Defoliation reduces soil biota - and modifies stimulating effects of elevated CO2.
ABSTRACT: To understand the responses to external disturbance such as defoliation and possible feedback mechanisms at global change in terrestrial ecosystems, it is necessary to examine the extent and nature of effects on aboveground-belowground interactions. We studied a temperate heathland system subjected to experimental climate and atmospheric factors based on prognoses for year 2075 and further exposed to defoliation. By defoliating plants, we were able to study how global change modifies the interactions of the plant-soil system. Shoot production, root biomass, microbial biomass, and nematode abundance were assessed in the rhizosphere of manually defoliated patches of Deschampsia flexuosa in June in a full-factorial FACE experiment with the treatments: increased atmospheric CO 2, increased nighttime temperatures, summer droughts, and all of their combinations. We found a negative effect of defoliation on microbial biomass that was not apparently affected by global change. The negative effect of defoliation cascades through to soil nematodes as dependent on CO 2 and drought. At ambient CO 2, drought and defoliation each reduced nematodes. In contrast, at elevated CO 2, a combination of drought and defoliation was needed to reduce nematodes. We found positive effects of CO 2 on root density and microbial biomass. Defoliation affected soil biota negatively, whereas elevated CO 2 stimulated the plant-soil system. This effect seen in June is contrasted by the effects seen in September at the same site. Late season defoliation increased activity and biomass of soil biota and more so at elevated CO 2. Based on soil biota responses, plants defoliated in active growth therefore conserve resources, whereas defoliation after termination of growth results in release of resources. This result challenges the idea that plants via exudation of organic carbon stimulate their rhizosphere biota when in apparent need of nutrients for growth.
Project description:Sub-arctic birch forests (<i>Betula pubescens</i> Ehrh. ssp. <i>czerepanovii</i>) periodically suffer large-scale defoliation events caused by the caterpillars of the geometrid moths <i>Epirrita autumnata</i> and <i>Operophtera brumata</i>. Despite their obvious influence on ecosystem primary productivity, little is known about how the associated reduction in belowground C allocation affects soil processes. We quantified the soil response following a natural defoliation event in sub-arctic Sweden by measuring soil respiration, nitrogen availability and ectomycorrhizal fungi (EMF) hyphal production and root tip community composition. There was a reduction in soil respiration and an accumulation of soil inorganic N in defoliated plots, symptomatic of a slowdown of soil processes. This coincided with a reduction of EMF hyphal production and a shift in the EMF community to lower autotrophic C-demanding lineages (for example, /russula-lactarius). We show that microbial and nutrient cycling processes shift to a slower, less C-demanding state in response to canopy defoliation. We speculate that, amongst other factors, a reduction in the potential of EMF biomass to immobilise excess mineral nitrogen resulted in its build-up in the soil. These defoliation events are becoming more geographically widespread with climate warming, and could result in a fundamental shift in sub-arctic ecosystem processes and properties. EMF fungi may be important in mediating the response of soil cycles to defoliation and their role merits further investigation.
Project description:Anthropogenic global change alters the activity and functional composition of soil communities that are responsible for crucial ecosystem functions and services. Two of the most pervasive global change drivers are drought and nutrient enrichment. However, the responses of soil organisms to interacting global change drivers remain widely unknown. We tested the interactive effects of extreme drought and fertilization on soil biota ranging from microbes to invertebrates across seasons. We expected drought to reduce the activity of soil organisms and fertilization to induce positive bottom-up effects via increased plant productivity. Furthermore, we hypothesized fertilization to reinforce drought effects through enhanced plant growth, resulting in even drier soil conditions. Our results revealed that drought had detrimental effects on soil invertebrate feeding activity and simplified nematode community structure, whereas soil microbial activity and biomass were unaffected. Microbial biomass increased in response to fertilization, whereas invertebrate feeding activity substantially declined. Notably, these effects were consistent across seasons. The dissimilar responses suggest that soil biota differ vastly in their vulnerability to global change drivers. Thus, important ecosystem processes like decomposition and nutrient cycling, which are driven by the interdependent activity of soil microorganisms and invertebrates, may be disrupted under future conditions.
