Soil amendment with biochar is evaluated globally as a means to improve soil fertility and to mitigate climate change. However, the effects of biochar on soil biota have received much less attention than its effects on soil chemical properties. A review of the literature reveals a significant number of early studies on biochar-type materials as soil amendments either for managing pathogens, as inoculant carriers or for manipulative experiments to sorb signaling compounds or toxins. However, no studies exist in the soil biologyliterature that recognize the observed largevariations ofbiochar physico-chemical properties. This shortcoming has hampered insight into mechanisms by which biochar influences soil microorganisms, fauna and plant roots. Additional factors limiting meaningful interpretation of many datasets are the clearly demonstrated sorption properties that interfere with standard extraction procedures for soil microbial biomass or enzyme assays, and the confounding effects of varying amounts of minerals. In most studies, microbial biomass has been found to increase as a result of biochar additions, with significant changes in microbial community composition and enzyme activities that may explain biogeochemical effects of biochar on element cycles, plant pathogens, and crop growth. Yet, very little is known about the mechanisms through which biochar affects microbial abundance and community composition. The effects of biochar on soil fauna are even less understood than its effects on microorganisms, apart from several notable studies on earthworms. It is clear, however, that sorption phenomena, pH and physical properties of biochars such as pore structure, surface area and mineral matter play important roles in determining how different biochars affect soil biota. Observations on microbial dynamics lead to the conclusion of a possible improved resource use due to co-location of various resources in and around biochars. Sorption and therebyinactivation of growth-inhibiting substances likelyplaysa rolefor increased abundance of soil biota. No evidence exists so far for direct negative effects of biochars on plant roots. Occasionally observed decreases in abundance of mycorrhizal fungi are likely caused by concomitant increases in nutrient availability,reducing theneedfor symbionts.Inthe shortterm,therelease ofavarietyoforganic molecules from fresh biochar may in some cases be responsible for increases or decreases in abundance and activity of soil biota. A road map for future biochar research must include a systematic appreciation of different biochar-types and basic manipulative experiments that unambiguously identify the interactions between biochar and soil biota.

http://www.css.cornell.edu/faculty/lehmann/pictures/publ/SoilBiolBiochem%2043,%201812–1836,%202011%20Lehmann.pdf

Biochar effects on soil biota – A review

Natural organic biomass burning creates black carbon which forms a considerable proportion of the soil’s organic carbon. Due to black carbon’s aromatic structure it is recalcitrant and has the potential for long-term carbon sequestration in soil. Soils within the Amazon-basin contain numerous sites where the ‘dark earth of the Indians’ (Terra preta de Indio, or Amazonian Dark Earths (ADE)) exist and are composed of variable quantities of highly stable organic black carbon waste (‘biochar’). The apparent high agronomic fertility of these sites, relative to tropical soils in general, has attracted interest. Biochars can be produced by ‘baking’ organic matter under low oxygen (‘pyrolysis’). The quantities of key mineral elements within these biochars can be directly related to the levels of these components in the feedstock prior to burning. Their incorporation in soils influences soil structure, texture, porosity, particle size distribution and density. The molecular structure of biochars shows a high degree of chemical and microbial stability. A key physical feature of most biochars is their highly porous structure and large surface area. This structure can provide refugia for beneficial soil micro-organisms such as mycorrhizae and bacteria, and influences the binding of important nutritive cations and anions. This binding can enhance the availability of macro-nutrients such as N and P. Other biochar soil changes include alkalisation of soil pH and increases in electrical conductivity (EC) and cation exchange capacity (CEC). Ammonium leaching has been shown to be reduced, along with N2O soil emissions. There may also be reductions in soil mechanical impedance. Terra preta soils contain a higher number of ‘operational taxonomic units’ and have highly distinctive microbial communities relative to neighbouring soils. The potential importance of biochar soil incorporation on mycorrhizal fungi has also been noted with biochar providing a physical niche devoid of fungal grazers. Improvements in soil field capacity have been recorded upon biochar additions. Evidence shows that bioavailability and plant uptake of key nutrients increases in response to biochar application, particularly when in the presence of added nutrients. Depending on the quantity of biochar added to soil significant improvements in plant productivity have been achieved, but these reports derive predominantly from studies in the tropics. As yet there is limited critical analysis of possible agricultural impacts of biochar application in temperate regions, nor on the likelihood of utilising such soils as long-term sites for carbon sequestration. This review aims to determine the extent to which inferences of experience mostly from tropical regions could be extrapolated to temperate soils and to suggest areas requiring study.

