The variable effects of soil nitrogen availability and insect herbivory on aboveground and belowground plant biomass in an old-field ecosystem

Nutrient availability and herbivory can regulate primary production in ecosystems, but little is known about how, or whether, they may interact with one another. Here, we investigate how nitrogen availability and insect herbivory interact to alter aboveground and belowground plant community biomass in an old-field ecosystem. In 2004, we established 36 experimental plots in which we manipulated soil nitrogen (N) availability and insect abundance in a completely randomized plot design. In 2009, after 6 years of treatments, we measured aboveground biomass and assessed root production at peak growth. Overall, we found a significant effect of reduced soil N availability on aboveground biomass and belowground plant biomass production. Specifically, responses of aboveground and belowground community biomass to nutrients were driven by reductions in soil N, but not additions, indicating that soil N may not be limiting primary production in this ecosystem. Insects reduced the aboveground biomass of subdominant plant species and decreased coarse root production. We found no statistical interactions between N availability and insect herbivory for any response variable. Overall, the results of 6 years of nutrient manipulations and insect removals suggest strong bottom-up influences on total plant community productivity but more subtle effects of insect herbivores on aspects of aboveground and belowground production.     

Drought effects on above- and belowground production of a grazed temperate grassland ecosystem

The effect of climatic variation on terrestrial aboveground productivity (ANPP) has been well studied. However, little is known about how variable climate, including drought, may influence belowground productivity (BNPP), which constitutes most of the annual primary production of grasslands. The objectives of this study were to (1) examine how a 3-year period of declining moisture, which began as climatically wet to average across Yellowstone National Park (YNP) and ended in drought, affected ANPP and BNPP in grasslands of YNP and (2) how herds of grazing ungulates, which were shown previously to stimulate grassland shoot and root growth in YNP, may have interacted with climatic conditions to influence grassland production. ANPP and 0–20 cm BNPP, representing the bulk of the root dynamics, were measured in grazed and ungrazed (fenced) grassland at nine sites ranging widely in elevation, soil conditions and plant production during the 3-year study. Results revealed that 0–20 cm BNPP was strongly influenced by drought (P = 0.0005) and declined from 1999 to 2001 among ungrazed and grazed grasslands by 39 and 49%, respectively. The greater reduction in 0–20 cm BNPP among grazed grasslands was due, in part, to a decline (P = 0.07) in the stimulatory effect of grazing, i.e., the ratio g BNPP stimulated: g shoot consumed. In contrast, ANPP was unaffected by drought in either type of grassland. Thus, the effect of this drought in YNP was a large reduction in BNPP, which was a function of (1) a direct negative influence of increased moisture stress on root growth and (2) a weak interaction between drought and herbivory that led to a decline in the positive feedback from grazers to BNPP. These findings highlight the need to better understand factors that control root growth and to study the effects of climatic variation on grasslands within an ecosystem framework to include potentially important climate–consumer interactions.     

Contrasting covariation of above- and belowground invertebrate species across plant genotypes

1. Invertebrate species generally do not respond independently to genotypic variation in plants, giving rise to clusters of species that naturally associate with or avoid certain genotypes. This covariation causes coevolution to be diffuse rather than pairwise. Studies on this topic, however, have never considered the belowground invertebrate community, leaving a critical gap in our understanding. 2. We investigated the covariation among naturally colonising above- and belowground invertebrate species across six genetically distinct populations of the dune grass Ammophila arenaria. After having grown from seed in a common garden, plants were randomised in a single field site to exclude all but broad-sense genetic variation. 3. Strong positive covariation across genotypes among both above- and belowground invertebrates was detected, while correlations between these two groups were negative. This clustering of above- and belowground species matched well with order level taxonomy. Host range, trophic level and food type on the other hand did not correspond well with the clusters. Within the cluster of aboveground fauna, subsequent groupings were not related to any phylogenetic or ecological characteristic, although correlations within these subgroups were very high. We furthermore demonstrated significant differences in multiple invertebrate species occurrence between plant genotypes, in general as well as at the above- and belowground level. 4. The observed strong covariation suggests diffuse coevolution between A. arenaria and its associated invertebrate species. The trade-off between root and shoot invertebrates could however hamper directional selection on resistance to either group. 5. Our results clearly demonstrate the need for studies of plant-animal interactions to include the belowground fauna, as this might drastically alter our general conception of how plants and their associated animal communities interact and how these interactions shape the process of evolution.     