Project description:In closed-canopy forests, gap formation and closure are thought to be major drivers of forest dynamics. Crown defoliation by insects, however, may also influence understory resource levels and thus forest dynamics. We evaluate the effect of a forest tent caterpillar outbreak on understory light availability, soil nutrient levels and tree seedling height growth in six sites with contrasting levels of canopy defoliation in a hardwood forest in northern lower Michigan. We compared resource levels and seedling growth of six hardwood species before, during and in the three years after the outbreak (2008-2012). Canopy openness increased strongly during the forest tent caterpillar outbreak in the four moderately and severely defoliated sites, but not in lightly defoliated sites. Total inorganic soil nitrogen concentrations increased in response to the outbreak in moderately and severely defoliated sites. The increase in total inorganic soil nitrogen was driven by a strong increase in soil nitrate, and tended to become stronger with increasing site defoliation. Seedling height growth increased for all species in the moderately and severely defoliated sites, but not in lightly defoliated sites, either during the outbreak year or in the year after the outbreak. Growth increases did not become stronger with increasing site defoliation, but were strongest in a moderately defoliated site with high soil nutrient levels. Growth increases tended to be strongest for the shade intolerant species Fraxinus americana and Prunus serotina, and the shade tolerant species Ostrya virginiana. The strong growth response of F. americana and P. serotina suggests that recurring forest tent caterpillar outbreaks may facilitate the persistence of shade intolerant species in the understory in the absence of canopy gaps. Overall, our results suggest that recurrent canopy defoliation resulting from cyclical forest insect outbreaks may be an additional driver of dynamics in temperate closed-canopy forests.
Project description:Plant ecologists have debated the mechanisms used by plants to cope with the impact of herbivore damage. While plant resistance mechanisms have received much attention, plant compensatory growth as a type of plant tolerance mechanisms has been less studied. We conducted a greenhouse experiment to evaluate compensatory growth for trembling aspen (Populus tremuloides) seedlings under varying intensities and frequencies of simulated defoliation, with or without nutrient enriched media. For the purpose of this study, changes in biomass production and non-structural carbohydrate concentrations (NSC) of roots and leaves were considered compensatory responses. All defoliated seedlings showed biomass accumulation under low defoliation intensity and frequency, regardless of resource availability; however, as defoliation intensity and frequency increased, compensatory growth of seedlings was altered depending on resource availability. Seedlings in a resource-rich environment showed complete compensation, in contrast responses ranged from undercompensation to complete compensation in a resource-limited environment. Furthermore, at the highest defoliation intensity and frequency, NSC concentrations in leaves and roots were similar between defoliated and non-defoliated seedlings in a resource-rich environment; in contrast, defoliated seedlings with limited resources sustained the most biomass loss, had lower amounts of stored NSC. Using these results, we developed a new predictive framework incorporating the interactions between frequency and intensity of defoliation and resource availability as modulators of plant compensatory responses.
Project description:Mediterranean pine forests display high resilience after extreme climatic events such as severe droughts. However, recent dry spells causing growth decline and triggering forest dieback challenge the capacity of some forests to recover following major disturbances. To describe how resilient the responses of forests to drought can be, we quantified growth dynamics in plantations of two pine species (Scots pine, black pine) located in south-eastern Spain and showing drought-triggered dieback. Radial growth was characterized at inter- (tree-ring width) and intra-annual (xylogenesis) scales in three defoliation levels. It was assumed that the higher defoliation the more negative the impact of drought on tree growth. Tree-ring width chronologies were built and xylogenesis was characterized 3 years after the last severe drought occurred. Annual growth data and the number of tracheids produced in different stages of xylem formation were related to climate data at several time scales. Drought negatively impacted growth of the most defoliated trees in both pine species. In Scots pine, xylem formation started earlier in the non-defoliated than in the most defoliated trees. Defoliated trees presented the shortest duration of the radial-enlargement phase in both species. On average the most defoliated trees formed 60% of the number of mature tracheids formed by the non-defoliated trees in both species. Since radial enlargement is the xylogenesis phase most tightly related to final growth, this explains why the most defoliated trees grew the least due to their altered xylogenesis phases. Our findings indicate a very limited resilience capacity of drought-defoliated Scots and black pines. Moreover, droughts produce legacy effects on xylogenesis of highly defoliated trees which could not recover previous growth rates and are thus more prone to die.
