Species richness alters spatial nutrient heterogeneity effects on above-ground plant biomass.
ABSTRACT: Previous studies have suggested that spatial nutrient heterogeneity promotes plant nutrient capture and growth. However, little is known about how spatial nutrient heterogeneity interacts with key community attributes to affect plant community production. We conducted a meta-analysis to investigate how nitrogen heterogeneity effects vary with species richness and plant density. Effect size was calculated using the natural log of the ratio in plant biomass between heterogeneous and homogeneous conditions. Effect sizes were significantly above zero, reflecting positive effects of spatial nutrient heterogeneity on community production. However, species richness decreased the magnitude of heterogeneity effects on above-ground biomass. The magnitude of heterogeneity effects on below-ground biomass did not vary with species richness. Moreover, we detected no modification in heterogeneity effects with plant density. Our results highlight the importance of species richness for ecosystem function. Asynchrony between above- and below-ground responses to spatial nutrient heterogeneity and species richness could have significant implications for biotic interactions and biogeochemical cycling in the long term.
Project description:Food web composition and resource levels can influence ecosystem properties such as productivity and elemental cycles. In particular, herbivores occupy a central place in food webs as the species richness and composition of this trophic level may simultaneously influence the transmission of resource and predator effects to higher and lower trophic levels, respectively. Yet, these interactions are poorly understood.Using an experimental seagrass mesocosm system, we factorially manipulated water column nutrient concentrations, food chain length, and diversity of crustacean grazers to address two questions: (1) Does food web composition modulate the effects of nutrient enrichment on plant and grazer biomasses and stoichiometry? (2) Do ecosystem fluxes of dissolved oxygen and nutrients more closely reflect above-ground biomass and community structure or sediment processes? Nutrient enrichment and grazer presence generally had strong effects on biomass accumulation, stoichiometry, and ecosystem fluxes, whereas predator effects were weaker or absent. Nutrient enrichment had little effect on producer biomass or net ecosystem production but strongly increased seagrass nutrient content, ecosystem flux rates, and grazer secondary production, suggesting that enhanced production was efficiently transferred from producers to herbivores. Gross ecosystem production (oxygen evolution) correlated positively with above-ground plant biomass, whereas inorganic nutrient fluxes were unrelated to plant or grazer biomasses, suggesting dominance by sediment microbial processes. Finally, grazer richness significantly stabilized ecosystem processes, as predators decreased ecosystem production and respiration only in the zero- and one- species grazer treatments.Overall, our results indicate that consumer presence and species composition strongly influence ecosystem responses to nutrient enrichment, and that increasing herbivore diversity can stabilize ecosystem flux rates in the face of perturbations.
Project description:Recent research has shown that biodiversity may has its greatest impact on ecosystem functioning in heterogeneous environments. However, the role of soil heterogeneity as a modulator of ecosystem responses to changes in biodiversity remains poorly understood, as few biodiversity studies have explicitly considered this important ecosystem feature.We conducted a microcosm experiment over two growing seasons to evaluate the joint effects of changes in plant functional groups (grasses, legumes, non-legume forbs and a combination of them), spatial distribution of soil nutrients (homogeneous and heterogeneous) and nutrient availability (50 and 100 mg of nitrogen [N] added as organic material) on plant productivity and surrogates of carbon, phosphorous and N cycling (?-glucosidase and acid phosphatase enzymes and in situ N availability, respectively).Soil nutrient heterogeneity interacted with nutrient availability and plant functional diversity to determine productivity and nutrient cycling responses. All the functional groups exhibited precise root foraging patterns. Above- and belowground productivity increased under heterogeneous nutrient supply. Surrogates of nutrient cycling were not directly affected by soil nutrient heterogeneity. Regardless of their above- and belowground biomass, legumes increased the availability of soil inorganic N and the activity of the acid phosphatase and ?-glucosidase enzymes.Our study emphasizes the role of soil nutrient heterogeneity as a modulator of ecosystem responses to changes in functional diversity beyond the species level. Functional group identity, rather than richness, can play a key role in determining the effects of biodiversity on ecosystem functioning.Synthesis. Our results highlight the importance of explicitly considering soil heterogeneity in diversity-ecosystem functioning experiments, where the identity of the plant functional group is of major importance. Such consideration will improve our ability to fully understand the role of plant diversity on ecosystem functioning in ubiquitous heterogeneous environments.
