Arbuscular mycorrhizal fungi in biochar-amended soils: a review
Jonna Rosenthal
A * and Kpoti M. Gunn B *
A
B
Handling Editor: Leo Condron
Abstract
Biochar has been a centre of advancement in agricultural and chemical technologies research in the past decade due to its effectiveness as a sustainable fertilizer and toxicological absorbent. However, this research has failed to address the variety of biochar’s effects on soil ecosystems. Due to this oversight, the impacts of biochar amendments on arbuscular mycorrhizal fungi (AMF) are largely overlooked. This work reviews a multitude of sources in an attempt to summarise current research surrounding biochar’s positive impacts on the ecological functioning of AMF and the implications for sustainable agricultural advancements. Findings vary in applicability; however, baseline trends show that infrequent biochar application leads to higher AMF root colonisation rates. Furthermore, biochar may also increase AMF biodiversity as well as spore count. This review is the starting point for the analysis of biochar on non-target soil microorganisms. Due to the current gaps in the literature, future research should focus on standardizing pyrolysis and biochar application methodology to advance potential positive applications of AMF to agricultural cultivations.
Keywords: adsorption, agricultural improvements, arbuscular mycorrhizal fungi, biochar, drought mitigation, field applications, nutrient uptake, soil conservation.
Introduction
Biochar is a carbon-based product of the pyrolytic transformation of organic materials (biomass). It has become a fairly common soil amendment in cultivated areas, contributing to soil health improvement, carbon sequestration, and contaminant remediation (Soltes and Elder 2018; Zama et al. 2018; Allohverdi et al. 2021; Gross et al. 2021; Ji et al. 2022). Pyrolysis turns biomass into a network of aromatic, aliphatic, and graphitic carbon molecules with functional groups, that more or less resist degradation or leaching (Lian and Xing 2017; Leng et al. 2019), and therefore represents an important carbon sink pool in the soil environment (Lal 2004; Allohverdi et al. 2021; Werner et al. 2022). Biochar is produced from a range of biomass materials (type and size of feedstock) and under various conditions (temperature and residence time), leading to changing behaviour in the soil environment as well as diverse biotic and abiotic interactions with soil materials (Joseph et al. 2021). Additionally, variability in land application (time of year, application rate, crop land use) has been found to enhance biochar’s changing behaviour (Lehmann et al. 2021; Amalina et al. 2022; Panwar and Pawar 2022). Due to the significant level of variability in biochar characteristics, recent studies have focused on optimising specific fabrication conditions that could lead to the most effective treatment for specific uses, though little consensus has been found (Wan et al. 2020).
Arbuscular mycorrhizal fungi (AMF) are a group of soil microorganisms that form symbiotic relationships with the roots of most plants, enhancing the critical supply of water and nutrients (nitrogen and phosphorus) in return for fixed carbon from the host plants. AMF develop a mycelium network that bridge plant roots with the surrounding soil microhabitats and penetrate the cortical cells of the roots to form arbuscules that expand the zone of capture of the root system (Simard et al. 2012; Luginbuehl and Oldroyd 2017). Fig. 1 shows the difference in AMF growth between a colonised and uncolonised plant, and the contribution of AMF arbuscules to the enhancement of host plant–soil environmental exchanges. Additionally to their known value in supporting water supply and nutrient cycling, AMF have been found to enhance plants’ resistance to various abiotic stresses, including drought, salinity, inhibiting metals, and extreme temperatures, which makes them important for sustainable agriculture and ecosystem health, especially in the context of adaptation to climate change impacts (Begum et al. 2019).
Growth of arbuscular mycorrhizal fungi (AMF) in the root zone and their effects on characteristic qualitative variables of soils. Differences are shown between uncolonised and soils colonised with AMF. Reprinted from ‘Trade-offs in Arbuscular Mycorrhizal Symbiosis: disease resistance, growth responses and perspectives for crop breeding’ by Jacott, C. N., Murray, J. D., and Ridout, C. J. 2017, Agronomy, 7(4), 75. Copyright 2017 under the Creative Commons Attribution (CC BY) licence. Reprinted with permission of the authors.