Project description:Drought stress causes a reduction in tree growth and forest productivity, which could be aggravated by climate warming and defoliation due to moth outbreaks. We investigate how European gypsy moth (<i>Lymantria dispar dispar</i> L., Lepidoptera: Erebidae) outbreak and related climate conditions affected growth and wood features in host and non-host tree species in north-western Spain. There, radiata pine (<i>Pinus radiata</i> D. Don) plantations and chestnut (<i>Castanea sativa</i> Mill.) stands were defoliated by the moth larvae, whereas Maritime pine (<i>Pinus pinaster</i> Ait.) was not defoliated. The gypsy moth outbreak peaked in 2012 and 2013, and it was preceded by very warm spring conditions in 2011 and a dry-warm 2011-2012 winter. Using dendrochronology we compared growth responses to climate and defoliation of host species (radiata pine, chestnut) with the non-host species (Maritime pine). We also analyzed wood density derived from X-ray densitometry in defoliated and non-defoliated trees of radiata pine. We aimed to: (i) disentangle the relative effects of defoliation and climate stress on radial growth, and (ii) characterize defoliated trees of radiata pine according to their wood features (ring-width, maximum and minimum density). Radial growth during the outbreak (2012-2013) decreased on average 74% in defoliated (>50% of leaf area removed) trees of radiata pine, 43% in defoliated trees of chestnut, and 4% in non-defoliated trees of Maritime pine. After applying a BACI (Before-After-Control-Impact) type analysis, we concluded that the difference in the pattern of radial growth before and during the defoliation event was more likely due to the differences in climate between these two periods. Radiata pines produced abundant latewood intra-annual density fluctuations in 2006 and 2009 in response to wet summer conditions, suggesting a high climatic responsiveness. Minimum wood density was lower in defoliated than in non-defoliated trees of radiata pine prior to the outbreak, but increased during the outbreak. The pre-outbreak difference in minimum wood density suggests that the trees most affected by the outbreak produced tracheids with wider lumen and were more susceptible to drought stress. Results of this study illustrate (i) that the pattern of radial growth alone may be not a good indicator for reconstructing past defoliation events and (ii) that wood variables are reliable indicators for assessing the susceptibility of radiata pine to defoliation by the gypsy moth.
Project description:Herbivory is one of the most globally distributed disturbances affecting carbon (C)-cycling in trees, yet our understanding of how it alters tree C-allocation to different functions such as storage, growth or rhizodeposition is still limited. Prioritized C-allocation to storage replenishment vs growth could explain the fast recovery of C-storage pools frequently observed in growth-reduced defoliated trees. We performed continuous 13C-labeling coupled to clipping to quantify the effects of simulated browsing on the growth, leaf morphology and relative allocation of stored vs recently assimilated C to the growth (bulk biomass) and non-structural carbohydrate (NSC) stores (soluble sugars and starch) of the different organs of two tree species: diffuse-porous (Betula pubescens Ehrh.) and ring-porous (Quercus petraea [Matt.] Liebl.). Carbon-transfers from plants to bulk and rhizosphere soil were also evaluated. Clipped birch and oak trees shifted their C-allocation patterns above-ground as a means to recover from defoliation. However, such increased allocation to current-year stems and leaves did not entail reductions in the allocation to the rhizosphere, which remained unchanged between clipped and control trees of both species. Betula pubescens and Q. petraea showed differences in their vulnerability and recovery strategies to clipping, the ring-porous species being less affected in terms of growth and architecture by clipping than the diffuse-porous. These contrasting patterns could be partly explained by differences in their C cycling after clipping. Defoliated oaks showed a faster recovery of their canopy biomass, which was supported by increased allocation of new C, but associated with large decreases in their fine root biomass. Following clipping, both species recovered NSC pools to a larger extent than growth, but the allocation of 13C-labeled photo-assimilates into storage compounds was not increased as compared with controls. Despite their different response to clipping, our results indicate no preventative allocation into storage occurred during the first year after clipping in either of the species.