Project description:Global change drivers are rapidly altering resource availability and biodiversity. While there is consensus that greater biodiversity increases the functioning of ecosystems, the extent to which biodiversity buffers ecosystem productivity in response to changes in resource availability remains unclear. We use data from 16 grassland experiments across North America and Europe that manipulated plant species richness and one of two essential resources-soil nutrients or water-to assess the direction and strength of the interaction between plant diversity and resource alteration on above-ground productivity and net biodiversity, complementarity, and selection effects. Despite strong increases in productivity with nutrient addition and decreases in productivity with drought, we found that resource alterations did not alter biodiversity-ecosystem functioning relationships. Our results suggest that these relationships are largely determined by increases in complementarity effects along plant species richness gradients. Although nutrient addition reduced complementarity effects at high diversity, this appears to be due to high biomass in monocultures under nutrient enrichment. Our results indicate that diversity and the complementarity of species are important regulators of grassland ecosystem productivity, regardless of changes in other drivers of ecosystem function.
Project description:Understory plant communities play critical ecological roles in forest ecosystems. Both above- and below-ground ecosystem properties and processes influence these communities but relatively little is known about such effects at fine (i.e., one to several meters within-stand) scales, particularly for forests in which the canopy is dominated by a single species. An improved understanding of these effects is critical for understanding how understory biodiversity is regulated in such forests and for anticipating impacts of changing disturbance regimes. Our primary objective was to examine the patterns of fine-scale variation in understory plant communities and their relationships to above- and below-ground resource and environmental heterogeneity within mature lodgepole pine forests. We assessed composition and diversity of understory vegetation in relation to heterogeneity of both the above-ground (canopy tree density, canopy and tall shrub basal area and cover, downed wood biomass, litter cover) and below-ground (soil nutrient availability, decomposition, forest floor thickness, pH, and phospholipid fatty acids (PLFAs) and multiple carbon-source substrate-induced respiration (MSIR) of the forest floor microbial community) environment. There was notable variation in fine-scale plant community composition; cluster and indicator species analyses of the 24 most commonly occurring understory species distinguished four assemblages, one for which a pioneer forb species had the highest cover levels, and three others that were characterized by different bryophyte species having the highest cover. Constrained ordination (distance-based redundancy analysis) showed that two above-ground (mean tree diameter, litter cover) and eight below-ground (forest floor pH, plant available boron, microbial community composition and function as indicated by MSIR and PLFAs) properties were associated with variation in understory plant community composition. These results provide novel insights into the important ecological associations between understory plant community composition and heterogeneity in ecosystem properties and processes within forests dominated by a single canopy species.
Project description:Soil respiration plays a crucial role in global carbon cycling. While the response of soil respiration to abiotic drivers like soil temperature and moisture is fairly well understood, less is known about the effects of biotic drivers, such as plant above- and belowground productivity or plant diversity, and their interactions with abiotic drivers on soil respiration. Thus, current predictions of soil respiration to summer droughts might miss relevant biological drivers and their interactions with abiotic drivers. Since drought events are expected to increase in Central Europe in the future, we simulated early summer drought using rainout shelters at 19 grassland sites, which differed in plant productivity and species richness in central Germany in 2002 and 2003. We tested the potentially interacting effects of drought with biotic drivers, i.e. annual above-ground productivity, species richness and root biomass, on the drought response of soil respiration in temperate grasslands. In both years, drought led to a significant reduction in soil respiration. The drought-induced reduction in soil respiration was largely driven by the reduction in above-ground productivity in response to drought. The extent of the drought response of soil respiration was dependent on the species richness level of the site and this interacting effect was explainable by the variation in root biomass (root biomass and species richness were positively correlated). Our findings highlight the importance of biotic drivers for the quantification of the drought response of soil respiration in grasslands.
