Drought legacy effects on root morphological traits and plant biomass via soil biota feedback.

Drought causes soil feedback effects on plant performance. However, how the linkages between conditioned soil biota and root traits contribute to explain plant-soil feedback (PSF) as a function of drought is unknown. We utilized soil inoculum from a conditioning experiment where grassland species grew under well-watered and drought conditions, and their soil fungi were analyzed. Under well-watered conditions, we grew 21 grassland species with those inocula from either conspecific or heterospecific soils. At harvest, plant biomass and root traits were measured. Negative PSF (higher biomass in heterospecific than in conspecific soils) was predominant, and favored in drought-conditioned soils. Previous drought affected the relationship between root traits and fungal groups. Specific root surface area (SRSA) was higher in heterospecific than in conspecific droughted soils and was linked to an increase in saprotroph richness. Overall, root diameter was higher in conspecific soils and was linked to mutualist and pathogen composition, whereas the decrease of root : shoot in heterospecific soils was linked to pathogenic fungi. Drought legacy affects biomass and root morphological traits via conditioned soil biota, even after the drought conditions have disappeared. This provides new insights into the role that soil biota have modulating PSF responses to drought.

Above- and below-ground plant traits are not consistent in response to drought and competition treatments.

BACKGROUND AND AIMS: Our understanding of plant responses to biotic and abiotic drivers is largely based on above-ground plant traits, with little focus on below-ground traits despite their key role in water and nutrient uptake. Here, we aimed to understand the extent to which above- and below-ground traits are co-ordinated, and how these traits respond to soil moisture gradients and plant intraspecific competition. METHODS: We chose seedlings of five tropical tree species and grew them in a greenhouse for 16 weeks under a soil moisture gradient [low (drought), medium and high (well-watered) moisture levels] with and without intraspecific competition. At harvest, we measured nine above- and five below-ground traits of all seedlings based on standard protocols. KEY RESULTS: In response to the soil moisture gradient, above-ground traits are found to be consistent with the leaf economics spectrum, whereas below-ground traits are inconsistent with the root economics spectrum. We found high specific leaf area and total leaf area in well-watered conditions, while high leaf dry matter content, leaf thickness and stem dry matter content were observed in drought conditions. However, below-ground traits showed contrasting patterns, with high specific root length but low root branching index in the low water treatment. The correlations between above- and below-ground traits across the soil moisture gradient were variable, i.e. specific leaf area was positively correlated with specific root length, while it was negatively correlated with root average diameter across moisture levels. However, leaf dry matter content was unexpectedly positively correlated with both specific root length and root branching index. Intraspecific competition has influenced both above- and below-ground traits, but interacted with soil moisture to affect only below-ground traits. Consistent with functional equilibrium theory, more biomass was allocated to roots under drought conditions, and to leaves under sufficient soil moisture conditions. CONCLUSIONS: Our results indicate that the response of below-ground traits to plant intraspecific competition and soil moisture conditions may not be inferred using above-ground traits, suggesting that multiple resource use axes are needed to understand plant ecological strategies. Lack of consistent leaf-root trait correlations across the soil moisture gradient highlight the multidimensionality of plant trait relationships which needs more exploration.

Drought induces shifts in soil fungal communities that can be linked to root traits across 24 plant species.

Root traits respond to drought in a species-specific manner, but little is known about how soil fungal communities and root traits respond to drought in concert. In a glasshouse experiment, we determined the response of soil pathogens, saprotrophs, and mutualistic and all fungi associated with the roots of 24 plant species subjected to drought. At harvest, soil fungal communities were characterized by sequencing. Data on root traits were extracted from a previously published work. Differences in fungal beta diversity between drought and control were plant species specific. For some species, saprotrophic fungi increased in relative abundance and richness with drought, whereas mutualistic fungi showed the opposite pattern. Community structure of pathogenic fungi was plant species specific but was slightly affected by drought. Pathogen composition was correlated with specific root surface area and root : shoot, saprotroph abundance with root tissue density, whereas mutualist composition was correlated with root : shoot. All these were the fungal attributes that best predicted shoot mass. Fungal response to drought depended highly on the fungal group and was related to root trait adjustments to water scarcity. This provides new insights into the role that root trait adjustments to drought may have in modulating plant-fungus interactions in grasslands ecosystems.

Myristate and the ecology of AM fungi: significance, opportunities, applications and challenges.

A recent study by Sugiura and coworkers reported the non-symbiotic growth and spore production of an arbuscular mycorrhizal (AM) fungus, Rhizophagus irregularis, when the fungus received an external supply of certain fatty acids, myristates (C:14). This discovery follows the insight that AM fungi receive fatty acids from their hosts when in symbiosis. If this result holds up and can be repeated under nonsterile conditions and with a broader range of fungi, it has numerous consequences for our understanding of AM fungal ecology, from the level of the fungus, at the plant community level, and to functional consequences in ecosystems. In addition, myristate may open up several avenues from a more applied perspective, including improved fungal culture and supplementation of AM fungi or inoculum in the field. We here map these potential opportunities, and additionally offer thoughts on potential risks of this potentially new technology. Lastly, we discuss the specific research challenges that need to be overcome to come to an understanding of the potential role of myristate in AM ecology.