Biochar has been shown to impact soil microbiology, including AMF (Jacott et al. 2017). Interactions between biochar and AMF in the soil environment have been found to range from beneficial to detrimental, likely related to chemical and structural characteristics of the pyrolysed feedstock (Joseph et al. 2021), the pyrolysis conditions, or the biotic and abiotic conditions of the soil environment. For example, the molecular structure and carbon content of biochar, affected by feedstock sources and pyrolysis conditions (Askeland et al. 2019), create conditions that could affect AMF abundance but decrease AMF composition (Neuberger et al. 2024). While reviews of the effects of biochar applications on soil characteristics, nutrient and contaminant movements, and plants have been extensively explored in the literature (Joseph et al. 2021), reviews related to the effects of biochar on AMF are limited, as only a few authors explore the mechanistic relations. While an increasing number of scientists have noted the positive effects of biochar on AMF colonisation in specific crops, documentation of conclusive findings is limited regarding the factors involved in AMF reproduction and infective potential. Therefore, given the increasing use of biochar as a soil amendment, it is imperative to expand on the correlative relationship between biochar amendments and AMF successes. This review summarises past literature detailing biochar’s impacts on AMF root colonisation, spore activity, infective potential, biodiversity, and group complexity. In particular, this review highlights the relationship between biochar amendments, AMF microbial potential, soil water-holding capacity, and crop production. Though other papers have focused on the impacts of biochar on overall soil biota or the impacts of biochar in conjunction with AMF, this review is unique in its focus on effects of biochar on AMF abundance and productivity. Findings will contribute to highlighting recommendations for future research into biochar fabrication conditions that support soil microbiological improvements.
Mechanistic interactions between biochar and AMF
Biochar is a solid material with a complex variable elemental composition, primarily containing carbon (C), hydrogen (H), and oxygen (O) at various H/C and O/C fractions, and smaller amounts of other elements like nitrogen, phosphorus, sulfur, calcium, magnesium, potassium, and sodium, depending on the types and sources of feedstock, and the pyrolysis conditions (Askeland et al. 2019; Wijitkosum and Jiwnok 2019). The major biochar elements (C, H, and O) are organised into a carbon-rich porous skeleton consisting mostly of aromatic rings and aliphatic chains, combined into layers, which may or may not be combined or stacked into graphitic structures (Fig. 2a). C, H, and O also contribute to the formation of hydroxyl, carbonyl/xyl, ester functional groups attached to the basic skeleton, which affect biochar properties (Fig. 2b). The elements listed above in smaller amounts are part of the biochar molecule as single elements attached directly to the carbon skeleton or through the functional groups.
(a) Biochar skeleton surface image showing cyclic aromatic rings as viewed with a scanning electron microscope (adapted from Askeland et al. (2019)). (b) Biochar aromatic ring layer example with functional groups (from Wood et al. (2024)).
The variability in elemental composition, skeletal structure, and functional groups makes biochar a material that demonstrates a large range of characteristics that may strongly affect its physicochemical and biologic behaviour in the environment (Tomczyk et al. 2020). Some of these characteristics include notably specific surface area (SSA), surface functional groups, porosity, cation exchange capacity (CEC), and pH. Biochar application to soils can promote an increase in SSA, functional groups, porosity, and CEC. Increase and change of functional groups in the soil environment can potentially lead to the formation and redistribution of soil microaggregates (Tomczyk et al. 2020). Soil aggregation has widely been known to improve soil physical characteristics that promote stability, aeration, and water retention (Halder et al. 2024), therefore creating and enhancing conditions for microbiological activities. Similar to the impact of functional groups alteration, increased SSA, CEC, and porosity due to biochar addition may affect not only aggregate formations in the soil, but also the amount, connectivity, and size of pores (i.e. macropores) in the soil environment. Changing macropore conditions in the soil environment may interfere (positively or negatively) with air and moisture content and movement, as well as the space available for microbiological population growth.
