In the context of a recent massive increase in research on plant root functions and their impact on the environment, root ecologists currently face many important challenges to keep on generating cutting-edge, meaningful and integrated knowledge. Consideration of the below-ground components in plant and ecosystem studies has been consistently called for in recent decades, but methodology is disparate and sometimes inappropriate. This handbook, based on the collective effort of a large team of experts, will improve trait comparisons across studies and integration of information across databases by providing standardised methods and controlled vocabularies. It is meant to be used not only as starting point by students and scientists who desire working on below-ground ecosystems, but also by experts for consolidating and broadening their views on multiple aspects of root ecology. Beyond the classical compilation of measurement protocols, we have synthesised recommendations from the literature to provide key background knowledge useful for: (1) defining below-ground plant entities and giving keys for their meaningful dissection, classification and naming beyond the classical fine-root vs coarse-root approach; (2) considering the specificity of root research to produce sound laboratory and field data; (3) describing typical, but overlooked steps for studying roots (e.g. root handling, cleaning and storage); and (4) gathering metadata necessary for the interpretation of results and their reuse. Most importantly, all root traits have been introduced with some degree of ecological context that will be a foundation for understanding their ecological meaning, their typical use and uncertainties, and some methodological and conceptual perspectives for future research. Considering all of this, we urge readers not to solely extract protocol recommendations for trait measurements from this work, but to take a moment to read and reflect on the extensive information contained in this broader guide to root ecology, including sections I-VII and the many introductions to each section and root trait description. Finally, it is critical to understand that a major aim of this guide is to help break down barriers between the many subdisciplines of root ecology and ecophysiology, broaden researchers' views on the multiple aspects of root study and create favourable conditions for the inception of comprehensive experiments on the role of roots in plant and ecosystem functioning.

https://hal.archives-ouvertes.fr/hal-02918834/document

A starting guide to root ecology: strengthening ecological concepts and standardising root classification, sampling, processing and trait measurements

The 4‰ initiative launched by the French government at COP21 in Paris in December 2015 aspires to increase global soil organic carbon (SOC) stocks at a rate of 0.4% per year. We conducted a systematic literature review on SOC storage under agroforestry and conservation agriculture systems in sub-Saharan Africa, where we reported 66 and 33 cases for both systems respectively. The results showed that SOC storage rates were significantly higher than 4‰ yr−1 in fallows and in multistrata agroforestry systems (P = 0.0001 and 0.0178, respectively), but not in alley cropping and parklands systems. For conservation agriculture, SOC storage rates were only significantly higher than 4‰ yr−1 (P = 0.0438) when all three principles were applied, i.e. no- or minimum tillage combined with crop residue retention and intercropping or rotation. The data showed very large variability in SOC storage rates as the result of various factors, including previous land-use history, experimental set up and approach used to determine SOC storage (diachronic versus synchronic approach), soil type, depth of soil sampling, type of crops and management, and duration of the experiment. SOC storage rates significantly decreased with time in the agroforestry systems (P = 0.0328). However, we were unable to find significant relationships with initial SOC stocks or tree density. Given the limited published data and the high variability in results, no significant relationships between SOC storage rates and site variables were found for conservation agriculture. We argue that there is a potential for SOC storage in agricultural soils of sub-Saharan Africa, as illustrated by SOC gaps observed on smallholder farms. Low SOC levels are, however, to a great extent the result of limited resources of most smallholder farmers. Practices such as agroforestry and conservation agriculture can restore SOC in these soils, but the 4‰ initiative has to be implemented on the grounds of the positive impact on crop productivity rather than on climate change mitigation. The efficiency in doing so will depend on the specific situations and will need economic support to smallholder farmers, including the promotion of good markets for sale of extra produce and for input supply, effective private support and policy, such as credit schemes and subsidies for inputs, and efficient extension services which incentivize farmers to invest in new technologies.

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The 4 per 1000 goal and soil carbon storage under agroforestry and conservation agriculture systems in sub-Saharan Africa

