Cardenolides, induced responses, and interactions between above- and belowground herbivores of milkweed (Asclepias spp.).

Theory has long predicted allocation patterns for plant defense against herbivory, but only recently have both above- and belowground plant defenses been considered simultaneously. Milkweeds in the genus Asclepias are a classic chemically defended clade of plants with toxic cardenolides (cardiac glycosides) and pressurized latex employed as anti-herbivore weapons. Here we combine a comparative approach to investigate broadscale patterns in allocation to root vs. shoot defenses across species with a species-specific experimental approach to identify the consequences of defense allocational shifts on a specialist herbivore. Our results show phylogenetic conservatism for inducibility of shoot cardenolides by an aboveground herbivore, with only four closely related tropical species showing significant induction; the eight temperate species examined were not inducible. Allocation to root and shoot cardenolides was positively correlated across species, and this relationship was maintained after accounting for phylogenetic nonindependence. In contrast to long-standing theoretical predictions, we found no evidence for a trade-off between constitutive and induced cardenolides; indeed the two were positively correlated across species in both roots and shoots. Finally, specialist root and shoot herbivores of common milkweed (A. syriaca) had opposing effects on latex production, and these effects had consequences for caterpillar growth consistent with latex providing resistance. Although cardenolides were not affected by our treatments, A. syriaca allocated 40% more cardenolides to shoots over roots. We conclude that constitutive and inducible defenses are not trading off across plant species, and shoots of Asclepias are more inducible than roots. Phylogenetic conservatism cannot explain the observed patterns of cardenolide levels across species, but inducibility per se was conserved in a tropical clade. Finally, given that above- and belowground herbivores can systemically alter the defensive phenotype of plants, we concur with recent calls for a whole-plant perspective in testing models of plant defense allocation.

ABOVEGROUND AND BELOWGROUND BIOMASS AND CARBON ESTIMATES FOR CLONAL EUCALYPTUS TREES IN SOUTHEAST BRAZIL

Eucalyptus plantations represent a short term and cost efficient alternative for sequestrating carbon dioxide from the atmosphere. Despite the known potential of forest plantations of fast growing species to store carbon in the biomass, there are relatively few studies including precise estimates of the amount of carbon in these plantations. In this study it was determined the carbon content in the stems, branches, leaves and roots of a clonal Eucalyptus grandis plantation in the Southeast of Brazil. We developed allometric equations to estimate the total amount of carbon and total biomass, and produced an estimate of the carbon stock in the stand level. Altogether, 23 sample trees were selected for aboveground biomass assessment. The roots of 9 of the 23 sampled trees were partially excavated to assess the belowground biomass at a singletree level. Two models with DBH, H and DBH2H were tested. The average relative share of carbon content in the stem, branch, leaf and root compartments was 44.6%, 43.0%, 46.1% and 37.8%, respectively, which is smaller than the generic value commonly used (50%). The best-fit allometric equations to estimate the total amount of carbon and total biomass had DBH2H as independent variable. The root-to-shoot ratio was relatively stable (C.V. = 27.5%) probably because the sub-sample was composed of clones. Total stand carbon stock in the Eucalyptus plantation was estimated to be 73.38 MgC ha-1, which is within the carbon stock range for Eucalyptus plantations.

Belowground biomass functions and expansion factors in high elevation Norway spruce

Biomass allocation to above- and belowground compartments in trees is thought to be affected by growth conditions. To assess the strength of such influences, we sampled six Norway spruce forest stands growing at higher altitudes. Within these stands, we randomly selected a total of 77 Norway spruce trees and measured volume and biomass of stem, above- and belowground stump and all roots over 0.5 cm diameter. A comparison of our observations with models parameterised for lower altitudes shows that models developed for specific conditions may be applicable to other locations. Using our observations, we developed biomass functions (BF) and biomass conversion and expansion factors (BCEF) linking belowground biomass to stem parameters. While both BF and BCEF are accurate in belowground biomass predictions, using BCEF appears more promising as such factors can be readily used with existing forest inventory data to obtain estimates of belowground biomass stock. As an example, we show how BF and BCEF developed for individual trees can be used to estimate belowground biomass at the stand level. In combination with existing aboveground models, our observations can be used to quantify total standing biomass of high altitude Norway spruce stands.

