the field, or in the percentages of parasitized larvae. This suggests that moderately elevated O(3) will not affect the behavior of this parasitoid.

Soil organic matter dynamics under soybean exposed to elevated [CO2

It is unclear how changing atmospheric composition will influence the plant–soil interactions that determine soil organic matter (SOM) levels in fertile agricultural soils. Positive effects of CO2 fertilization on plant productivity and residue returns should increase SOM stocks unless mineralization or biomass removal rates increase in proportion to offset gains. Our objectives were to quantify changes in SOM stocks and labile fractions in prime farmland supporting a conventionally managed corn–soybean system and the seasonal dynamics of labile C and N in soybean in plots exposed to elevated [CO2] (550 ppm) under free-air concentration enrichment (FACE) conditions. Changes in SOM stocks including reduced C/N ratios and labile N stocks suggest that SOM declined slightly and became more decomposed in all plots after 3 years. Plant available N (>273 mg N kg−1) and other nutrients (Bray P, 22–50 ppm; extractable K, 157–237 ppm; Ca, 2,378–2,730 ppm; Mg, 245–317 ppm) were in the high to medium range. Exposure to elevated [CO2] failed to increase particulate organic matter C (POM-C) and increased POM-N concentrations slightly in the surface depth despite known increases (≈30%) in root biomass. This, and elevated CO2 efflux rates indicate accelerated decay rates in fumigated plots (2001: elevated [CO2]: 10.5 ± 1.2 μmol CO2 m−2 s−1 vs. ambient: 8.9 ± 1.0 μmol CO2 m−2 s−1). There were no treatment-based differences in the within-season dynamics of SOM. Soil POM-C and POM-N contents were slightly greater in the surface depth of elevated than ambient plots. Most studies attribute limited ability of fumigated soils to accumulate SOM to N limitation and/or limited plant response to CO2 fertilization. In this study, SOM turnover appears to be accelerated under elevated [CO2] even though soil moisture and nutrients are non-limiting and plant productivity is consistently increased. Accelerated SOM turnover rates may have long-term implications for soil’s productive potential and calls for deeper investigation into C and N dynamics in highly-productive row crop systems.

EFFECT OF ELEVATED CO2ON COARSE-ROOT BIOMASS IN FLORIDA SCRUB DETECTED BY GROUND-PENETRATING RADAR

Growth and distribution of coarse roots in time and space represent a gap in our understanding of belowground ecology. Large roots may play a critical role in carbon sequestration belowground. Using ground-penetrating radar (GPR), we quantified coarse-root biomass from an open-top chamber experiment in a scrub-oak ecosystem at Kennedy Space Center, Florida, USA. GPR propagates electromagnetic waves directly into the soil and reflects a portion of the energy when a buried object is contacted. In our study, we utilized a 1500 MHz antenna to establish correlations between GPR signals and root biomass. A significant relationship was found between GPR signal reflectance and biomass (R2 = 0.68). This correlation was applied to multiple GPR scans taken from each open-top chamber (elevated and ambient CO2). Our results showed that plots receiving elevated CO2 had significantly (P = 0.049) greater coarse-root biomass compared to ambient plots, suggesting that coarse roots may play a large role in carbon sequestration in scrub-oak ecosystems. This nondestructive method holds much promise for rapid and repeatable quantification of coarse roots, which are currently the most elusive aspect of long-term belowground studies.

Sequestration and turnover of plant‐ and microbially derived sugars in a temperate grassland soil during 7 years exposed to elevated atmospheric pCO2