When biochar is added to the soil by broadcast and tillage, trench incorporation, or combined with fertiliser (biochar compound fertiliser), it absorbs water into its pore network, which contributes in the dissolution and release of chemical species, including soluble organic and mineral molecules located on the surface of the biochar particles (Joseph et al. 2021). This initial release of biochar-held compounds contributes to changes in the chemical composition of the soil solution, hence in soil pH and CEC, leading to increased development of fungal spores in the soil environment. Schreiter et al. (2020) found that the release of compounds from the biochar surface allows an increase in the biochar’s surface area and porosity, creating the opportunity for cations, anions, nutrients, and minerals storage. Such compound storage on the surface of biochar tends to promote an increase in microbiological activity near biochar particles. Simard et al. (2012) found that AMF in particular grows a mycelium network within the pores of biochar particles in soils poor in P, which allows plants grown in low P soils to take advantage of the P stored in the biochar interstices. To summarise, AMF and biochar particles do not interact directly, but biochar operates like a storage medium that helps hold nutrients in the soil environment until accessed by plants via root extensions created by the mycelium network (Neuberger et al. 2024). Consequently, the higher the density of biochar particles conditioned to hold available nutrients is in the soil, the more AMF mycelium would grow to access these nutrients. Note that the linearity of the relation between AMF activity and biochar density has been challenged by some authors and may not hold at high biochar rate application (Joseph et al. 2021).
Positive effects of biochar on AMF
AMF root colonisation
AMF enhances nutrient and water availability and quality, and soil structural integrity (Wright 2005) that directly affect plant physiology and yield. Notably, Chen et al. (2018) cites 231 field trials that averaged 9.5% increased total yield for crops with AMF root colonisation as compared to AMF-absent crops. It is therefore expected that strong AMF root colonisation will lead to higher plant productivity and health, which provides a strong imperative to further study this fungus. While past literature has focused on crop health concerning AMF, recent studies have shown that biochar addition to soils has corresponded with significant changes in AMF root colonisation (Warnock et al. 2007) (Table 1). The mechanisms proposed to explain these changes in colonisation of AMF include additional root surface area for growth, enhanced nutrient profiles of soil, and increased refuge for spores, all due to biochar addition (Neuberger et al. 2024). In their original research on the effects of mallee biochar (rates of 0, 1.5, 3.0, and 6 t ha−1) on wheat (Triticum aestivum) growth in Western Australia, Solaiman et al. (2010a) found that additional colonisation of AMF resulted in an increased nutrient uptake, (especially phosphorus) for surrounding plants and microorganisms. An additional study conducted using a twin chamber system with woody biochar and harvesting dates 35, 49, and 63 days after planting, noted a 47% increase in AMF root colonisation on clover plants grown in biochar-amended soils (Mickan et al. 2016). Roots of wheat, soy (Glycine max), corn (Zea mays), and other legumes were reported to show higher rates of AMF colonisation under biochar-amended soils, However, the literature seems to gloss over the significance of this observation, following past research that does not focus on soil microbiology in remediation studies (Solaiman et al. 2010b; Videgain-Marco et al. 2021). Considerable improvements in soil and plant conditions are observed under biochar conditions, which could be due in part to AMF contributions, necessitating researchers to highlight recent AMF colonisation findings to inspire further research. For this reason, it is imperative that future research considers AMF root colonisation as a key variable in biochar-related research.