(Table 1) Biomass and total standing crop of major wet sedge tundra species at five sites at Alexandra Fiord in 1981 and 2005

The global climate is changing rapidly and Arctic regions are showing responses to recent warming. Responses of tundra ecosystems to climate change have been examined primarily through short-term experimental manipulations, with few studies of long-term ambient change. We investigated changes in above- and belowground biomass of wet sedge tundra to the warming climate of the Canadian High Arctic over the past 25 years. Aboveground standing crop was harvested from five sedge meadow sites and belowground biomass was sampled from one of the sites in the early 1980s and in 2005 using the same methods. Aboveground biomass was on average 158% greater in 2005 than in the early 1980s. The belowground biomass was also much greater in 2005: root biomass increased by 67% and rhizome biomass by 139% since the early 1980s. Dominant species from each functional group (graminoids, shrubs and forbs) showed significant increases in aboveground biomass. Responsive species included the dominant sedge species Carex aquatilis stans, C. membranacea, and Eriophorum angustifolium, as well as the dwarf shrub Salix arctica and the forb Polygonum viviparum. However, diversity measures were not different between the sample years. The greater biomass correlated strongly with increased annual and summer temperatures over the same time period, and was significantly greater than the annual variation in biomass measured in 1980-1983. Increased decomposition and mineralization rates, stimulated by warmer soils, were likely a major cause of the elevated productivity, as no differences in the mass of litter were found between sample periods. Our results are corroborated by published short-term experimental studies, conducted in other wet sedge tundra communities which link warming and fertilization with elevated decomposition, mineralization and tundra productivity. We believe that this is the first study to show responses in High Arctic wet sedge tundra to recent climate change.

Allocation of Biomass and Net Primary Productivity of Mangrove Forests along Environmental Gradients in the Florida Coastal Everglades, USA

Vegetation patterns of mangroves in the Florida Coastal Everglades (FCE) result from the interaction of environmental gradients and natural disturbances (i.e., hurricanes), creating an array of distinct riverine and scrub mangroves across the landscape. We investigated how landscape patterns of biomass and total net primary productivity (NPPT), including allocation in above- and below-ground mangrove components, vary inter-annually (2001–2004) across gradients in soil properties and hydroperiod in two distinct FCE basins: Shark River Estuary and Taylor River Slough. We propose that the allocation of belowground biomass and productivity (NPPB) relative to aboveground allocation is greater in regions with P limitation and permanent flooding. Porewater sulfide was significantly higher in Taylor River (1.2 ± 0.3 mM) compared to Shark River (0.1 ± 0.03 mM) indicating the lack of a tidal signature and more permanent flooding in this basin. There was a decrease in soil P density and corresponding increase in soil N:P from the mouth (28) to upstream locations (46–105) in Shark River that was consistent with previous results in this region. Taylor River sites showed the highest P limitation (soil N:P > 60). Average NPPT was double in higher P environments (17.0 ± 1.1 Mg ha−1 yr−1) compared to lower P regions (8.3 ± 0.3 Mg ha−1 yr−1). Root biomass to aboveground wood biomass (BGB:AWB) ratio was 17 times higher in P-limited environments demonstrating the allocation strategies of mangroves under resource limitation. Riverine mangroves allocated most of the NPPT to aboveground (69%) while scrub mangroves showed the highest allocation to belowground (58%). The total production to biomass (P:B) ratios were lower in Shark River sites (0.11 yr−1); whereas in Taylor River sites P:B ratios were higher and more variable (0.13–0.24 yr−1). Our results suggest that the interaction of lower P availability in Taylor River relative to Shark River basin, along with higher sulfide and permanent flooding account for higher allocation of belowground biomass and production, at expenses of aboveground growth and wood biomass. These distinct patterns of carbon partitioning between riverine and scrub mangroves in response to environmental stress support our hypothesis that belowground allocation is a significant contribution to soil carbon storage in forested wetlands across FCE, particularly in P-limited scrub mangroves. Elucidating these biomass strategies will improve analysis of carbon budgets (storage and production) in neotropical mangroves and understanding what conditions lead to net carbon sinks in the tropical coastal zone.

