Timing and intensity of heat and drought stress determine wheat yield losses in Germany.

Crop yields are increasingly affected by climate change-induced weather extremes in Germany. However, there is still little knowledge of the specific crop-climate relations and respective heat and drought stress-induced yield losses. Therefore, we configure weather indices (WIs) that differ in the timing and intensity of heat and drought stress in wheat (Triticum aestivum L.). We construct these WIs using gridded weather and phenology time series data from 1995 to 2019 and aggregate them with Germany-wide municipality level on-farm wheat yield data. We statistically analyze the WI's explanatory power and region-specific effect size for wheat yield using linear mixed models. We found the highest explanatory power during the stem elongation and booting phase under moderate drought stress and during the reproductive phase under moderate heat stress. Furthermore, we observed the highest average yield losses due to moderate and extreme heat stress during the reproductive phase. The highest heat and drought stress-induced yield losses were observed in Brandenburg, Saxony-Anhalt, and northern Bavaria, while similar heat and drought stresses cause much lower yield losses in other regions of Germany.

Simulating Root Growth as a Function of Soil Strength and Yield With a Field-Scale Crop Model Coupled With a 3D Architectural Root Model.

Accurate prediction of root growth and related resource uptake is crucial to accurately simulate crop growth especially under unfavorable environmental conditions. We coupled a 1D field-scale crop-soil model running in the SIMPLACE modeling framework with the 3D architectural root model CRootbox on a daily time step and implemented a stress function to simulate root elongation as a function of soil bulk density and matric potential. The model was tested with field data collected during two growing seasons of spring barley and winter wheat on Haplic Luvisol. In that experiment, mechanical strip-wise subsoil loosening (30-60 cm) (DL treatment) was tested, and effects on root and shoot growth at the melioration strip as well as in a control treatment were evaluated. At most soil depths, strip-wise deep loosening significantly enhanced observed root length densities (RLDs) of both crops as compared to the control. However, the enhanced root growth had a beneficial effect on crop productivity only in the very dry season in 2018 for spring barley where the observed grain yield at the strip was 18% higher as compared to the control. To understand the underlying processes that led to these yield effects, we simulated spring barley and winter wheat root and shoot growth using the described field data and the model. For comparison, we simulated the scenarios with the simpler 1D conceptual root model. The coupled model showed the ability to simulate the main effects of strip-wise subsoil loosening on root and shoot growth. It was able to simulate the adaptive plasticity of roots to local soil conditions (more and thinner roots in case of dry and loose soil). Additional scenario runs with varying weather conditions were simulated to evaluate the impact of deep loosening on yield under different conditions. The scenarios revealed that higher spring barley yields in DL than in the control occurred in about 50% of the growing seasons. This effect was more pronounced for spring barley than for winter wheat. Different virtual root phenotypes were tested to assess the potential of the coupled model to simulate the effect of varying root traits under different conditions.

Nutrient deficiency effects on root architecture and root-to-shoot ratio in arable crops.

Plant root traits play a crucial role in resource acquisition and crop performance when soil nutrient availability is low. However, the respective trait responses are complex, particularly at the field scale, and poorly understood due to difficulties in root phenotyping monitoring, inaccurate sampling, and environmental conditions. Here, we conducted a systematic review and meta-analysis of 50 field studies to identify the effects of nitrogen (N), phosphorous (P), or potassium (K) deficiencies on the root systems of common crops. Root length and biomass were generally reduced, while root length per shoot biomass was enhanced under N and P deficiency. Root length decreased by 9% under N deficiency and by 14% under P deficiency, while root biomass was reduced by 7% in N-deficient and by 25% in P-deficient soils. Root length per shoot biomass increased by 33% in N deficient and 51% in P deficient soils. The root-to-shoot ratio was often enhanced (44%) under N-poor conditions, but no consistent response of the root-to-shoot ratio to P-deficiency was found. Only a few K-deficiency studies suited our approach and, in those cases, no differences in morphological traits were reported. We encountered the following drawbacks when performing this analysis: limited number of root traits investigated at field scale, differences in the timing and severity of nutrient deficiencies, missing data (e.g., soil nutrient status and time of stress), and the impact of other conditions in the field. Nevertheless, our analysis indicates that, in general, nutrient deficiencies increased the root-length-to-shoot-biomass ratios of crops, with impacts decreasing in the order deficient P > deficient N > deficient K. Our review resolved inconsistencies that were often found in the individual field experiments, and led to a better understanding of the physiological mechanisms underlying root plasticity in fields with low nutrient availability.