| Experimental treatment | BC feedstock | BC quantity | Effect A | Plant response to AM | Source | |
|---|---|---|---|---|---|---|
| AMF reactions to BC applications and mechanisms | Mallee trees | 50–200 mm depth on plot | + | 18–46% increase in colonisation activity | Solaiman et al. (2010a) | |
| BC impact on AMF under water-stressed conditions | Jarrah wood | BC at 2% w/w | + | 47% increase in root colonisation | Mickan et al. (2016) | |
| AMF colonisation under different BC treatments | Wood chips, fibre sludge, and grain husks | 0, 0.5%, 2.5%, and 5.0% w/w | 0 | Higher co-dependency between AMF and soil parameters | Barna et al. (2020) | |
| BC effect on AMF production | Orchard prunings | 10 t ha−1 | 0 | No difference in measured biomass | Amendola et al. (2017) | |
| BC effect on AMF community structure | Forest logging residues | Not listed | + | Increased community complexity and biodiversity | Yan et al. (2021) | |
| Legacy effects of BC and compost on AMF | Rice straw | 10 t ha−1 | + | Increased biodiversity and biomass | Xin et al. (2022) | |
| BC effects on AMF in agro environments | Vine shoots | 1.5% w/w, 3.0% w/w | + | Increased AMF root colonisation, spores, and infective potential | Videgain-Marco et al. (2021) | |
| BC effects on AMF in aqueous crops | Jarrah wood | Not listed | + | Higher AMF colonisation | Solaiman et al. (2010b) | |
| BC and AMF effects under saline-alkali stress | Corn stalks | 0%, 2%, 5%, and 10% w/w | Increased AMF infection and biodiversity | Wen et al. (2024) | ||
| BC aiding AMF colonisation in secondary metabolite production | Straw | (1%, 2%, 3% v/v) | Increased AMF colonisation | Chen et al. (2025) | ||
| BC and AMF for phosphorus cycling in paddy soils | Rice husks | Not listed | Increased AMF spore distribution and colonisation | Chen et al. (2024) |
AMF spore activity
Fungi primarily reproduce through aerial dispersion. The spores of AMF are thick-walled, multinucleate structures. Although not dependent on host plants for dispersal, spores can be influenced by their symbiotic relationship with the host plant (Wright 2005). It has been noted that biochar amendments tend to benefit the number of spores relased by a 10-fold scale (Feng et al. 2021; Videgain-Marco et al. 2021). Additionally, on an experimental basis, biochar raises soil temperatures by an average of 2°C and reduces fluctuations between night and day temperatures, which leads to increases in AMF spore development (Mau and Utami 2014). Through the process of tilling biochar into soils, soil porosity, and carbon availability are increased, which directly correlate with the expansion of AMF spore activity (Neuberger et al. 2024). Additionally, biochar increases dissolved organic carbon, cations, and anions in the soil solution, which augments the electrical conductivity, leading to spore development (Joseph et al. 2021). Increasing the spore dispersal of AMF is incredibly beneficial economically and environmentally, as higher AMF presence in the soil may lead to higher yield, more drought resistance, and greater overall crop health.
Infective potential of AMF
Infective potential in plant microbiology is defined as the amount of energy available for a pathogen to infect a host (Kumar and Rao 1979). While too much AMF colonisation can be detrimental to plant growth (Poveda 2020), high AMF infective potential can also be a method of defence for host plants. Evidence suggests that high AMF infective potential can increase host disease rates; however, this is countered by literature that shows that previous inoculation with AMF decreases the severity of impact from other diseases (Eck et al. 2022). Videgain-Marco et al. (2021) found that soils amended with biochar had increased AMF infective potential and that host plants showed lower disease rates than areas not amended with biochar and AMF. The increased infective potential is especially high in these soils due to biochar’s ability to optimise the chemical structure and porosity of topsoils (Videgain-Marco et al. 2021). Through increased AMF infective potential, biochar could be part of the solution to disease control with reduced agrochemical usage.