The effects of above- and belowground mutualisms on orchid speciation and coexistence

Both pollination by animals and mycorrhizal symbioses with fungi are believed to have been important for the diversification of flowering plants. However, the mechanisms by which these above- and belowground mutualisms affect plant speciation and coexistence remain obscure. We provide evidence that shifts in pollination traits are important for both speciation and coexistence in a diverse group of orchids, whereas shifts in fungal partner are important for coexistence but not for speciation. Phylogenetic analyses show that recently diverged orchid species tend either to use different pollinator species or to place pollen on different body parts of the same species, consistent with the role of pollination-mode shifts in speciation. Field experiments provide support for the hypothesis that colonization of new geographical areas requires adaptation to new pollinator species, whereas co-occurring orchid species share pollinator species by placing pollen on different body parts. In contrast to pollinators, fungal partners are conserved between closely related orchid species, and orchids recruit the same fungal species even when transplanted to different areas. However, co-occurring orchid species tend to use different fungal partners, consistent with their expected role in reducing competition for nutrients. Our results demonstrate that the two dominant mutualisms in terrestrial ecosystems can play major but contrasting roles in plant community assembly and speciation.     

Effects of nitrogen additions on above- and belowground carbon dynamics in two tropical forests

Anthropogenic nitrogen (N) deposition is increasing rapidly in tropical regions, adding N to ecosystems that often have high background N availability. Tropical forests play an important role in the global carbon (C) cycle, yet the effects of N deposition on C cycling in these ecosystems are poorly understood. We used a field N-fertilization experiment in lower and upper elevation tropical rain forests in Puerto Rico to explore the responses of above- and belowground C pools to N addition. As expected, tree stem growth and litterfall productivity did not respond to N fertilization in either of these N-rich forests, indicating a lack of N limitation to net primary productivity (NPP). In contrast, soil C concentrations increased significantly with N fertilization in both forests, leading to larger C stocks in fertilized plots. However, different soil C pools responded to N fertilization differently. Labile (low density) soil C fractions and live fine roots declined with fertilization, while mineral-associated soil C increased in both forests. Decreased soil CO₂ fluxes in fertilized plots were correlated with smaller labile soil C pools in the lower elevation forest (R² = 0.65, p < 0.05), and with lower live fine root biomass in the upper elevation forest (R² = 0.90, p < 0.05). Our results indicate that soil C storage is sensitive to N deposition in tropical forests, even where plant productivity is not N-limited. The mineral-associated soil C pool has the potential to respond relatively quickly to N additions, and can drive increases in bulk soil C stocks in tropical forests.     

Interactions between above- and belowground insect herbivores as mediated by the plant defense system

Plants are frequently attacked by both above- and belowground arthropod herbivores. Nevertheless, studies rarely consider root and shoot herbivory in conjunction. Here we provide evidence that the root-feeding insect Agriotes lineatus reduces the performance of the foliage feeding insect Spodoptera exigua on cotton plants. In a bioassay, S. exigua larvae were allowed to feed on either undamaged plants, or on plants that had previously been exposed to root herbivory, foliar herbivory, or a combination of both. Previous root herbivory reduced the relative growth rates as well as the food consumption of S. exigua by more than 50% in comparison to larvae feeding on the undamaged controls. We found no effects in the opposite direction, as aboveground herbivory by S. exigua did not affect the relative growth rates of root-feeding A. lineatus . Remarkably, neither did the treatment with foliar herbivory affect the food consumption and relative growth rate of S. exigua in the bioassay. However, this treatment did result in a significant change in the distribution of S. exigua feeding. Plants that had been pre-exposed to foliar herbivory suffered significantly less damage on their young terminal leaves. While plant growth and foliar nitrogen levels were not affected by any of the treatments, we did find significant differences between treatments with respect to the level and distribution of plant defensive chemicals (terpenoids). Exposure to root herbivores resulted in an increase in terpenoid levels in both roots as well as in mature and immature foliage. Foliar damage, on the other hand, resulted in high terpenoid levels in young, terminal leaves only. Our results show that root-feeding herbivores may change the level and distribution of plant defenses aboveground. Our data suggest that the reported interactions between below- and aboveground insect herbivores are mediated by induced changes in plant secondary chemistry.     

Plant production and nitrogen accumulation above- and belowground in low and tall birch tundra communities: the influence of snow and litter

Background and Aim

A vegetation transition to taller and denser deciduous shrub tundra is currently occurring in many locations across the low Arctic, and is associated with climate change. Here, we investigated if deeper snow is a mechanism for enhanced shrub growth.

Methods

To determine if a moderate and climatically realistic increase in snow depth can enhance shrub productivity, we compared growth responses between ambient and experimentally deepened snow plots in low birch hummock tundra. To determine the potential influence of factors other than deepened snow that are associated with taller, denser shrubs, we also compared shrub growth between low birch hummock and tall birch-dominated tundra.