AbstractTemperate grasslands contribute about 20% to the global terrestrial carbon (C) budget with sugars contributing 10–50% to this soil C pool. Whether the observed increase of the atmospheric CO2 concentration (pCO2) leads to additional C sequestration into these ecosystems or enhanced mineralization of soil organic matter (SOM) is still unclear. Therefore, the aim of the presented study was to investigate the impact of elevated atmospheric pCO2 on C sequestration and turnover of plant‐ (arabinose and xylose) and microbially derived (fucose, rhamnose, galactose, mannose) sugars in soil, representing a labile SOM pool. The study was carried out at the Swiss Free Air Carbon Dioxide Enrichment (FACE) experiment near Zurich. For 7 years, Lolium perenne swards were exposed to ambient and elevated pCO2 (36 and 60 Pa, respectively). The additional CO2 in the FACE plots was depleted in 13C compared with ambient plots, so that ‘new’ (<7 years) C inputs could be determined by means of compound‐specific stable isotope analysis (13C : 12C). Samples were fractionated into clay, silt, fine sand and coarse sand, which yielded relatively stable and labile SOM pools with different turnover rates. Total sugar sequestration into bulk soil after 7 years of exposure to elevated pCO2 was about 28% compared with the control plots. In both ambient and elevated plots, total sugar concentrations in particle size fractions increased in the order sand<silt<clay whereas ratios of microbially‐ to plant‐derived sugars showed an increasing dominance of microbially derived sugars in the order sand<silt<clay. In particle size fractions, total sugar amounts were higher under elevated pCO2 for coarse sand, fine sand and silt (about 274%, 17% and 96%, respectively) but about 14% lower for clay compared with the control plots, corroborating that sugars belong to the labile SOM pool. The fraction of newly produced sugars gradually increased by up to 50% in bulk soil samples after 7 years under elevated pCO2. In the ambient plots, sugars were enriched in 13C by up to 10‰ when compared with bulk soil samples from the same plots. The enrichment of 13C in plant‐derived sugars was up to 13.4‰ when compared with parent plant material. After 7 years, the δ13C values of individual sugars decreased under elevated (13C‐depleted) CO2 in bulk soil and particle size fractions, varying between −13.7‰ and −37.8‰ under elevated pCO2. In coarse and fine sand, silt and clay fractions newly produced sugars made up 106%, 63%, 60% and 45%, respectively, of the total sugars present after 7 years. Mean residence time (MRT) of the sugars were calculated according to two models revealing a few decades, mean values increasing in the order coarse sand<fine sand<silt<clay thus corroborating the model of increasing SOM stability in the same order. Our work clearly demonstrated that higher biomass production under elevated pCO2 led to a net sequestration of about 30% of labile SOM (sugars) while no increase of total organic C was observed at the same plots. The additional labile SOM is gradually incorporated into more stable SOM pools such as silt and clay fractions in the medium term (<7 years). MRT of labile (sugar) SOM under elevated pCO2 is in the same order of magnitude when compared with studies under ambient pCO2 though no direct comparison of elevated and ambient plots was possible.

Impact of changes in rainfall amounts predicted by climate-change models on decomposition in a deciduous forest

Climate-change models predict a more intense hydrological cycle, with both increased and decreased amounts of rainfall in areas covered with temperate deciduous forests. These changes could alter rates of litter decomposition, with consequences for rates of nutrient cycling in the forest ecosystem. To examine impacts of predicted changes in precipitation on the rate of decay of canopy leaves, we placed litterbags in replicated, fenced 14 m 2 low-rainfall and high-rainfall plots located under individual rainout shelters. Unfenced, open plots served as an ambient treatment. Litter in the high-rainfall and ambient plots decayed 50% and 78% faster, respectively, than litter in the low-rainfall plots. Litter in the ambient plots disappeared 20% faster than in the high-rainfall treatment, perhaps via greater leaching during heavy rainfall events. Ambient rainfall during the experiment was similar in total amount to the high-rainfall treatment, but was more variable in intensity and timing. We used litterbags of different mesh sizes to examine whether changes in rainfall might alter the impacts of major categories of the fauna on litter decay. However, we found no consistent evidence that excluding arthropods of different sizes affected litter decay rate within any of the three rainfall treatments. This research reveals that changes in rainfall predicted to occur with global climate change will likely strongly alter rates of litter decay in deciduous forests.