Effects of mixing two legume species at seedling stage under different environmental conditions.

While intercropping is known to have positive effects on crop productivity, it is unclear whether the effects of mixing species start at the early plant stage, that is, during germination. We tested whether the germination of two legume species, alsike clover and black medic, characterized by a contrasting response to water availability and temperature is affected by mixing. We set up four experiments in each of which we compared a 1:1 mixture against the two monocultures, and combined this with various other experimental factors. These additional factors were (i) varied seed densities (50%, 100% and 150% of a reference density) in two field trials in 2016 and 2017, (ii) varied seed densities (high and low) and water availability (six levels, between 25% and 100% of water holding capacity (WHC)) in a greenhouse pot trial, (iii) varied seed spacing in a climate chamber, and (iv) varied temperatures (12 °C, 20 °C and 28 °C) and water availability (four levels between 25% and 100% of WHC) in a climate chamber. Across all experiments, the absolute mixture effects (AME) on germination ranged between -9% and +11%, with a median of +1.3%. Within experiments, significant mixture effects were observed, but the direction of these effects was inconsistent. In the field, AME on germination was significantly negative at some of the tested seed densities. A positive AME was observed in the climate chamber at 12 °C, and the mean AME decreased with increasing temperature. Higher density was associated with decreased germination in the field, indicating negative interaction through competition or allelopathy, among seedlings. Our findings indicate that interaction among seeds in species mixtures may be ongoing during germination, but that the direction of the mixture effect is affected by complex interactions with abiotic and biotic factors.

Crop Resilience to Drought With and Without Response Diversity.

In the face of increasingly frequent droughts threatening crop performance, ecological theory suggests that higher species diversity may help buffering productivity by making systems more resistant through resource complementarity and more resilient through higher response diversity. However, empirical evidence for these diversity effects under drought stress has remained patchy. In two pot experiments, we explored whether mixing two legume species with a contrasting response to water availability, alsike clover (AC) and black medic (BM), promotes resistance to cumulative drought stress, and resilience of aboveground crop biomass to a transient drought event. The mixture was more productive than the average of the sole crops, and this mixture effect was higher in the non-stressed than in the drought-stressed plants. However, with six levels of constant drought intensities, the mixture effect was not consistently affected by drought level. Response diversity was evident as asynchrony of growth in the two species after the drought event, with BM re-growing faster than AC. Significant resilience to drought was observed in sole AC, i.e., without response diversity. Resilience was larger in AC than in BM and increased from 44 to 72 days after sowing (DAS). The mixture was more resilient than the average resilience of the sole crops at 72 DAS, but it was never more resilient than AC, indicating that resilience is promoted by, but not dependent on response diversity. We conclude that crop diversity may contribute to drought resilience through growth asynchrony, but that species identity plays a crucial role in making systems more drought-resilient.

Distinct communities of Cercozoa at different soil depths in a temperate agricultural field.

Protists are the most important predators of soil microbes like bacteria and fungi and are highly diverse in terrestrial ecosystems. However, the structure of protistan communities throughout the soil profile is still poorly explored. Here, we used Illumina sequencing to track differences in the relative abundance and diversity of Cercozoa, a major group of protists, at two depths; 10-30 cm (topsoil) and 60-75 cm (subsoil) in an agricultural field in Germany. At the two depths, we also distinguished among three soil compartments: rhizosphere, drilosphere (earthworm burrows) and bulk soil. With increasing depth, we found an overall decline in richness, but we were able to detect subsoil specific phylotypes and contrasting relative abundance patterns between topsoil and subsoil for different clades. We also found that the compartment effect disappeared in the subsoil when compared to the topsoil. More studies are now needed to describe and isolate these possibly subsoil specific phylotypes and better understand their ecology and function.

Bacteria utilizing plant-derived carbon in the rhizosphere of Triticum aestivum change in different depths of an arable soil.