Potential mechanisms for achieving agricultural benefits from biochar application to temperate soils: a review

Biochar is a carbon-rich coproduct resulting from pyrolyzing biomass. When applied to the soil it resists decomposition, effectively sequestering the applied carbon and mitigating anthropogenic CO2 emissions. Other promoted benefits of biochar application to soil include increased plant productivity and reduced nutrient leaching. However, the effects of biochar are variable and it remains unclear if recent enthusiasm can be justified. We evaluate ecosystem responses to biochar application with a meta-analysis of 371 independent studies culled from 114 published manuscripts. We find that despite variability introduced by soil and climate, the addition of biochar to soils resulted, on average, in increased aboveground productivity, crop yield, soil microbial biomass, rhizobia nodulation, plant K tissue concentration, soil phosphorus (P), soil potassium (K), total soil nitrogen (N), and total soil carbon (C) compared with control conditions. Soil pH also tended to increase, becoming less acidic, following the addition of biochar. Variables that showed no significant mean response to biochar included belowground productivity, the ratio of aboveground : belowground biomass, mycorrhizal colonization of roots, plant tissue N, and soil P concentration, and soil inorganic N. Additional analyses found no detectable relationship between the amount of biochar added and aboveground productivity. Our results provide the first quantitative review of the effects of biochar on multiple ecosystem functions and the central tendencies suggest that biochar holds promise in being a win-win-win solution to energy, carbon storage, and ecosystem function. However, biochar's impacts on a fourth component, the downstream nontarget environments, remain unknown and present a critical research gap.

https://www.onlinelibrary.wiley.com/doi/pdf/10.1111/gcbb.12037

Biochar and its effects on plant productivity and nutrient cycling: a meta‐analysis

Experiments suggest that biomass-derived black carbon (biochar) affects microbial populations and soil biogeochemistry. Both biochar and mycor- rhizal associations, ubiquitous symbioses in terrestrial ecosystems, are potentially important in various ecosystem services provided by soils, contributing to sustainable plant production, ecosystem restoration, and soil carbon sequestration and hence mitigation of global climate change. As both biochar and mycor- rhizal associations are subject to management, under- standing and exploiting interactions between them could be advantageous. Here we focus on biochar effects on mycorrhizal associations. After reviewing the experimental evidence for such effects, we critically examine hypotheses pertaining to four mechanisms by which biochar could influence mycorrhizal abundance and/or functioning. These mechanisms are (in decreas- ing order of currently available evidence supporting them): (a) alteration of soil physico-chemical proper- ties; (b) indirect effects on mycorrhizae through effects on other soil microbes; (c) plant-fungus signaling interference and detoxification of allelochemicals on biochar; and (d) provision of refugia from fungal grazers. We provide a roadmap for research aimed at testing these mechanistic hypotheses.

http://www.css.cornell.edu/faculty/lehmann/publ/PlantSoil%20300,%209-20,%202007,%20Warnock.pdf

Mycorrhizal responses to biochar in soil – concepts and mechanisms

Arbuscular mycorrhizal fungi and organic farming

Seedlings of asparagus (Asparagus officinalis L., cv. Mary Washington 500 W) were inoculated with arbuscular mycorrhizal (AM) fungus (Glomus sp. R10) and grown in the presence of different soil amendments : carbonized chaff (CC), coconut charcoal (CO) and manure of coffee residue (MC) mixed with the soil in the ratio of 1 : 9 (v/v, 10% plot) or 3 : 7 (30% plot). Nine weeks later, the seedlings were exposed to fusarium root rot, caused by Fusarium oxysporum f. sp. asparagi (Foa) to test their tolerance. When asparagus seedlings were examined 13 weeks after Foa inoculation, incidence of fusarium root rot reached 90-100% in all the AM fungus-noninoculated plants, irrespective of the soil amendments. However, the incidences of root rot were 14.3% in the 30% CO and 20% in the 10% MC plots among the AM fungus-inoculated plants. In the non-AM inoculated plants, the disease indices reached 92.5 in the soil only plot and 48.9 in the 30% CC plot. As for AM-treated plants, the indices attained remarkably low values of 2.9 in the 30% CO and 4.0 in the 10% MC plots. More storage roots developed in the AM-treated plants than in the control, whether they were diseased or healthy. AM fungal infection levels were higher in the root systems of 10% and 30% CO and 10% MC plots than in the other plots ; little difference occurred among 10% and 30% CO and 10% MC plots. Furthermore, the infection area of the AM fungus in a root system was most extensive in the CO plots. These findings suggest that the addition of coconut charcoal or manure of coffee residue to bed soil is effective for increasing the tolerance to fusarium root rot in AM fungus-infected asparagus plants, though the effect differed with the quantity of the soil amendments.