Results

Neither deciduous shrub above- nor belowground production nor nitrogen accumulation was enhanced by deepened snow. However, deciduous birch shrub new shoot production was 23× larger and total vascular shoot to belowground biomass ratios were higher in the tall birch tundra than the birch hummock (~0.7 and ~0.4, respectively), indicating that the combination of deeper snow together with other internal feedbacks greatly enhanced birch growth.

Conclusions

Together, our results strongly suggest that the much larger litter production in tall birch ecosystems is an important internal feedback that may or may not interact with deeper snow to promote birch growth in tall shrub tundra.

     

Effects of two contrasting hemiparasitic plant species on biomass production and nitrogen availability

Hemiparasitic plants can substantially change plant community structure; the drainage of host resources has a direct negative effect on host biomass and, as a consequence, promotes non-host biomass production (parasitism pathway); on the other hand, hemiparasitic litter inputs can enhance nutrient cycling which may have an indirect positive effect on both host and non-host biomass production (litter pathway). We evaluated the net effect of both pathways on total shoot biomass (with and without the hemiparasite) and shoot biomass of graminoids, forbs and ericaceous shrubs using a removal experiment in three sites infested with the annual Rhinanthus angustifolius, and three sites infested with the biennial Pedicularis sylvatica. We addressed the potential importance of litter effects by determination of litter quantity and quality, as well as modeling N release during decomposition. In the second year after removing the hemiparasites, total plant biomass at Rhinanthus sites was 24 % higher in weeded plots than in control plots, while weeding had no significant effect at Pedicularis sites. The increase in total biomass following Rhinanthus removal was mainly due to a higher biomass of graminoids. The amount of litter produced by Rhinanthus was only half of that produced by Pedicularis; N contents were similar. The amount of N in the litter was 9 and 30 % of the amount removed by mowing for Rhinanthus and Pedicularis sites, respectively. Within 2 months, about 45 % of the N in both hemiparasitic litter types was released by decomposition. Our results suggest that in addition to the suppression of host biomass due to parasitism, positive litter feedbacks on host and non-host biomass—via an increase in nutrient availability—also affect plant community structure. We propose that, depending on the particular hemiparasite and/or site conditions, these positive litter feedbacks on shoot biomass can compensate for the negative effect of parasitism.     

Integrating nitrogen fixing structures into above- and belowground functional trait spectra in soy (Glycine max)

Plant and Soil - Phenotypic trait variation across environmental gradients and through plant ontogeny is critical in driving ecological processes, especially in agroecosystems where single...     

Earthworm and belowground competition effects on plant productivity in a plant diversity gradient

Diversity is one major factor driving plant productivity in temperate grasslands. Although decomposers like earthworms are known to affect plant productivity, interacting effects of plant diversity and earthworms on plant productivity have been neglected in field studies. We investigated in the field the effects of earthworms on plant productivity, their interaction with plant species and functional group richness, and their effects on belowground plant competition. In the framework of the Jena Experiment we determined plant community productivity (in 2004 and 2007) and performance of two phytometer plant species [Centaurea jacea (herb) and Lolium perenne (grass); in 2007 and 2008] in a plant species (from one to 16) and functional group richness gradient (from one to four). We sampled earthworm subplots and subplots with decreased earthworm density and reduced aboveground competition of phytometer plants by removing the shoot biomass of the resident plant community. Earthworms increased total plant community productivity (+11%), legume shoot biomass (+35%) and shoot biomass of the phytometer C. jacea (+21%). Further, phytometer performance decreased, i.e. belowground competition increased, with increasing plant species and functional group richness. Although single plant functional groups benefited from higher earthworm numbers, the effects did not vary with plant species and functional group richness. The present study indicates that earthworms indeed affect the productivity of semi-natural grasslands irrespective of the diversity of the plant community. Belowground competition increased with increasing plant species diversity. However, belowground competition was modified by earthworms as reflected by increased productivity of the phytometer C. jacea. Moreover, particularly legumes benefited from earthworm presence. Considering also previous studies, we suggest that earthworms and legumes form a loose mutualistic relationship affecting essential ecosystem functions in temperate grasslands, in particular decomposition and plant productivity. Further, earthworms likely alter competitive interactions among plants and the structure of plant communities by beneficially affecting certain plant functional groups.     