Project description:Climate change is expected to increase future abiotic stresses on ecosystems through extreme weather events leading to more extreme drought and rainfall incidences [Jentsch A, et al. (2007) Front Ecol Environ 5(7):365-374]. These fluctuations in precipitation may affect soil biota, soil processes [Evans ST, Wallenstein MD (2012) Biogeochemistry 109:101-116], and the proportion of exotics in invaded plant communities [Jiménez MA, et al. (2011) Ecol Lett 14:1277-1235]. However, little is known about legacy effects in soil on the performance of exotics and natives in invaded plant communities. Here we report that drought and rainfall effects on soil processes and biota affect the performance of exotics and natives in plant communities. We performed two mesocosm experiments. In the first experiment, soil without plants was exposed to drought and/or rainfall, which affected soil N availability. Then the initial soil moisture conditions were restored, and a mixed community of co-occurring natives and exotics was planted and exposed to drought during growth. A single stress before or during growth decreased the biomass of natives, but did not affect exotics. A second drought stress during plant growth resetted the exotic advantage, whereas native biomass was not further reduced. In the second experiment, soil inoculation revealed that drought and/or rainfall influenced soil biotic legacies, which promoted exotics but suppressed natives. Our results demonstrate that extreme weather events can cause legacy effects in soil biota, promoting exotics and suppressing natives in invaded plant communities, depending on the type, frequency, and timing of extreme events.
Project description:This study mainly aimed to investigate the effects of dark septate endophytes (DSE) (Acrocalymma vagum, Paraboeremia putaminum, and Fusarium acuminatum) on the growth and microbial community composition in the rhizosphere soil of a medicinal plant, licorice (Glycyrrhiza uralensis), grown in the non-sterile soil under drought stress. The results showed that three DSE strains could effectively colonize the plant roots and form a strain-dependent symbiosis with licorice. Although drought stress declined the growth of licorice plants, these decreases were partly recovered by DSE inoculation. Specifically, the inoculation of A. vagum and P. putaminum significantly increased the biomass and glycyrrhizin content, whereas A. vagum and F. acuminatum increased glycyrrhizic acid content of host plants under drought stress. However, the inoculation of F. acuminatum showed significant negative effects on the shoot, root, and total biomass of licorice plants. In addition, the effects of DSE inoculation on the morphological, photosynthetic, and antioxidant parameters of licorice plants, and mineral nutrient and microbial community composition in the rhizosphere soil were dependent on the DSE species as well as water regime. Interestingly, DSE inoculation significantly increased AM fungi content under drought stress. In addition, DSE associated with water had a significant positive influence on soil organic matter, available phosphorus (P), AM fungi, leaf number, soluble protein, SOD activity, total root length, root branch, and glycyrrhizic acid content. Based on the results of variance partitioning analysis, 17.0, 34.0, 14.9, 40.1, 28.2, and 18.0% variations in shoot morphology, root morphology, plant biomass, active ingredient, photosynthetic parameters, and antioxidant parameters, respectively, were attributable to the presence of certain soil microorganisms. These findings suggest the possibility that DSE inoculation improved the root development and nutrient absorption of host plants, altered the soil microbiota, and might also contribute to plant growth and survival under drought conditions. As A. vagum exhibited positive effects on the plant biomass, morphological and physiological parameters, and active ingredient content in licorice plants under drought stress, it was considered to be the best fungus for licorice cultivation. These results contribute to the understanding of the ecological function of DSE fungi in dryland agriculture.
Project description:The role of soil fauna in crucial ecosystem services such as nutrient cycling remains poorly quantified, mainly because of the overly reductionistic approach adopted in most experimental studies. Given that increasing nitrogen inputs in various ecosystems influence the structure and functioning of soil microbes and the activity of fauna, we aimed to quantify the role of the entire soil nematode community in nutrient mineralization in an experimental set-up emulating nutrient-rich field conditions and accounting for crucial interactions amongst the soil microbial communities and plants. To this end, we reconstructed a complex soil foodweb in mesocosms that comprised largely undisturbed native microflora and the entire nematode community added into defaunated soil, planted with Lolium perenne as a model plant, and amended with fresh grass-clover residues. We determined N and P availability and plant uptake, plant biomass and abundance and structure of the microbial and nematode communities during a three-month incubation. The presence of nematodes significantly increased plant biomass production (+9%), net N (+25%) and net P (+23%) availability compared to their absence, demonstrating that nematodes link below- and above-ground processes, primarily through increasing nutrient availability. The experimental set-up presented allows to realistically quantify the crucial ecosystem services provided by the soil biota.