Project description:Anthropogenic activities have severely altered fluxes of nitrogen and phosphorus in ecosystems worldwide. In grasslands, subsequent negative effects are commonly attributed to competitive exclusion of plant species following increased above-ground biomass production. However, some studies have shown that this does not fully account for nutrient enrichment effects, questioning whether lowering competition by reducing grassland productivity through mowing or herbivory can mitigate the environmental impact of nutrient pollution. Furthermore, few studies so far discriminate between nitrogen and phosphorus pollution. We performed a full factorial experiment in greenhouse mesocosms combining nitrogen and phosphorus addition with two clipping regimes designed to relax above-ground competition. Next, we studied the survival and growth of seedlings of eight common European grassland species and found that five out of eight species showed higher survival under the clipping regime with the lowest above-ground competition. Phosphorus addition negatively affected seven plant species and nitrogen addition negatively affected four plant species. Importantly, the negative effects of nutrient addition and higher above-ground competition were independent of each other for all but one species. Our results suggest that at any given level of soil nutrients, relaxation of above-ground competition allows for higher seedling survival in grasslands. At the same time, even at low levels of above-ground competition, nutrient enrichment negatively affects survival as compared to nutrient-poor conditions. Therefore, although maintaining low above-ground competition appears essential for species' recruitment, for instance through mowing or herbivory, these management efforts are likely to be insufficient and we conclude that environmental policies aimed to reduce both excess nitrogen and particularly phosphorus inputs are also necessary.
Project description:Climate change has profound influences on plant community composition and ecosystem functions. However, its effects on plant community composition and biomass production are not well understood. A four-year field experiment was conducted to examine the effects of warming, nitrogen (N) addition, and their interactions on plant community composition and biomass production in a temperate meadow ecosystem in northeast China. Experimental warming had no significant effect on plant species richness, evenness, and diversity, while N addition highly reduced the species richness and diversity. Warming tended to reduce the importance value of graminoid species but increased the value of forbs, while N addition had the opposite effect. Warming tended to increase the belowground biomass, but had an opposite tendency to decrease the aboveground biomass. The influences of warming on aboveground production were dependent upon precipitation. Experimental warming had little effect on aboveground biomass in the years with higher precipitation, but significantly suppressed aboveground biomass in dry years. Our results suggest that warming had indirect effects on plant production via its effect on the water availability. Nitrogen addition significantly increased above- and below-ground production, suggesting that N is one of the most important limiting factors determining plant productivity in the studied meadow steppe. Significant interactive effects of warming plus N addition on belowground biomass were also detected. Our observations revealed that environmental changes (warming and N deposition) play significant roles in regulating plant community composition and biomass production in temperate meadow steppe ecosystem in northeast China.
Project description:Nutrient enrichment, particularly nitrogen, is an important determinant of plant community productivity, diversity and invasibility in a wetland ecosystem. It may contribute to increasing colonization and dominance of invasive species, such as Phragmites australis, especially during wetland restoration. Providing native species a competitive advantage over invasive species, manipulating soil nutrients (nitrogen) may be an effective strategy to control the invasive species and that management tool is essential to restore the degraded ecosystems. Therefore, we examined competition between Phragmites australis and Melaleuca ericifolia in a greenhouse setting with activated carbon (AC) treatments, followed by cutting of Phragmites shoots in nutrient-rich soils. Additionally, we evaluated the effect of AC on plant-free microcosms in the laboratory, to differentiate direct effects of AC on soil microbial functions from indirect effects. Overall, the objective was to test whether lowering nitrogen might be an effective approach for reducing Phragmites invasion in the wetland. The AC reduced Phragmites total biomass more significantly in repeated cut regime (57%) of Phragmites shoots compared to uncut regime (39%). Conversely, it increased Melaleuca total biomass by 41% and 68% in uncut and repeated cut regimes, respectively. Additionally, AC decreased more total nitrogen in above-ground biomass (41 to 55%) and non-structural carbohydrate in rhizome (21 to 65%) of Phragmites, and less total nitrogen reduction in above-ground biomass (25 to 24%) of Melaleuca in repeated cut compared to uncut regime. The significant negative correlation between Phragmites and Melaleuca total biomass was observed, and noticed that Phragmites acquired less biomass comparatively than Melaleuca in AC-untreated versus AC-treated pots across the cutting frequency. AC also caused significant changes to microbial community functions across Phragmites populations, namely nitrogen mineralization, nitrification, nitrogen microbial biomass and dehydrogenase activity (P???0.05) that may potentially explain changes in plant growth competition between Phragmites and Melaleuca. The overall effects on plant growth, however, may be partially microbially mediated, which was demonstrated through soil microbial functions. Results support the idea that reducing community vulnerability to invasion through nutrient (nitrogen) manipulations by AC with reducing biomass of invasive species may provide an effective strategy for invasive species management and ecosystem restoration.