Root exudates shape microbial communities at the plant-soil interface. Here we compared bacterial communities that utilize plant-derived carbon in the rhizosphere of wheat in different soil depths, including topsoil, as well as two subsoil layers up to 1 m depth. The experiment was performed in a greenhouse using soil monoliths with intact soil structure taken from an agricultural field. To identify bacteria utilizing plant-derived carbon, 13 C-CO2 labelling of plants was performed for two weeks at the EC50 stage, followed by isopycnic density gradient centrifugation of extracted DNA from the rhizosphere combined with 16S rRNA gene-based amplicon sequencing. Our findings suggest substantially different bacterial key players and interaction mechanisms between plants and bacteria utilizing plant-derived carbon in the rhizosphere of subsoils and topsoil. Among the three soil depths, clear differences were found in 13 C enrichment pattern across abundant operational taxonomic units (OTUs). Whereas, OTUs linked to Proteobacteria were enriched in 13 C mainly in the topsoil, in both subsoil layers OTUs related to Cohnella, Paenibacillus, Flavobacterium showed a clear 13 C signal, indicating an important, so far overseen role of Firmicutes and Bacteriodetes in the subsoil rhizosphere.

Prokaryotes in Subsoil-Evidence for a Strong Spatial Separation of Different Phyla by Analysing Co-occurrence Networks.

Microbial communities in soil provide a wide range of ecosystem services. On the small scale, nutrient rich hotspots in soil developed from the activities of animals or plants are important drivers for the composition of microbial communities and their functional patterns. However, in subsoil, the spatial heterogeneity of microbes with differing lifestyles has been rarely considered so far. In this study, the phylogenetic composition of the bacterial and archaeal microbiome based on 16S rRNA gene pyrosequencing was investigated in the soil compartments bulk soil, drilosphere, and rhizosphere in top- and in the subsoil of an agricultural field. With co-occurrence network analysis, the spatial separation of typically oligotrophic and copiotrophic microbes was assessed. Four bacterial clusters were identified and attributed to bulk topsoil, bulk subsoil, drilosphere, and rhizosphere. The bacterial phyla Proteobacteria and Bacteroidetes, representing mostly copiotrophic bacteria, were affiliated mainly to the rhizosphere and drilosphere-both in topsoil and subsoil. Acidobacteria, Actinobacteria, Gemmatimonadetes, Planctomycetes, and Verrucomicrobia, bacterial phyla which harbor many oligotrophic bacteria, were the most abundant groups in bulk subsoil. The bacterial core microbiome in this soil was estimated to cover 7.6% of the bacterial sequencing reads including both oligotrophic and copiotrophic bacteria. In contrast the archaeal core microbiome includes 56% of the overall archaeal diversity. Thus, the spatial variability of nutrient quality and quantity strongly shapes the bacterial community composition and their interaction in subsoil, whereas archaea build a stable backbone of the soil prokaryotes due to their low variability in the different soil compartments.

Biochar affected by composting with farmyard manure.

Biochar applications to soils can improve soil fertility by increasing the soil's cation exchange capacity (CEC) and nutrient retention. Because biochar amendment may occur with the applications of organic fertilizers, we tested to which extent composting with farmyard manure increases CEC and nutrient content of charcoal and gasification coke. Both types of biochar absorbed leachate generated during the composting process. As a result, the moisture content of gasification coke increased from 0.02 to 0.94 g g, and that of charcoal increased from 0.03 to 0.52 g g. With the leachate, the chars absorbed organic matter and nutrients, increasing contents of water-extractable organic carbon (gasification coke: from 0.09 to 7.00 g kg; charcoal: from 0.03 to 3.52 g kg), total soluble nitrogen (gasification coke: from not detected to 705.5 mg kg; charcoal: from 3.2 to 377.2 mg kg), plant-available phosphorus (gasification coke: from 351 to 635 mg kg; charcoal: from 44 to 190 mg kg), and plant-available potassium (gasification coke: from 6.0 to 15.3 g kg; charcoal: from 0.6 to 8.5 g kg). The potential CEC increased from 22.4 to 88.6 mmol kg for the gasification coke and from 20.8 to 39.0 mmol kg for the charcoal. There were little if any changes in the contents and patterns of benzene polycarboxylic acids of the biochars, suggesting that degradation of black carbon during the composting process was negligible. The surface area of the biochars declined during the composting process due to the clogging of micropores by sorbed compost-derived materials. Interactions with composting substrate thus enhance the nutrient loads but alter the surface properties of biochars.