Biodiversity and complexity of AMF
Soil biodiversity is defined as the complex network of life that exists within soil environments, including, but not limited to microbial communities, bacteria, and invertebrates (Wardle 2006). In general, higher levels of biodiversity are indicators of healthy and resilient ecosystems. On a smaller scale, higher levels of AMF biodiversity and complexity also lead to more productive host plants (Rawat et al. 2019). Biochar application to soil has been reported to increase the scale and complexity of microbial co-occurrence networks through changes in soil structure that promote more AMF hyphae growth (Yan et al. 2021). Biochar application also enhanced the presence of keystone species, which are essential to ecosystem health, especially in the fungal orders Diversisporales and Glomerales. Keystone species play a large role in carbon sequestration, nutrient cycling, and soil structure maintenance. Shannon diversity and Pielou evenness indices are measures used to evaluate living organisms’ genetic makeup in terms of overall diversity and species evenness (Spellerberg and Fedor 2003). Data from a two-way factorial design experiment in north-east China suggests that once-a-year biochar amendment increases AMF Shannon diversity and Pielou evenness indices, indicating that higher biodiversity could be achieved using a low-effort method (Xin et al. 2022). Consequently, biochar application to agricultural soils could be a key strategy in global soil restoration efforts. Although further research is required in this specific subsection of AMF functioning, these soils should be analysed for an array of AMF qualities to determine the beneficial impacts of biochar on ecosystem functioning.
Negative effects of biochar on AMF
It is incorrect to assume that biochar amendments will consistently result in beneficial outcomes in terms of AMF population or that AMF will consistently benefit plants. It should be noted that extremely high levels of AMF colonisation can be detrimental if left unchecked. Furthermore, though many studies have noted increases in root colonisation, plant productivity, and contamination decrease, biochar applied at the wrong time or rate can lead to suppression of AMF. A key variable for AMF success is high soil nutrient availability, which can be limited by biochar amendments. Some biochar application methods have resulted in decreases in phosphorus and nitrogen availability, hence limiting plant root growth and creating an unfavourable environment for AMF colonisation (Gaur and Adholeya 2000; Wallstedt et al. 2002). Though the mechanisms for this activity are still not fully understood, it is important to note the possibility of overall AMF biomass decrease under biochar conditions.
Combined application of biochar and AMF
The interaction between biochar and AMF inocula has shown high potential, specifically in the area of nutrient absorption, with studies showing that Na content increases by 54.2% and Fe increases by 1.8-fold in maize crops treated with biochar and AMF inocula in comparison with the control treatment (Sun et al. 2022). Not only does the interaction between biochar and AMF provide pathways to critical nutrient uptake, it also decreases the concentrations of heavy metals in crops, thereby reducing the potential for produce contamination hence protecting farmers from potential economic hurdles, and leading to improved health outcomes for humans and animals (Zhuo et al. 2020).
In addition to studies describing the positive impacts of biochar and AMF on nutrient uptake, other studies have observed the potential for biochar and AMF on drought tolerance (Barna et al. 2020; Videgain-Marco et al. 2021). Plants udner drought stress take up less nutrients, which decreases yield and quality. With the increasing likelihood of global food demand and changes in climate conditions that could create a strong reduction in available soil and water availability (Mansoor et al. 2022), increasing drought resistance in staple crops is crucial. In a study (Hashem et al. 2019) on chickpea (Cicer arietinum) drought tolerance, biochar application with AMF inoculation decreased the effect of drought on morphological traits (root and shoot length, total leaf area, and number of secondary branches). Though the mechanism behind this result is not fully understood, it has found that biochar pores hold significant amounts of water with dissolved nutrients that can be used by AMF to support the key nutrition of plants in drought-like conditions (Mickan et al. 2016). Furthermore, biochar’s ability to optimise the chemical structure and porosity of topsoils allows for greater water retention (Videgain-Marco et al. 2021). Additional analysis of the effects of biochar pore size is necessary to gain insight into possible drought mitigation strategies. Given the impacts of feedstocks and pyrolysis conditions on biochar characteristics, notably pore size, the results of such analysis could guide the selection of feedstock and pyrolysis conditions that would apply to specific applications.