The strategy of the wheat plant in acclimating growth and grain production to nitrogen availability

Two cultivars of spring wheat (Triticum aestivum L.) were grown to maturity in hydroponic cultures. Nitrogen accumulation was controlled by daily growth-limiting additions of nitrate together with all other nutrients in excess. Six different curves of N accumulation were used, with the same relative changes from day to day, but with different amplitudes. These curves were obtained by using the same mathematic formula of the N accumulation curves but varying the value of initial N content. The total amount of nitrogen added varied from 20 mg plant(-1) to 65 mg plant(-1). Plant bioproductivity showed a linear response to accumulated N. The number of grains per plant increased linearly with increased N availability whereas grain weights were essentially unaffected. Grain N concentrations and N content varied slightly, with highest values generally at the lower N availability levels. The quantitatively most important response to increased N availability was an increased number of earbearing tillers per plant. This varied from 0.1 tiller plant(-1) at maturity when given 20 mg N plant(-1), up to about 2 tillers plant(-1) when given 65 mg N plant(-1). Not all tillers that were initiated developed ears. The reduction of tillers seems to be one important mechanism in adapting plant productivity to N availability. Other individual characters influenced by N availability were straw height and the number of spikelets per spike. The two cultivars behaved in a qualitatively similar manner over the range of N availability even though they quantitatively differed in grain size, N concentrations and yield.     

Inter-annual variation in above- and belowground herbivory on a native,annual legume

The relative importance of subterranean versus aboveground insect damage to plants is not well understood. In particular, the simultaneous effects of above- and belowground herbivory, and the importance of highly variable abiotic factors such as rainfall, have received little attention in diverse natural ecosystems. We investigated the influence of both above- and belowground herbivory on Lupinus nanus (Fabaceae), an annual plant native to coastal California. A number of insect species damage L. nanus aboveground, and a weevil larva consumes nodules belowground. To manipulate herbivory in the field, we employed a combination of insecticides and simulated herbivory during two different years. In 1997, simulated belowground damage reduced L. nanus survival, and insecticide application to roots increased seed production and seed mass. By contrast, in 1998, only aboveground folivory significantly reduced L. nanus reproduction, and, in combination, above- and belowground insecticides did not affect flower or seed number relative to controls. A growth chamber experiment conducted in the absence of herbivory revealed that the aboveground insecticide marginally reduced flower production and the belowground insecticide marginally increased flower production compared to controls; these non-target effects made our field experiments for aboveground herbivory conservative. Finally, ambient levels of herbivory differed among years (1997, 1998, and 2000), which varied greatly in rainfall due to the effects of El Nino. The results suggest that the impacts of herbivores are temporally variable and that abiotic factors, particularly those related to large-scale changes in weather patterns, may be more important than insect herbivory to L. nanus in some years. This revised version was published online in August 2006 with corrections to the Cover Date.     

Combined effects of soil moisture and nitrogen availability variations on grass productivity in African savannas

Savannas cover about 20% of the Earth’s land area and 50% of Africa. As an indispensable component of savanna, grasses play an important role in these ecosystems. A better understanding of grass productivity and its controlling factors in savanna ecosystems could therefore be a key to understand the functioning of savannas and predict savanna responses to future climatic changes. In this study, a stable isotope fertilization experiment was conducted to determine how factors limiting grass production in savannas differ across regional climate gradients. The study was conducted on the geomorphically homogenous Kalahari Transect (KT), which offers an ideal setting to study nutrient and vegetation dynamics independently of confounding soil effects. The results show that the grasses assimilated the added fertilizer at all the sites but they did not respond to nitrogen fertilization for both dry and wet years, and at both dry and wet ends of the Transect. Although prior studies have proposed a switch between water and nitrogen limitations between arid and mesic savannas, our results suggest that nitrogen availability may not limit grass productivity across the whole KT. Thus, although the traditional classifications as nutrient poor (broad-leaf) and nutrient rich (fine-leaf) savanna ecosystems may still be useful, it does not necessarily imply the existence of nitrogen limitation in the nutrient poor area; in fact, it is more likely that the herbaceous species found in the more humid sites (nutrient poor sites) are already adapted to lower nitrogen availability.     

Herbivore impact on moss depth, soil temperature and arctic plant growth

We provide evidence for a mechanism by which herbivores may influence plant abundance in arctic ecosystems. These systems are commonly dominated by mosses, the thickness of which influences the amount of heat reaching the soil surface. Herbivores can reduce the thickness of the moss layer by means of trampling and consumption. Exclusion of grazing by barnacle geese and reindeer over a period of 7?years at Ny-Ålesund, Spitsbergen, caused an increase in the thickness of the moss layer, and a reduction in soil temperature of 0.9?°C. Soil temperature was negatively correlated with moss-layer thickness across sites, with highest soil temperatures where moss layers were shallow. We found that moss growth did not respond to experimental manipulation of soil temperature, but the grass Poa arctica (arctic meadow-grass) and the dicot Cardamine nymanii (polar cress) suffered a 50% reduction in biomass when growing in chilled soils.     