Functional diversity of macrophyte communities within and between Pyrenean lakes

Submersed vegetation is a common feature in about 70% Pyrene an high mountain (>1500 m a.s.l.) lakes. Isoetids and soft-water elodeids are common elements of this underw ater flora and can form distinct vegetation units (i.e. patches of vegetation dominated by different species) within complex mosaics of vegetation in shallow waters (<7 m). Since is oetids exert a strong influence on sediment biogeochemistry due to high radial oxygen loss, we examined the small scale characteristics of the lake environment (water and sediment) associated to vegetation patches in order to ascertain potential functional differences among them. To do so, we characterised the species composition and biomass of the main vegetation units from 11 lakes, defined plant communities based on biomass data, and then related each community with sediment properties (redox and dissolved nutrient concentration in the pore water) and water nutrient concentration within plant canopy. We also characterised lake water and sediment in areas without vegetation as a reference. A total of twenty-one vegetation units were identified, ranging from one to five per lake. A cluster analysis on biomass species composition suggested seven different macrophyte communities that were named after the most dominant species:Nitella sp.,Potamogeton praelongus, Myriophyllum alterniflorum, Sparganium angustifolium , Isoetes echinospora,Isoetes lacustris and Carex rostrata . Coupling between macrophyte communities and their immediate environment (overlying water and sediment) was manifested mainly as variation in sediment redox conditions and the dominant form of inorganic nitrogen in pore-water. These effects depended on the specific compositi on of the community, and on the allocation between above- and belowground biomass, and could be predicted with a model relating the average and standard deviation of sediment redox potential from 0 down to -20 cm, across macrophyte communities. Differences in pore-water total dissolved phosphorus were related to the trophic state of the lakes. There was no correlation between sediment and water column dissolved nutrients. However, nitrate concentrations tended to be lower in the water overlaying isoetid communities, in apparent contradiction to the patterns of dissolved nitrates in the pore-water. These tendencies were robust even when comparing the water over laying communities within the same lake, thus pointing towards a potential effect of isoetids in reducing dissolved nitrogen in the lakes.

Predicted soil organic carbon stocks and changes in the Brazilian Amazon between 2000 and 2030

Currently we have little understanding of the impacts of land use change on soil C stocks in the Brazilian Amazon. Such information is needed to determine impacts'6n the global C cycle and the sustainability of agricultural systems that are replacing native forest. The aim of this study was to predict soil carbon stocks and changes in the Brazilian Amazon during the period between 2000 and 2030, using the GEFSOC soil carbon (C) modelling system. In order to do so, we devised current and future land use scenarios for the Brazilian Amazon, taking into account: (i) deforestation, rates from the past three decades, (ii) census data on land use from 1940 to 2000, including the expansion and intensification of agriculture in the region, (iii) available information on management practices, primarily related to well managed pasture versus degraded pasture and conventional systems versus no-tillage systems for soybean (Glycine max) and (iv) FAO predictions on agricultural land use and land use changes for the years 2015 and 2030. The land use scenarios were integrated with spatially explicit soils data (SOTER database), climate, potential natural vegetation and land management units using the recently developed GEFSOC soil C modelling system. Results are presented in map, table and graph form for the entire Brazilian Amazon for the current situation (1990 and 2000) and the future (2015 and 2030). Results include soil organic C (SOC) stocks and SOC stock change rates estimated by three methods: (i) the Century ecosystem model, (ii) the Rothamsted C model and (iii) the intergovernmental panel on climate change (IPCC) method for assessing soil C at regional scale. In addition, we show estimated values of above and belowground biomass for native vegetation, pasture and soybean. The results on regional SOC stocks compare reasonably well with those based on mapping approaches. The GEFSOC system provided a means of efficiently handling complex interactions among biotic-edapho-climatic conditions (> 363,000 combinations) in a very large area (similar to 500 Mha) such as the Brazilian Amazon. All of the methods used showed a decline in SOC stock for the period studied; Century and RothC simulated values for 2030 being about 7% lower than those in 1990. Values from Century and RothC (30,430 and 25,000 Tg for the 0-20 cm layer for the Brazilian Amazon region were higher than those obtained from the IPCC system (23,400 Tg in the 0-30 cm layer). Finally; our results can help understand the major biogeochemical cycles that influence soil fertility and help devise management strategies that enhance the sustainability of these areas and thus slow further deforestation. (C) 2007 Elsevier B.V. All rights reserved.