Project description:Plants play a pivotal role in soil stabilization, with above-ground vegetation and roots combining to physically protect soil against erosion. It is possible that diverse plant communities boost root biomass, with knock-on positive effects for soil stability, but these relationships are yet to be disentangled.We hypothesize that soil erosion rates fall with increased plant species richness, and test explicitly how closely root biomass is associated with plant diversity.We tested this hypothesis in salt marsh grasslands, dynamic ecosystems with a key role in flood protection. Using step-wise regression, the influences of biotic (e.g. plant diversity) and abiotic variables on root biomass and soil stability were determined for salt marshes with two contrasting soil types: erosion-resistant clay (Essex, southeast UK) and erosion-prone sand (Morecambe Bay, northwest UK). A total of 132 (30-cm depth) cores of natural marsh were extracted and exposed to lateral erosion by water in a re-circulating flume.Soil erosion rates fell with increased plant species richness (R2 = 0.55), when richness was modelled as a single explanatory variable, but was more important in erosion-prone (R2 = 0.44) than erosion-resistant (R2 = 0.18) regions. As plant species richness increased from two to nine species·m-2, the coefficient of variation in soil erosion rate decreased significantly (R2 = 0.92). Plant species richness was a significant predictor of root biomass (R2 = 0.22). Step-wise regression showed that five key variables accounted for 80% of variation in soil erosion rate across regions. Clay-silt fraction and soil carbon stock were linked to lower rates, contributing 24% and 31%, respectively, to variation in erosion rate. In regional analysis, abiotic factors declined in importance, with root biomass explaining 25% of variation. Plant diversity explained 12% of variation in the erosion-prone sandy region.Our study indicates that soil stabilization and root biomass are positively associated with plant diversity. Diversity effects are more pronounced in biogeographical contexts where soils are erosion-prone (sandy, low organic content), suggesting that the pervasive influence of biodiversity on environmental processes also applies to the ecosystem service of erosion protection.
Project description:Seedlings of co-occurring species vary in their response to resource availability and this has implications for the conservation and management of forests. Differential shade-tolerance is thought to influence seedling performance in mixed Nothofagus betuloides-Nothofagus pumilio forests of Patagonia. However, these species also vary in their soil nutrient requirements. To determine the effects of light and soil nutrient resources on small seedlings we examined responses to an experimental reduction in canopy tree root competition through root trenching and restricting soil nutrient depletion through the addition of fertilizer. To understand the effect of light these treatments were undertaken in small canopy gaps and nearby beneath undisturbed canopy with lower light levels. Seedling diameter growth was greater for N. pumilio and height growth was greater for N. betuloides. Overall, diameter and height growth were greater in canopy gaps than beneath undisturbed canopy. Such growths were also greater with fertilizer and root trenching treatments, even beneath undisturbed canopy. Seedling survival was lower under such treatments, potentially reflecting thinning facilitated by resource induced growth. Finally, above-ground biomass did not vary among species although the less shade tolerant N. pumilio had higher below-ground biomass and root to shoot biomass ratio than the more shade tolerant N. betuloides. Above- and below-ground biomass were higher in canopy gaps so that the root to shoot biomass ratio was similar to that beneath undisturbed canopy. Above-ground biomass was also higher with fertilizer and root trenching treatments and that lowered the root to shoot biomass ratio. Restricting soil nutrient depletion allowed seedlings of both species to focus their responses above-ground. Our results support a view that soil nutrient resources, as well as the more commonly studied light resources, are important to seedlings of Nothofagus species occurring on infertile soils.