Most studies on plant nutrition tend to focus on the topsoil (plough layer) and frequently neglect subsoil processes. However, cereal roots can potentially acquire nutrients including organic and inorganic nitrogen (N) from deep in the soil profile. Greater knowledge on the interaction of plants and microbes in subsoil environments is required to evaluate whether deep rooting traits in cereals will achieve greater nutrient use efficiency and greater soil carbon (C) storage in cropping systems. This study aimed to evaluate the relationship between root distribution, organic and inorganic N availability and potential N supply at the critical growth period during the wheat cropping cycle in a sand textured Eutric Cambisol. Our results provide evidence of significant microbial capacity in the subsoil. The rate of plant residue turnover and the mineralization of organic C and N substrates (glucose, amino acids, peptides, protein) declined slightly with increasing soil depth; however, these rates were not correlated with basal soil respiration, microbial biomass or community structure. This suggests that the microbial population in subsoil is more C limited but that its activity can be readily stimulated upon C substrate addition. A significant potential for organic and inorganic N turnover was also demonstrated at depth with a similar abundance of ammonifiers and ammonia oxidizing bacteria (AOB) and archaea (AOA) throughout the soil profile. Again, N mineralization in subsoils appears to be substrate limited. Root density declined rapidly down the soil profile with few roots present past 50 cm; suggesting that this is the major factor limiting C recharge of soil organic matter and microbial activity in subsoils. Greater root proliferation at depth could allow greater capture of water and the recapture of N lost by leaching; however, our results suggest that plant-microbial competition for C and N is as intense in the subsoil as in the topsoil. We conclude that while deeper rooting may improve nutrient and water use efficiency it may not lead to much greater C sequestration in subsoils, at least in the short term.

/pdf/microbial-competition-for-nitrogen-and-carbon-is-as-intense-puvu2lylfd.pdf

Microbial competition for nitrogen and carbon is as intense in the subsoil as in the topsoil

Digging Deeper for Agricultural Resources, the Value of Deep Rooting

Extreme drought is likely to become more frequent and intense as a result of global climate change, which may significantly impact plant root traits and responses (i.e., morphology, production, turnover, and biomass). However, a comprehensive understanding of how drought affects root traits and responses remains elusive. Here, we synthesized data from 128 published studies under field conditions to examine the responses of 17 variables associated with root traits to drought. Our results showed that drought significantly decreased root length and root length density by 38.29% and 11.12%, respectively, but increased root diameter by 3.49%. However, drought significantly increased root:shoot mass ratio and root cortical aerenchyma by 13.54% and 90.7%, respectively. Our results suggest that drought significantly modified root morphological traits and increased root mortality, and the drought-induced decrease in root biomass was less than shoot biomass, causing higher root:shoot mass ratio. The cascading effects of drought on root traits and responses may need to be incorporated into terrestrial biosphere models to improve prediction of the climate-biosphere feedback.

Drought-induced changes in root biomass largely result from altered root morphological traits: Evidence from a synthesis of global field trials

Understanding deep roots and their functions in ecosystems: an advocacy for more unconventional research

Fine root dynamics is a main driver of soil carbon stocks, particularly in tropical forests, yet major uncertainties still surround estimates of fine root production and turnover. This lack of knowledge is largely due to the fact that studying root dynamics in situ, particularly deep in the soil, remains highly challenging. We explored the interactions between fine root dynamics, soil depth and rainfall in mature rubber trees (Hevea brasiliensis Mull. Arg.) exposed to sub-optimal edaphic and climatic conditions. A root observation access well was installed in northern Thailand to monitor root dynamics along a 4.5 m deep soil profile. Image-based measurements of root elongation and lifespan of individual roots were carried out at monthly intervals over 3 years. Soil depth was found to have a significant effect on root turnover. Surprisingly, root turnover increased with soil depth and root half-life was 16, 6 - 8 and only 4 months at 0.5, 1.0, 2.5 and 3.0 m deep, respectively (with the exception of roots at 4.5 m which had a half-life similar to that found between depths of 1.0 and 2.5 m). Within the first two meters of the soil profile, the highest rates of root emergence occurred about three months after the onset of the rainy season, while deeper in the soil, root emergence was not linked to the rainfall pattern. Root emergence was limited during leaf flushing (between March and May), particularly within the first two meters of the profile. Between soil depths of 0.5 and 2.0 m, root mortality appeared independent of variations in root emergence, but below 2.0 m, peaks in root emergence and death were synchronized. Shallow parts of the root were more responsive to rainfall than their deeper counterparts. Increased root emergence in deep soil towards the onset of the dry season could correspond to a drought acclimation mechanism, with the relative importance of deep water capture increasing once rainfall ceased. The considerable soil depth regularly explored by fine roots, even though significantly less than in surface layers in terms of root length density and biomass, will impact strongly the evaluation of soil carbon stocks

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Seasonal Patterns of Fine Root Production and Turnover in a Mature Rubber Tree (Hevea brasiliensis Müll. Arg.) Stand- Differentiation with Soil Depth and Implications for Soil Carbon Stocks.

Corentin Clément

Papers

Understanding deep roots and their functions in ecosystems: an advocacy for more unconventional research

Seasonal Patterns of Fine Root Production and Turnover in a Mature Rubber Tree (Hevea brasiliensis Müll. Arg.) Stand- Differentiation with Soil Depth and Implications for Soil Carbon Stocks.

Root and xylem anatomy varies with root length, root order, soil depth, and environment in intermediate wheatgrass (Kernza®) and alfalfa.