Effects of Microplastic Fibers and Drought on Plant Communities.

Microplastics in soils can affect plant performance, as shown in studies using individual plants. However, we currently have no information about potential effects on plant community productivity and structure. In a plant community consisting of seven plant species that co-occur in temperate grassland ecosystems, we thus investigated the effect of microplastics (i.e., microfibers) and drought, a factor with which microfibers might interact, on plant productivity and community structure. Our results showed that at the community level, shoot and root mass decreased with drought but increased with microfibers, an effect likely linked to reduced soil bulk density, improved aeration, and better penetration of roots in the soil. Additionally, we observed that microfibers affected plant community structure. Species such as Calamagrostis, invasive in Europe, and the allelophatic Hieracium, became more dominant with microfibers, while species that potentially have the ability to facilitate the establishment of other plant species (e.g., Holcus), decreased in biomass. As microfibers affect plant species dominance, the examination of cascade effects on ecosystem functions should be a high priority for future research.

Disentangling above- and below-ground facilitation drivers in arid environments: the role of soil microorganisms, soil properties and microhabitat.

Nurse plants promote establishment of other plant species by buffering climate extremes and improving soil properties. Soil biota plays an important role, but an analysis to disentangle the effects of soil microorganisms, soil properties and microclimate on facilitation is lacking. In three microhabitats (gaps, small and large Retama shrubs), we placed six microcosms with sterilized soil, two per soil origin (i.e. from each microhabitat). One in every pair received an alive, and the other a sterile, inoculum from its own soil. Seeds of annual plants were sown into the microcosms. Germination, survival and biomass were monitored. Soil bacterial communities were characterized by pyrosequencing. Germination in living Retama inoculum was nearly double that of germination in sterile inoculum. Germination was greater under Retama canopies than in gaps. Biomass was up to three times higher in nurse than in gap soils. Soil microorganisms, soil properties and microclimate showed a range of positive to negative effects on understory plants depending on species identity and life stage. Nurse soil microorganisms promoted germination, but the effect was smaller than the positive effects of soil properties and microclimate under nurses. Nurse below-ground environment (soil properties and microorganisms) promoted plant growth and survival more than nurse microhabitat.

Drought induces moderate, diverse changes in the odour of grassland species.

Plants react to drought stress with numerous changes including altered emissions of volatile organic compounds (VOC) from leaves, which provide protection against oxidative tissue damage and mediate numerous biotic interactions. Despite the share of grasslands in the terrestrial biosphere, their importance as carbon sinks and their contribution to global biodiversity, little is known about the influence of drought on VOC profiles of grassland species. Using coupled gas chromatography-mass spectrometry, we analysed the odorants emitted by 22 European grassland species exposed to an eight-week-lasting drought treatment (DT; 30% water holding capacity, WHC). We focused on the odorants emitted during the light phase from whole plant shoots in their vegetative stage. Emission rates were standardised to the dry weight of each shoot. Well-watered (WW) plants (70% WHC) served as control. Drought-induced significant changes included an increase in total emission rates of plant VOC in six and a decrease in three species. Diverging effects on the number of emitted VOC (chemical richness) or on the Shannon diversity of the VOC profiles were detected in 13 species. Biosynthetic pathways-targeted analyses revealed 13 species showing drought-induced higher emission rates of VOC from one, two, three, or four major biosynthetic pathways (lipoxygenase, shikimate, mevalonate and methylerythritol phosphate pathway), while six species exhibited reduced emission rates from one or two of these pathways. Similarity trees of odorant profiles and their drought-induced changes based on a biosynthetically informed distance metric did not match species phylogeny. However, a phylogenetic signal was detected for the amount of terpenoids released by the studied species under WW and DT conditions. A comparative analysis of emission rates of single compounds released by WW and DT plants revealed significant VOC profile dissimilarities in four species only. The moderate drought-induced changes in the odorant emissions of grassland species are discussed with respect to their impact on trophic interactions across the food web. (294 words).

The promoting effects of soil microplastics on alien plant invasion depend on microplastic shape and concentration.