An inexpensive method for remotely monitoring nest activity

ABSTRACT In studies of avian nest success, investigators often face the difficult task of periodically checking nest status while at the same time limiting observer influence on nest survival. Remotely monitoring nests using temperature data loggers is one method that allows for continuous data capture regarding nest status (i.e., active vs. inactive) without the negative effects associated with repeated nest checks. We used small temperature data loggers (Thermochron iButtons) to remotely monitor nests of Long-billed Curlews (Numenius americanus) in northeastern Nevada. Data loggers programmed to record temperature at 10-min and 20-min intervals were placed in curlew nests. Data loggers were set to collect data throughout the nesting cycle to determine onset of incubation and timing of nest failure. On average, Long-billed Curlews began incubating approximately 3 d after the first egg was laid and onset of incubation coincided with the laying of the third egg. iButtons allowed us to determine when incubation was terminated in 17 of 23 unsuccessful Long-billed Curlew nests, including 13 of 17 depredated nests. The presence of iButtons in Long-billed Curlew nests did not affect daily survival rate, egg hatchability or rate of nest abandonment. iButtons are an efficient and practical means for remotely monitoring nests of large egg-laying birds, such as the Long-billed Curlew. SINOPSIS En estudios de exito de anidamiento, los investigadores muchas veces se encuentran con la dificultad de examinar el nido, tratando de limitar la infuencia del observador en la sobrevivencia de las aves. Uno de los metodos de monitoreo remoto es el uso de bitacoras electronicas de temperatura que permiten tomar datos de forma continua en un nido (e.g., activo vs inactivo) sin los efectos negativos asociados con el examen periodico del mismo. Utilizamos bitacoras electronicas de temperatura (Thermochron iButtons) para monitorear nidos de chorlo (Numenius americanus) en el norte de Nevada. El instrumento se programo para tomar la temperatura dentro del nido a intervalos de 10 a 20 minutos. El instrumento se utilizo para tomar datos a traves del ciclo de anidamiento del ave para determinar el comienzo de la incubacion y el momento en que el nido fuera abandonado. En promedio los chorlos comienzan a incubar aproximadamente tres dias despues de puesto el primer huevo, lo que coincide con la puesta del tercer huevo. iButtons permitio determinar el final del periodo de incubacion de 13 de 17 nidos depredados. La presencia del instrumento en los nidos de chorlo no afecto la tasa diaria de sobrevivencia, tampoco el eclosionamiento o la tasa de abandono. Las bitacoras utilizadas son eficientes y ademas una forma practica de monitorear a remoto los nidos de aves grandes tales como el chorlo estudiado.

Effects of Arenicola marina on polychaete functional diversity revealed by large-scale experimental lugworm exclusion

The lugworm Arenicola marina was excluded from sandy sediment areas in the mid and low intertidal zone of the Wadden Sea. Exclusion, control and ambient plots were 400 m 2 each, replicated six times and sampled in August of three consecutive years. Responses were analysed with respect to functional trait groups in the associated polychaete assemblage using uni- and multivariate statistical techniques. Tube-building worms and predacious worms were most abundant in exclusion plots, while subsurface deposit feeders tended to dominate in the presence of lugworms. Lugworm effects were stronger in low intertidal fine sand than in mid intertidal medium sand. In the third year, lugworm densities strongly decreased at the study site. Nevertheless the polychaete functional group composition in lugworm exclusion plots still significantly differed from that in control and ambient plots. We assume that the permanent exclusion of lugworms may have entailed a cumulative change in sediment properties in the exclusion plots. Overall, lugworm effects were highly dependent on space and time as well as on differential recruitment success in this intertidal polychaete assemblage. Sediment-mediated effects of an ecosystem engineer on associated species appear to be subtle and contingent in variable environments.

Sequestration and turnover of bacterial‐ and fungal‐derived carbon in a temperate grassland soil under long‐term elevated atmospheric pCO2