No shift to a deeper water uptake depth in response to summer drought of two lowland and sub-alpine C3-grasslands in Switzerland

Temperate C3-grasslands are of high agricultural and ecological importance in Central Europe. Plant growth and consequently grassland yields depend strongly on water supply during the growing season, which is projected to change in the future. We therefore investigated the effect of summer drought on the water uptake of an intensively managed lowland and an extensively managed sub-alpine grassland in Switzerland. Summer drought was simulated by using transparent shelters. Standing above- and belowground biomass was sampled during three growing seasons. Soil and plant xylem waters were analyzed for oxygen (and hydrogen) stable isotope ratios, and the depths of plant water uptake were estimated by two different approaches: (1) linear interpolation method and (2) Bayesian calibrated mixing model. Relative to the control, aboveground biomass was reduced under drought conditions. In contrast to our expectations, lowland grassland plants subjected to summer drought were more likely (43–68 %) to rely on water in the topsoil (0–10 cm), whereas control plants relied less on the topsoil (4–37 %) and shifted to deeper soil layers (20–35 cm) during the drought period (29–48 %). Sub-alpine grassland plants did not differ significantly in uptake depth between drought and control plots during the drought period. Both approaches yielded similar results and showed that the drought treatment in the two grasslands did not induce a shift to deeper uptake depths, but rather continued or shifted water uptake to even more shallower soil depths. These findings illustrate the importance of shallow soil depths for plant performance under drought conditions.

The contribution of two common shrub species to aboveground and belowgraound carbon pool in Iberian Dehesas

Shrubs play an important role in water-limited agro-silvo-pastoral systems by providing shelter and forage for livestock, for erosion control, to maintain biodiversity, diversifying the landscape, and above all, facilitating the regeneration of trees. Furthermore, the carbon sink capacity of shrubs could also help to mitigate the effects of climate change since they constitute a high proportion of total plant biomass. The contribution of two common extensive native shrub species (Cistus ladanifer L. and Retama sphaerocarpa (L.) Boiss.) to the carbon pool of Iberian dehesas (Mediterranean agro-silvo-pastoral systems) is analyzed through biomass models developed at both individual (biovolume depending) and community level (height and cover depending). The total amount of carbon stored in these shrubs, including above- and belowground biomass, ranges from 1.8 to 11.2 Mg C ha_1 (mean 6.8 Mg C ha_1) for communities of C. ladanifer and from 2.6 to 8.6 Mg C ha_1 (mean 4.5 Mg C ha_1) for R. sphaerocarpa. These quantities account for over 20e30% of the total plant biomass in the system. The potential for carbon sequestration of these shrubs in the studied system ranges 0.10e1.32 Mg C ha_1 year_1 and 0.25e1.25 Mg C ha_1 year_1 for the C. ladanifer and R. sphaerocarpa communities’ respectively

The role of soil in vegetated gravelly river braid plains: more than just a passive response?

This paper reviews the role of alluvial soils in vegetated gravelly river braid plains. When considering decadal time scales of river evolution, we argue that it becomes vital to consider soil development as an emergent property of the developing ecosystem. Soil processes have been relatively overlooked in accounts of the interactions between braided river processes and vegetation, although soils have been observed on vegetated fluvial landforms. We hypothesise that soil development plays a major role in the transition (speed and pathway) from a fresh sediment deposit to a vegetated soil-covered landform. Disturbance (erosion and/or deposition), vertical sediment structure (process history), vegetation succession, biological activity and water table fluctuation are seen as the main controls on early alluvial soil evolution. Erosion and deposition processes may not only act as soil disturbing agents, but also as suppliers of ecosystem resources, because of their role in delivering and changing access (e.g. through avulsion) to fluxes of water, fine sediments and organic matter. In turn, the associated initial ecosystem may influence further fluvial landform development, such as through the trapping of fine-grained sediments (e.g. sand) by the engineering action of vegetation and the deposit stabilisation by the developing above and belowground biomass. This may create a strong feedback between geomorphological processes, vegetation succession and soil evolution which we summarise in a conceptual model. We illustrate this model by an example from the Allondon River (CH) and identify the research questions that follow.