Both alien plant invasions and soil microplastic pollution have become a concerning threat for terrestrial ecosystems, with consequences on the human well-being. However, our current knowledge of microplastic effects on the successful invasion of plants remains limited, despite numerous studies demonstrating the direct and indirect impacts of microplastics on plant performance. To address this knowledge gap, we conducted a greenhouse experiment involving the mixtures of soil and low-density polyethylene (LDPE) microplastic pellets and fragments at the concentrations of 0, 0.5 % and 2.0 %. Additionally, we included Solidago decurrens (native plant) and S. canadensis (alien invasive plant) as the target plants. Each pot contained an individual of either species, after six-month cultivation, plant biomass and antioxidant enzymes, as well as soil properties including soil moisture, pH, available nutrient, and microbial biomass were measured. Our results indicated that microplastic effects on soil properties and plant growth indices depended on the Solidago species, microplastic shapes and concentrations. For example, microplastics exerted positive effects on soil moisture of the soil with native species but negative effects with invasive species, which were impacted by microplastic shapes and concentrations, respectively. Microplastics significantly impacted catalase (P < 0.05) and superoxide dismutase (P < 0.01), aboveground biomass (P < 0.01), and belowground/aboveground biomass (P < 0.01) of the native species depending on microplastic shapes, but no significant effects on those of the invasive species. Furthermore, microplastics effects on soil properties, nutrient, nutrient ratio, and plant antioxidant enzyme activities contributed to plant biomass differently among these two species. These results suggested that the microplastics exerted a more pronounced impact on native Solidago plants than the invasive ones. This implies that the alien invasive species displays greater resistance to microplastic pollution, potentially promoting their invasion. Overall, our study contributes to a better understanding of the promoting effects of microplastic pollution on plant invasion.

Legacy effect of microplastics on plant-soil feedbacks.

Microplastics affect plants and soil biota and the processes they drive. However, the legacy effect of microplastics on plant-soil feedbacks is still unknown. To address this, we used soil conditioned from a previous experiment, where Daucus carota grew with 12 different microplastic types (conditioning phase). Here, we extracted soil inoculum from those 12 soils and grew during 4 weeks a native D. carota and a range-expanding plant species Calamagrostis epigejos in soils amended with this inoculum (feedback phase). At harvest, plant biomass and root morphological traits were measured. Films led to positive feedback on shoot mass (higher mass with inoculum from soil conditioned with microplastics than with inoculum from control soil). Films may decrease soil water content in the conditioning phase, potentially reducing the abundance of harmful soil biota, which, with films also promoting mutualist abundance, microbial activity and carbon mineralization, would positively affect plant growth in the feedback phase. Foams and fragments caused positive feedback on shoot mass likely via positive effects on soil aeration in the conditioning phase, which could have increased mutualistic biota and soil enzymatic activity, promoting plant growth. By contrast, fibers caused negative feedback on root mass as this microplastic may have increased soil water content in the conditioning phase, promoting the abundance of soil pathogens with negative consequences for root mass. Microplastics had a legacy effect on root traits: D. carota had thicker roots probably for promoting mycorrhizal associations, while C. epigejos had reduced root diameter probably for diminishing pathogenic infection. Microplastic legacy on soil can be positive or negative depending on the plant species identity and may affect plant biomass primarily via root traits. This legacy may contribute to the competitive success of range-expanding species via positive effects on root mass (foams) and on shoot mass (PET films). Overall, microplastics depending on their shape and polymer type, affect plant-soil feedbacks.

Microplastic Shape, Polymer Type, and Concentration Affect Soil Properties and Plant Biomass.

Microplastics may enter the soil in a wide range of shapes and polymers. However, little is known about the effects that microplastics of different shapes, polymers, and concentration may have on soil properties and plant performance. To address this, we selected 12 microplastics representing different shapes (fibers, films, foams, and fragments) and polymers, and mixed them each with soil at a concentration of 0.1, 0.2, 0.3, and 0.4%. A phytometer (Daucus carota) grew in each pot during 4 weeks. Shoot, root mass, soil aggregation, and microbial activity were measured. All shapes increased plant biomass. Shoot mass increased by ∼27% with fibers, ∼60% with films, ∼45% with foams, and by ∼54% with fragments, as fibers hold water in the soil for longer, films decrease soil bulk density, and foams and fragments can increase soil aeration and macroporosity, which overall promote plant performance. By contrast, all shapes decreased soil aggregation by ∼25% as microplastics may introduce fracture points into aggregates and due to potential negative effects on soil biota. The latter may also explain the decrease in microbial activity with, for example, polyethylene films. Our findings show that shape, polymer type, and concentration are key properties when studying microplastic effects on terrestrial systems.

Fungal diversity regulates plant-soil feedbacks in temperate grassland.

Feedbacks between plants and soil microbial communities play an important role in vegetation dynamics, but the underlying mechanisms remain unresolved. Here, we show that the diversity of putative pathogenic, mycorrhizal, and saprotrophic fungi is a primary regulator of plant-soil feedbacks across a broad range of temperate grassland plant species. We show that plant species with resource-acquisitive traits, such as high shoot nitrogen concentrations and thin roots, attract diverse communities of putative fungal pathogens and specialist saprotrophs, and a lower diversity of mycorrhizal fungi, resulting in strong plant growth suppression on soil occupied by the same species. Moreover, soil properties modulate feedbacks with fertile soils, promoting antagonistic relationships between soil fungi and plants. This study advances our capacity to predict plant-soil feedbacks and vegetation dynamics by revealing fundamental links between soil properties, plant resource acquisition strategies, and the diversity of fungal guilds in soil.