AbstractTemperate grasslands contribute about 20% to the global C budget. Elevation of atmospheric CO2 concentration (pCO2) could lead to additional C sequestration into these ecosystems. Microbial‐derived C in the soil comprising about 1–5% of total soil organic carbon may be an important ‘pool’ for long‐term storage of C under future increased atmospheric CO2 concentrations. In our study, the impact of elevated pCO2 on bacterial‐ and fungal‐derived C in the soil of Lolium perenne pastures was investigated under free air carbon dioxide enrichment (FACE) conditions. For 7 years, L. perenne swards were exposed to ambient and elevated pCO2 (36 and 60 Pa pCO2, respectively). The additional CO2 in the FACE plots was depleted in 13C compared with ambient plots, so that ‘new’ (<7 years) C inputs in the form of microbial‐derived residues could be determined by means of stable C isotope analysis. Amino sugars in soil are reliable organic biomarkers for indicating the presence of microbial‐derived residues, with particular amino sugars indicative of either bacterial or fungal origin. It is assumed that amino sugars are stabilized to a significant extent in soil, and so may play an important role in long‐term C storage. In our study, we were also able to discriminate between ‘old’ (> 7 years) and ‘new’ microbial‐derived C using compound‐specific δ13C analysis of individual amino sugars. This new tool was very useful in investigating the potential for C storage in microbial‐derived residues and the turnover of this C in soil under increased atmospheric pCO2.The 13C signature of individual amino sugars varied between −17.4‰ and −39.6‰, and was up to 11.5% depleted in 13C in the FACE plots when compared with the bulk δ13C value of the native C3 L. perenne soil. New amino sugars in the bulk soil contributed up to 16% to the overall amino sugar pool after the first year and between 62% and 125% after 7 years of exposure to elevated pCO2. Amounts of new glucosamine increased by the greatest amount (16–125%) during the experiment, followed by mannosamine (−9% to 107%), muramic acid (−11% to 97%), and galactosamine (15–62%). Proportions of new amino sugars in particle size fractions varied between 38% for muramic acid in the clay fraction and 100% for glucosamine and galactosamine in the coarse sand fraction.Summarizing, during the 7‐year period, amino sugars constituted only between 0.9% and 1.6% of the total SOC content. Therefore, their absolute significance for long‐term C sequestration is limited. Additionally new amino sugars were only sequestered in the silt fraction upon elevated pCO2 exposure while amino sugar concentrations in the clay fraction decreased. Overall, amino sugar concentrations in bulk soil did not change significantly upon exposure to elevated pCO2. The calculated mean residence time of amino sugars was surprisingly low varying between 6 and 90 years in the bulk soil, and between 3 and 30 years in the particle size fractions, representing soil organic matter pools with different but relatively low turnover times. Therefore, compound‐specific δ13C analysis of individual amino sugars clearly revealed a high amino sugar turnover despite more or less constant amino sugar concentrations over a 7 years period of exposure to elevated pCO2.

Effects of Elevated Atmospheric Carbon Dioxide on Biomass and Carbon Accumulation in a Model Regenerating Longleaf Pine Community

Plant species vary in response to atmospheric CO2 concentration due to differences in physiology, morphology, phenology, and symbiotic relationships. These differences make it very difficult to predict how plant communities will respond to elevated CO2. Such information is critical to furthering our understanding of community and ecosystem responses to global climate change. To determine how a simple plant community might respond to elevated CO2, a model regenerating longleaf pine community composed of five species was exposed to two CO2 regimes (ambient, 365 micromol mol(-1) and elevated, 720 micromol mol(-1)) for 3 yr. Total above- and belowground biomass was 70 and 49% greater, respectively, in CO2-enriched plots. Carbon (C) content followed a response pattern similar to biomass, resulting in a significant increase of 13.8 Mg C ha(-1) under elevated CO2. Responses of individual species, however, varied. Longleaf pine (Pinus palustris Mill.) was primarily responsible for the positive response to CO2 enrichment. Wiregrass (Aristida stricta Michx.), rattlebox (Crotalaria rotundifolia Walt. Ex Gmel.), and butterfly weed (Asclepias tuberosa L.) exhibited negative above- and belowground biomass responses to elevated CO2, while sand post oak (Quercus margaretta Ashe) did not differ significantly between CO2 treatments. As with pine, C content followed patterns similar to biomass. Elevated CO2 resulted in alterations in community structure. Longleaf pine comprised 88% of total biomass in CO2-enriched plots, but only 76% in ambient plots. In contrast, wiregrass, rattlebox, and butterfly weed comprised 19% in ambient CO2 plots, but only 8% under high CO2. Therefore, while longleaf pine may perform well in a high CO2 world, other members of this community may not compete as well, which could alter community function. Effects of elevated CO2 on plant communities are complex, dynamic, and difficult to predict, clearly demonstrating the need for more research in this important area of global change science.