Responses of soil biota and nitrogen availability to an invasive plant under aboveground herbivory

Plant and Soil - Recently, much attention has been paid to the plant-mediated effects of aboveground herbivory on soil ecosystems. However, studies about the herbivore-induced effects of invasive...     

Initial effects of experimental warming on above- and belowground components of net ecosystem CO2 exchange in arctic tundra

The Arctic contains extensive soil carbon reserves that could provide a substantial positive feedback to atmospheric CO₂ concentrations and global warming. Evaluation of this hypothesis requires a mechanistic understanding of the in situ responses of individual components of tundra net ecosystem CO₂ exchange (NEE) to warming. In this study, we measured NEE, total ecosystem respiration and respiration from below ground in experimentally warmed plots within Alaskan acidic tussock tundra. Soil warming of 2-4°C during a single growing season caused strong increases in total ecosystem respiration and belowground respiration from moss-dominated inter-tussock areas, and similar trends from sedge-dominated tussocks. Consequently, the overall effect of the manipulation was to substantially enhance net ecosystem carbon loss during mid-summer. Components of vascular plant biomass were closely correlated with total ecosystem respiration and belowground respiration in control plots of both microsites, but not in warmed plots. By contrast, in the warmed inter-tussock areas, belowground respiration was most closely correlated with organic-layer depth. Warming in tussock areas was associated with increased leaf nutrient pools, indicating enhanced rates of soil nutrient mineralisation. Together, these results suggest that warming enhanced net ecosystem CO₂ efflux primarily by stimulating decomposition of soil organic matter, rather than by increasing plant-associated respiration. Our short-term experiment provides field evidence to support previous growth chamber and modelling studies indicating that arctic soil C reserves are relatively sensitive to warming and could supply an initial positive feedback to rising atmospheric CO₂ concentrations/changing climate.     

Interactive effects of soil nitrogen and water availability on leaf mass loss in a temperate steppe

Although the link between leaf mass loss and assessment of ecosystem nutrient use efficiency and plant nutrient resorption efficiency has received considerable attention in various ecosystems, there has been relatively little effort to assess plant leaf mass loss during senescence, especially for herbaceous species. We conducted experimental studies to assess leaf mass loss during senescence in five dominant herbaceous species and examined the effects of increasing nitrogen (N) and water availability on leaf mass loss of four species in a temperate steppe in northern China. We nondestructively estimated mature leaf mass based on leaf length and width. Leaf mass loss varied substantially among species, ranging from 20–50%. On average across all species, N and water addition increased leaf mass loss by 30% and 19%, respectively. N and water addition interacted to affect leaf mass loss, as water addition had a significant positive effect on leaf mass loss under enriched N conditions but showed no effect under ambient N levels. We conclude that leaf mass loss of herbaceous plants was considerable and can potentially be more pronounced with increasing N and water availability. It is notable that the responses of plant species to N and water addition were variable. We suggest that leaf mass loss during senescence should be given full consideration in assessing nutrient use and resorption efficiency in semi-arid areas.     

Drought timing differentially affects above- and belowground productivity in a mesic grassland

Climate models forecast an intensification of the global hydrological cycle with droughts becoming more frequent and severe, and shifting to times when they have been historically uncommon. Droughts, or prolonged periods of precipitation deficiency, are characteristic of most temperate grasslands, yet few experiments have explored how variation in the seasonal timing of drought may impact ecosystem function. We investigated the response of above- and belowground net primary production (ANPP & BNPP) to altered drought timing in a mesic grassland in NE Kansas. Moderate drought treatments (25% reduction from the mean growing season precipitation [GSP]) were imposed by erecting rainout shelters in late spring (LSP), early summer (ESM), and mid-summer (MSM, n = 10 plots/treatment). These treatments were compared to two controls (long-term average GSP [LTA] and ambient GSP [AMB]) and a wet treatment (+30% of the long-term average GSP [WET]). We found that LSP drought did not significantly reduce ANPP relative to control plots while the ESM and MSM drought did despite equivalent reductions in soil moisture. In contrast, the WET treatment did not affect ANPP. Neither the WET nor the drought treatments altered BNPP as compared to the controls. Our results suggest that forecasts of ecosystem responses to climate change will be improved if both the seasonal timing of alterations in precipitation as well as differential responses of above- and belowground productivity to drought are incorporated into models.