Besides physical characteristic objectives, using biochar to promote AMF population growth in agricultural soils would align with the US Department of Agriculture’s strategic plan for 2022–2026 (U.S. Department of Agriculture 2021), which has core goals to: (1) address climate change via climate-smart agriculture, forestry, and renewable energy; (2) advance racial justice, equity, and opportunity; (3) create more/better markets for producers and consumers at home and abroad; and (4) tackle food and nutrition insecurity and food safety’ (further subgoals and specifics can be found at USDA.gov). Biochar and AMF have been proven to help increase crop yield, protect consumers from emerging contaminants in food systems, and benefit overall crop health. For these reasons, stakeholders must invest in scientific work to improve development and application techniques.
Though many peer-reviewed papers have published results noting the impacts of biochar on the total abundance and activity of AMF in soils, the mechanisms of these impacts are not entirely understood. Some suggested mechanisms include biochar altering the: (1) soil nutrient availability, which in turn affects AMF, (2) soil nutrient availability, which in turn affects other microbial communities that outcompete AMF, (3) plant–AMF signalling pathways, which then alter root colonisation, and (4) habitat characteristics that give rise to a refuge for AMF (Warnock et al. 2007). It is important to note that these mechanisms are not mutually exclusive nor scientifically proven; however, there still could be ways to explain the changes in Table 1.
The effectiveness of biochar application and implications for AMF and other soil microbes are highly variable and depend on the physical characteristics of the biochar. For this reason, it is hard to compare and standardise results as each study uses different methods. Furthermore, the confounding variables observed during research add another layer of uncertainty to the data. Soil type, structure, texture, nutrient composition, and weather conditions tend to be determining factors of success in most studies that examine the relationship between biochar and AMF (Gujre et al. 2021). To this end, a standardised pyrolysis and application methodology must be developed (especially from a research point of view), and ongoing reporting must be more detailed to increase the credibility and applicability of research in this field. Expanding knowledge in this field allows for a more sustainable agricultural scheme to be developed, which can have crucial implications for economic gain, food stability, and environmental health.
Conclusion
This review focused on biochar’s range of impacts on soil conditions and ecosystems and highlights the need for ongoing studies with standardised methodologies when evaluating biochars. Biochar application to agricultural soils provides many beneficial outcomes, including increased crop yield, decreased chemical contamination, higher nutrient loads, and increased AMF colonisation and biomass. However, due to the lack of understanding surrounding AMF behaviour in soils, using biochar to target this fungus is challenging. This is despite the many studies showing the positive correlation between biochar application and AMF growth. Increased root colonisation, spore density, infective potential, and general biodiversity have all been observed in past analyses; however, the lack of standardised pyrolysis and application methods leads to less conclusive results. Therefore, this review concludes that future scientific work must focus on understanding the impacts and mechanisms of biochar’s interaction with AMF and other soil biota to maximise potential positive outcomes on a long-term scale. Specifically, understanding how biochar pore size affects AMF and water-holding capacity will be especially useful as drought conditions become more common. Therefore, continued research efforts are needed to improve the efficiency of biochar–AMF interactions in complex agricultural schemes.
Data availability
Data sharing is not applicable as no new data were generated or analysed during this study.
Declaration of funding
This work was supported by the University of Wisconsin–Green Bay Research Enhancement Program under the grant number AAN1283. Summer undergraduate research support was also provided by the Freshwater Collaborative of Wisconsin and the Freshwater@UW Summer Research Opportunities Program under grant SL3.09, along with the University of Wisconsin Sea Grant Institute, under grants from the National Sea Grant College Program, National Oceanic and Atmospheric Administration, US Department of Commerce, and from the State of Wisconsin, federal grant number (NA18OAR4170097) and project number (A/HCE-01); the University of Wisconsin Water Resources Institute under grants from the U.S. Geological Survey, US Department of the Interior, and from the State of Wisconsin, federal grant number (G21AP10608); and the University of Wisconsin–Madison.
Acknowledgements
The authors thank the University of Wisconsin-Green Bay and the Freshwater Collaborative of Wisconsin for support during this work. The authors also thank all additional funders and lab collaborators who have contributed to this work.
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