Effet de l’irrigation par l’eau usée sur la biomasse aérienne et souterraine d’une culture intensive de saules en courtes rotations

Le but de cette étude est de comprendre l’effet d’une irrigation par les eaux usées et /ou de la fertilisation par les engrais chimiques sur la productivité aérienne et souterraine d’une plantation de saule Salix miyabeana SX67 en CICR dans un contexte de filtre végétal. Nous avons d’une part évalué l’impact de diverses doses d’eau usées et/ou de la fertilisation minérale sur les rendements en biomasse ligneuse d’une culture de saules au cours d’un cycle de croissance de deux ans. D’autre part et pour la même période nous avons comparé le développement racinaire (biomasse, morphologie et distribution dans le sol) suite aux divers traitements. Les résultats ont montré qu’au terme de deux ans de croissance, les traitements par les eaux usées aussi bien que celle par les engrais a permis l’augmentation des rendements de la biomasse aérienne de notre culture de saules avec un effet plus prononcé suite au traitements des eaux usées qu’à celui du fertilisant chimique. Nous avons mesuré des productivités en biomasse aussi élevées que 39,4 Mg ha-1 et 54,7 Mg ha-1 et ce pour les parcelles qui ont reçu la plus grande quantité d'eaux usées, respectivement pour les saules non fertilisé et fertilisé (D3-NF et D3-F). La majeure partie du système racinaire était en superficie avec 92-96% des racines (racine fine et racine grosse) concentrées dans les premiers 40 cm de sol et nous avons trouvé que la biomasse des racines fines était comprise entre 1,01 et 1,99 Mg ha-1. Généralement la fertilisation chimique n’a pas eu d’effet sur les rendements en biomasse des racines totales et/ou fines. Bien que l’irrigation par les eaux usées ait entraîné une réduction statistiquement significative de la biomasse racinaire, néanmoins cette réduction n'était pas linéaire (avec une réduction de la biomasse de D0 à D1, une augmentation de D1 à D2 pour réduire de nouveau de D2 à D3). Cette tendance porte à penser qu'au-delà d'une certaine quantité d'eau et de nutriments (suite à l’irrigation par les eaux usées), le développement du système racinaire des saules est affecté négativement, et bien que la biomasse aérienne soit restée élevée sous le traitement D3, nous pensons que le développement de la plante a été quelque peu déséquilibré. Aucun changement significatif n'a été constaté dans les traits morphologiques liés à l'irrigation par les eaux usées.

Soil organic carbon dynamics under perennial energy crops

The European renewable energy directive 2009/28/EC (E.C. 2009) provides a legislative framework for reducing GHG emissions by 20%, while achieving a 20% share of energy from renewable sources by 2020. Perennial energy crops could significantly contribute to limit GHG emissions through replacing equivalent fossil fuels and by sequestering a considerable amount of carbon into the soil through the large amounts of belowground biomass produced. The objective of this study is to evaluate the effects of land use change that perennial energy crops have on croplands (switchgrass) and marginal grasslands (miscanthus). For that purpose above and belowground biomass, SOC variation and Net Ecosystem Exchange were evaluated after five years of growth. At aboveground level both crops produced high biomass under cropland conditions as well as under marginal soils. At belowground level they also produced large amounts of biomass, but no significant influences on SOC in the upper layer (0-30 cm) were found. This is probably because of the "priming effect" that caused fast carbon substitution. In switchgrass only it was found a significant SOC increase in deeper layers (30-60 cm), while in the whole soil profile (0-60 cm) SOC increased from 42 to 51 ha-1. However, the short experimental periods (for both switchgrass and miscanthus), in which land use change was evaluated, do not permit to determine the real capacity of perennial energy crops to accumulate SOC. In conclusion the large amounts of belowground biomass enhanced the SOC dynamic through the priming effect resulting in increased SOC in cropland but not in marginal grassland.