Carrion fly (Diptera: Calliphoridae) larval colonization of sunlit and shaded pig carcasses in West Virginia, USA

Two pig ( Sus scrofta L.) carcasses were placed in sunlit and shaded plots in September 2003, and again in May 2004. Mean ambient temperatures between sunlit and shaded plots were not significantly different in either September or May, but mean ambient temperatures at sunlit and shaded plots in 2004 were significantly higher than corresponding means for sunlit and shaded plots in 2003. Mean maggot mass temperatures were significantly higher than ambient plot temperatures for all four experimental plots (i.e., sunlit and shaded carcasses in both 2003 and 2004). In addition, maggot mass temperatures on sunlit carcasses were positively, and significantly, correlated with ambient temperatures, whereas there was no significant correlation between maggot mass and ambient temperatures at shaded plots. Carcass decomposition proceeded more rapidly in 2004 in the presence of higher ambient temperatures, and sunlit carcasses decomposed faster than shaded ones in both 2003 and 2004 experiments. Phaenecia coeruleiviridis (Macquart) and Phormia regina (Meigen) third instars dominated collections on all four carcasses, but there was little temporal overlap between these species with third instars of the former dominating collections in the early portion (≈40%) of each experimental period (with the exception of the shaded carcass in 2004 where both species were co-dominant), and the latter assuming dominance in the latter portion (≈60%). Lower accumulated degree hour values were calculated for instar development on 2004 carcasses subjected to higher ambient temperatures.

Vascular Plant Responses to Elevated CO2 in a Temperate Lowland Sphagnum Peatland

Vascular plant responses to experimental enrichment with atmospheric carbon dioxide (CO2), using MINIFACE technology, were studied in a Dutch lowland peatland dominated by Sphagnum and Phragmites for 3 years. We hypothesized that vascular plant carbon would accumulate in this peatland in response to CO2 enrichment owing to increased productivity of the predominant species and poorer quality (higher C/N ratios) and consequently lower decomposability of the leaf litter of these species. Carbon isotope signatures demonstrated that the extra 180 ppmv CO2 in enriched plots had been incorporated into vegetation biomass accordingly. However, on the CO2 sequestration side of the ecosystem carbon budget, there were neither any significant responses of total aboveground abundance of vascular plants, nor of any of the individual species. On the CO2 release side of the carbon budget (decomposition pathway), litter quantity did not differ between ambient and CO2 treatments, while the changes in litter quality (N and P concentration, C/N and C/P ratio) were marginal and inconsistent. It appeared therefore that the afterlife effects of significant CO2-induced changes in green-leaf chemistry (lower N and P concentrations, higher C/N and C/P) were partly offset by greater resorption of mobile carbohydrates from green leaves during senescence in CO2-enriched plants. The decomposability of leaf litters of three predominant species from ambient and CO2-enriched plots, as measured in a laboratory litter respiration assay, showed no differences. The relatively short time period, environmental spatial heterogeneity and small plot sizes might explain part of the lack of CO2 response. When our results are combined with those from other Sphagnum peatland studies, the common pattern emerges that the vascular vegetation in these ecosystems is genuinely resistant to CO2-induced change. On decadal time-scales, water management and its effects on peatland hydrology, N deposition from anthropogenic sources and land management regimes that arrest the early successional phase (mowing, tree and shrub removal), may have a greater impact on the vascular plant species composition, carbon balance and functioning of lowland Sphagnum-Phragmites reedlands than increasing CO2 concentrations in the atmosphere.

Sensitivity of canopy transpiration to altered precipitation in an upland oak forest: evidence from a long‐term field manipulation study

AbstractClimate‐induced changes in regional precipitation could have important implications for the carbon, water, and nutrient cycles of forest ecosystems. However, few studies have examined the response of deciduous forests to increases or decreases in precipitation. Therefore, the throughfall displacement experiment (TDE) was established in 1993 near Oak Ridge, Tennessee to examine the sensitivity of an upland oak (Quercus spp.) forest to ambient, wet (+33%), and dry (−33%) precipitation regimes. Sap flux measurements on co‐occurring tree species were scaled using species‐specific estimates of stand sapwood area to derive daily and seasonal rates of canopy transpiration (EC) from 2000 to 2003. With the exception of 2003, which was an extremely wet year, daily EC in the dry plot, and occasionally during extended droughts in the ambient and wet plots, declined as water potential in the upper 0.35 m soil profile approached ‐3.0 MPa. Seasonal patterns of soil water potential and treatment‐specific differences in EC were dependent on precipitation frequency and intensity. Supplemental precipitation added to the wet plot increased seasonal EC on average by 9% (range −1% to 19%), whereas extended periods of drought on the dry plot in 2000, 2001, and 2002 were sufficient to reduce seasonal EC by 26–30% compared with the ambient plot. There was a strong correlation between seasonal EC and the water stress integral, a cumulative index of drought severity and duration. A polynomial fitted to these data indicated that reductions in seasonal EC on the order of 40% were possible given TDE‐imposed reductions in soil water potential. Application of this equation to all years of the TDE (1994–2003) revealed considerable interannual and treatment‐specific variation in canopy transpiration. In general, a 33% removal of throughfall on the dry plot during 1995, 1998, and 2002 resulted in a 23–32% reduction in seasonal EC compared with the ambient plot. While droughts in deciduous forests are often limited in duration and tend to occur late in the growing season, soil water deficits of the magnitude observed in this study have the potential to impact local and regional forest water budgets.