Incidence of Fusarium Root Rot in Asparagus Seedlings Infected with Arbuscular Mycorrhizal Fungus as Affected by Several Soil Amendments

Root and stem rot of cut-flower roses (Rosa spp.) was observed in commercial glasshouse-grown roses in 10 prefectures of Japan from 1998 through 2001. Binucleate-like Rhizoctonia spp. were isolated mainly from the disease plants. In all, 670 isolates were divided into two types based on cultural appearance; 168 isolates of light brown to brown type and 502 isolates of whitish type. A hyphal anastomosis reaction using representative isolates from each type revealed that the light brown to brown type belonged to anastomosis group G (AG-G), whereas the whitish type (AG-CUT) failed to anastomose with tester strains of binucleate Rhizoctonia AG-A through AG-S. Neither isolates of AG-G nor AG-CUT anastomosed with tester strains of a previously reported unknown AG (AG-MIN) of binucleate Rhizoctonia spp. collected from miniature roses. In pathogenicity tests, randomly selected isolates of the three groups caused root and stem rot on cut-flower and miniature roses. To differentiate AG-CUT and AG-MIN from known AGs of binucleate Rhizoctonia spp., restriction fragment length polymorphism (RFLP) and sequence analyses of a ribosomal (r)DNA internal transcribed spacer (ITS) region were conducted. Among the eight restriction enzymes used, HaeIII produced DNA banding patterns for AG-CUT that differed from those of tester strains and AG-MIN. Additionally, restriction profiles of AG-MIN differed from those of all tester strains. AG-G isolates from cut-flower roses had the same RFLP pattern as the tester strains of AG-G. Based on the results of hyphal anastomosis and RFLP and sequence analysis of an rDNA-ITS region, we propose that AG-CUT be designated AG-T and AG-MIN be designated AG-U, two new AGs of binucleate Rhizoctonia spp. The phylogenetic tree based on the sequence data of the rDNA-ITS region showed that isolates of AG-MIN were in a distinct clade from other AGs, whereas isolates of AG-CUT were in the same clade as those of AG-A. More detailed phylogenetic analysis besides rDNA-ITS region might be necessary for AG classification of binucleate Rhizoctonia spp.

https://apsjournals.apsnet.org/doi/pdf/10.1094/PHYTO-95-0784

New Anastomosis Groups, AG-T and AG-U, of Binucleate Rhizoctonia spp. Causing Root and Stem Rot of Cut-Flower and Miniature Roses

Effect of Arbuscular Mycorrhizal Fungus Infection on the Incidence of Fusarium Root Rot in Asparagus Seedlings

Phylogenetic analyses of the genus Rosa using the matK sequence : molecular evidence for the narrow genetic background of modern roses

The use of heavy ion beams is an effective method for inducing mutations in plants. After irradiation with beams of Nitrogen (N) or Neon (Ne), the frequency of flower color mutants increased through the regeneration of stem or leaf without lateral meristem in an interspecific hybrid of Torenia. From anthocyanin assay results, mutants were divided into two groups. One group involved the deletion of the blue gene (DFR). The other group involved the deletion or duplication of a gene related to pigment production. The results show that heavy ion beams are effective in obtaining artificial mutants.

Hirokazu Fukui

Papers

Incidence of Fusarium Root Rot in Asparagus Seedlings Infected with Arbuscular Mycorrhizal Fungus as Affected by Several Soil Amendments

New Anastomosis Groups, AG-T and AG-U, of Binucleate Rhizoctonia spp. Causing Root and Stem Rot of Cut-Flower and Miniature Roses

Effect of Arbuscular Mycorrhizal Fungus Infection on the Incidence of Fusarium Root Rot in Asparagus Seedlings

Phylogenetic analyses of the genus Rosa using the matK sequence : molecular evidence for the narrow genetic background of modern roses

Flower pigment mutations induced by heavy ion beam irradiation in an interspecific hybrid of Torenia