Assessing visual disturbance conditions on the Custer National Forest

Assessment of soil disturbance on the Custer National Forest was conducted during two summers to determine if the U.S. Forest Service Forest Soil Disturbance Monitoring Protocol (FSDMP) was able to distinguish post-harvest soil conditions in a chronological sequence of sites harvested using different ground-based logging systems. Results from the first year of sampling suggested that the FSDMP point sampling method may not be sensitive enough to measure post-harvest disturbance in stands with low levels of disturbance. Therefore, a revised random transect method was used during the second sampling season to determine the actual extent of soil disturbance in these cutting units. Using combined data collected from both summers I detected statistically significant differences (p < 0.05) in fine fraction bulk density measurements between FSDMP disturbance classes across all sites. Disturbance class 3 (most severe) had the highest reported bulk density, which suggest that the FSDMP visual class estimates are defined adequately allowing for correlations to be made between visual disturbance and actual soil physical characteristics. Forest site productivity can be defined by its ability to retain carbon and convert it to above- and belowground biomass. However, forest management activities that alter basic site characteristics have the potential to alter productivity. Soil compaction is one critical management impact that is important to understand; compaction has been shown to impede the root growth potential of plants, reduce water infiltration rates increasing erosion potential, and alter plant available water and nutrients, depending on soil texture. A new method to assess ground cover, erosion, and other soil disturbances was recently published by the U.S. Forest Service, as the Forest Soil Disturbance Protocol (FSDMP). The FSDMP allows soil scientists to visually assign a disturbance class estimate (0 – none, 1, 2, 3 – severe) from field measures of consistently defined soil disturbance indicators (erosion, fire, rutting, compaction, and platy/massive/puddled structure) in small circular (15 cm) plots to compare soil quality properties pre- and post- harvest condition. Using this protocol we were able to determine that ground-based timber harvesting activities occurring on the Custer National Forest are not reaching the 15% maximum threshold for detrimental soil disturbance outlined by the Region 1 Soil Quality Standards.

The effects of carbon dioxide fertilization on the ecology of tropical seagrass communities

Increasing atmospheric CO2 concentrations associated with climate change will likely influence a wide variety of ecosystems. Terrestrial research has examined the effects of increasing CO2 concentrations on the functionality of plant systems; with studies ranging in scale from the short-term responses of individual leaves, to long-term ecological responses of complete forests. While terrestrial plants have received much attention, studies on the responses of marine plants (seagrasses) to increased CO 2(aq) concentrations remain relatively sparse, with most research limited to small-scale, ex situ experimentation. Furthermore, few studies have attempted to address similarities between terrestrial and seagrass responses to increases in CO2(aq). The goals of this dissertation are to expand the scope of marine climate change research, and examine how the tropical seagrass, Thalassia testudinum responds to increasing CO 2(aq)concentrations over multiple spatial and temporal scales. ^ Manipulative laboratory and field experimentation reveal that, similar to terrestrial plants, seagrasses strongly respond to increases in CO 2(aq) concentrations. Using a novel field technique, in situ field manipulations show that over short time scales, seagrasses respond to elevated CO2(aq) by increasing leaf photosynthetic rates and the production of soluble carbohydrates. Declines in leaf nutrient (nitrogen and phosphorus) content were additionally detected, paralleling responses from terrestrial systems. Over long time scales, seagrasses increase total above- and belowground biomass with elevated CO2(aq), suggesting that, similar to terrestrial research, pervasive increases in atmospheric and oceanic CO2(aq) concentrations stand to influence the productivity and functionality of these systems. Furthermore, field experiments reveal that seagrass epiphytes, which comprise an important component of seagrass ecosystems, additionally respond to increased CO2(aq) with strong declines in calcified taxa and increases in fleshy taxa. ^ Together, this work demonstrates that increasing CO2(aq) concentrations will alter the functionality of seagrass ecosystems by increasing plant productivity and shifting the composition of the epiphyte community. These results have implications for future rates of carbon storage and sediment production within these widely distributed systems.^