Ecological resistance, seed density and their interactions determine patterns of invasion in a California coastal grassland

Relatively little experimental evidence is available regarding how ecological resistance and propagule density interact in their effects on the establishment of invasive exotic species. We examined the independent and interactive effects of neighbour cover (biotic resistance), winter vs. spring water addition (abiotic resistance) and seed density on the invasion of the European perennial grass Holcus lanatus into a California coastal grassland dominated by exotic annual grasses. We found that decreased competition from resident exotic grasses had no effect. In contrast, increased late-season water availability eroded the abiotic resistance offered by naturally dry conditions, facilitating invasion. Finally, watering treatment and seed density interacted strongly in determining seedling survival: while seedling mortality was close to 100% in ambient and winter water addition plots, survivor numbers increased with seed density in spring-watered plots. Thus, decreased abiotic resistance can amplify the effect of increased propagule density on seedling establishment, thereby increasing the likelihood of invasion.

Extracellular Enzyme Activities and Soil Organic Matter Dynamics for Northern Hardwood Forests receiving Simulated Nitrogen Deposition

Anthropogenic nitrogen enrichment alters decomposition processes that control the flux of carbon (C) and nitrogen (N) from soil organic matter (SOM) pools. To link N-driven changes in SOM to microbial responses, we measured the potential activity of several extracellular enzymes involved in SOM degradation at nine experimental sites located in northern Michigan. Each site has three treatment plots (ambient, +30 and +80 kg N ha 1 y 1 ). Litter and soil samples were collected on five dates over the third growing season of N treatment. Phenol oxidase, peroxidase and cellobiohydrolase activities showed significant responses to N additions. In the Acer saccha- rum-Tilia americana ecosystem, oxidative activity was 38% higher in the litter horizon of high N treatment plots, relative to ambient plots, while oxidative activity in mineral soil showed little change. In the A. saccharum-Quercus rubra and Q. velutina-Q. alba ecosystems, oxidative activities declined in both litter (15 and 23%, respectively) and soil (29 and 38%, respectively) in response to high N treatment while cellobiohydrolase activity increased (6 and 39% for litter, 29 and 18% for soil, respectively). Over 3 years, SOM content in the high N plots has decreased in the Acer-Tilia ecosystem and increased in the two Quercus ecosystems, relative to ambient plots. For all three ecosystems, differences in SOM content in relation to N treatment were directly related (r 2 = 0.92) to an enzyme activity factor that included both oxidative and hydrolytic enzyme responses.

Effect of Free-Air CO2 Enrichment (FACE) on the Community Structure of Aquatic Organisms in Floodwater of a Japanese Paddy Field during the Flooding Period of Rice Cultivation

In the present study, the effect of free-air CO2 enrichment (FACE) on the diversity and the community structure of aquatic organisms in the floodwater of a paddy field in northern Japan was evaluated. The varieties and the number of aquatic organisms ranging from 30 μm to 2 cm were compared four times at intervals of 20 d between the FACE (599 ppm CO2) and ambient air (384 ppm CO2) plots during the flooding period of rice cultivation. Aquatic organisms were classified mainly at the order level. Twenty-eight taxonomical groups were detected, and the number of groups in the FACE plots was significantly lower than that in the ambient plots. Zygnematales and Chaetonotida were less frequently detected in the FACE plots. The population densities of Zygnematales, and Chaetonotida also tended to be lower in the FACE plots than in the ambient plots in the late flooding period (after 50 d of flooding). No groups showed a significantly higher population density and frequency of presence in the FACE plots than in the ambient plots. The seasonal variations in the number of taxonomical groups and the community structure of aquatic organisms were less conspicuous in the FACE plots than in the ambient plots. We concluded that the FACE conditions adversely affected the communities of aquatic organisms in paddy fields.

Differential ozone sensitivity in an old and a modern Swedish wheat cultivar—grain yield and quality, leaf chlorophyll and stomatal conductance

Field-grown crops of one modern (‘Dragon’) and one 100-year old (‘Lantvete’) wheat ( Triticum aestivum L.) cultivar were exposed to charcoal-filtered air (CF) or non-filtered air plus additional ozone (NF+) in open-top chambers ( n = 6). Ambient air plots ( n = 3) were used to monitor the effects of the chamber enclosure. Quantitative (ear weight, harvest index and 1000-grain weight), as well as qualitative (grain concentration of N, P, K, Ca, Mg, Cd), aspects of crop yield were studied. In addition, the chlorophyll content of the top leaf was followed during the 10-week-long ozone exposure and flag leaf stomatal conductance was monitored. Ear weight was negatively affected by ozone. The effect was smaller in the lower yielding ‘Lantvete’ compared to the modern cultivar ‘Dragon’. A similar ozone effect was found for the harvest index and 1000-grain weight. The difference between cultivars was small for 1000-grain weight, with harvest index significantly lower in ‘Lantvete’ compared with ‘Dragon’. The crude protein concentration of the grain was higher in ‘Lantvete’, and was enhanced by ozone exposure. No significant treatment or cultivar effects were observed on grain Ca and Cd concentrations, but the greatest chamber effect was on Cd, which was considerably higher in grain from treatment chambers compared with ambient air (possibly due to the higher rate of transpiration in the plants enclosed in the chambers). There were strong differences with respect to grain P, K and Mg concentration between cultivars; ‘Lantvete’ exhibiting higher concentrations than ‘Dragon’ in all cases, apart from K. Weak, but significant, increases in grain P, K and Mg concentrations were evident in O 3-treated plants. The chlorophyll concentration of the flag leaf was negatively affected by ozone exposure after approximately 1 month, but chlorophyll content declined faster in ‘Dragon’ than ‘Lantvete’. Flag leaf chlorophyll concentration was significantly lower in ‘Lantvete’ compared with ‘Dragon’ before onset of ozone effects in NF+ and during most of the experiment in CF. Stomatal conductance was also significantly lower in ‘Lantvete’ than in ‘Dragon’. The most important conclusion drawn from the present study is that the ‘older’ cultivar, ‘Lantvete’, was less affected by ozone than the modern, bred cultivar, ‘Dragon’. This observation might to a large extent be explained by the higher stomatal conductance exhibited by the modern wheat cultivar. The result is consistent with earlier investigations for Greek wheat cultivars, and suggests that the findings reported by others, based on vegetative growth, may extend to grain yield. Growth dilution effects (i.e. the lower observed concentration of an element at higher biomass yield) may explain observed differences between cultivars, and between ozone treatments, in the level of various elements, except for potassium.

Effect of Free-Air CO2 Enrichment (FACE) on Methanogenic Archaeal Communities Inhabiting Rice Roots in a Japanese Rice Field

The effect of free-air CO2 enrichment (FACE) on the methanogenic archaeal communities inhabiting rice roots was studied in a Japanese rice field by separately collecting rice roots three times (at mid-tillering, panicle initiation, and heading stages) according to their nodal number, extracting DNA from the roots and subjecting it to polymerase-chain reaction (PCR)-denaturing gradient gel electrophoresis (DGGE) and sequence analyses. Appearance of roots indicated that aging and senescence occurred faster under the FACE conditions than under the ambient conditions. The number of DGGE bands of methanogenic archaeal communities tended to increase with the growth stages. Cluster analysis showed that the succession of methanogenic archaeal communities in the ambient plot preceded that in the FACE plot, while the trend of the appearance of rice roots was opposite. All the closest relatives associated with the DGGE bands belonged to Methanomicrobiales and Rice cluster I, and FACE did not affect the phylogenetic position of the closest relatives associated with the characteristic DGGE bands. Faster succession of methanogenic archaeal communities in the ambient plot and similar phylogenetic members between the plots were observed in rice roots in years with both warmer (1999) and cooler (2003) weather during the rice cultivation period than in average years.