MINI REVIEW

published: 07 March 2022

doi: 10.3389/ffunb.2022.723892

Roles of Arbuscular Mycorrhizal

Fungi on Soil Fertility: Contribution in
the Improvement of Physical,
Chemical, and Biological Properties
of the Soil
Abdoulaye Fofana Fall 1,2*, Grace Nakabonge 3 , Joseph Ssekandi 1 ,
Hassna Founoune-Mboup 4 , Samuel Obeng Apori 5 , Abibatou Ndiaye 1 , Arfang Badji 6 and
Khady Ngom 1
1
African Center of Excellence in Agroecology and Livelihood Systems, Faculty of Agriculture, Uganda Martyrs University,
Nkozi, Uganda, 2 Fungi Biotechnology Laboratory, Plant Biology Department, Cheikh Anta Diop University of Dakar (UCAD),
Dakar, Senegal, 3 College of Agriculture and Environmental Sciences, Makerere University, Kampala, Uganda, 4 ISRA_LNRPV,
Laboratoire National de Recherches sur les Productions Végétales (LNRPV), Dakar, Senegal, 5 School of Food Science and
Environmental Health, Technological University Dublin, Dublin, Ireland, 6 Department of Agricultural Production, Makerere
University, Kampala, Uganda
Edited by:
Paola Angelini,
University of Perugia, Italy Many of the world’s soils are experiencing degradation at an alarming rate. Climate
Reviewed by: change and some agricultural management practices, such as tillage and excessive
Birinchi Kumar Sarma, use of chemicals, have all contributed to the degradation of soil fertility. Arbuscular
Banaras Hindu University, India
Mahaveer P. Sharma,
Mycorrhizal Fungi (AMFs) contribute to the improvement of soil fertility. Here, a short
ICAR Indian Institute of Soybean review focusing on the role of AMF in improving soil fertility is presented. The aim of
Research, India
this review was to explore the role of AMF in improving the chemical, physical, and
Ees Ahmad,
National Bureau of Agriculturally biological properties of the soil. We highlight some beneficial effects of AMF on soil
Important Microorganisms carbon sequestration, nutrient contents, microbial activities, and soil structure. AMF has
(ICAR), India
a positive impact on the soil by producing organic acids and glomalin, which protect
*Correspondence:
Abdoulaye Fofana Fall
from soil erosion, chelate heavy metals, improve carbon sequestration, and stabilize
abdoulaye.fofana@stud.umu.ac.ug soil macro-aggregation. AMF also recruits bacteria that produce alkaline phosphatase,
a mineralization soil enzyme associated with organic phosphorus availability. Moreover,
Specialty section:
AMFs influence the composition, diversity, and activity of microbial communities in the soil
This article was submitted to
Fungi-Plant Interactions, through mechanisms of antagonism or cooperation. All of these AMF activities contribute
a section of the journal to improve soil fertility. Knowledge gaps are identified and discussed in the context of
Frontiers in Fungal Biology
future research in this review. This will help us better understand AMF, stimulate further
Received: 11 June 2021
Accepted: 19 January 2022
research, and help in sustaining the soil fertility.
Published: 07 March 2022
Keywords: macro-aggregation, microorganisms, glomalin, rock phosphate, P legacy
Citation:
Fall AF, Nakabonge G, Ssekandi J,
Founoune-Mboup H, Apori SO, INTRODUCTION
Ndiaye A, Badji A and Ngom K (2022)
Roles of Arbuscular Mycorrhizal Fungi
The most significant threats to soil function at the global level are soil erosion, soil organic
on Soil Fertility: Contribution in the
Improvement of Physical, Chemical,
carbon, excessive use of input, and nutrient imbalance (Montanarella et al., 2016). The depletion
and Biological Properties of the Soil. of soil fertility in the world has increased due to unsustainable land management practices,
Front. Fungal Biol. 3:723892. such as overgrazing, bush burning, continuous crop cultivations, and tillage practices (Dewitte
doi: 10.3389/ffunb.2022.723892 et al., 2013). However, inoculation with Arbuscular Mycorrhizae Fungi (AMFs) has been

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Fall et al. Mycorrhizal Fungi and Soil Fertility

identified as an eco-friendly approach to improve soil fertility benefits carbohydrates from the host plants (Diagne et al., 2020).
(Dal Cortivo et al., 2018). AMF is the most widespread soil Many authors demonstrated that AMF obtains up to 20% of
microorganisms that form a symbiotic relationship with more photosynthetic carbohydrates from the host plant (Bonfante and
than 80% of plants (Prasad et al., 2017), except for a few plant Desirò, 2015; Kaiser et al., 2015). In addition to carbohydrates,
families, such as Amaranthaceae, Brassicaceae, Cruciferae, lipids are a major source of organic carbon delivered to the
Chenopodiaceae, Caryophyllaceae, Juncaceae, Cyperaceae, and fungus (Luginbuehl et al., 2017). It has been discovered that
Polygonaceae, which do not exhibit any association (Brundrett, plants provide the fungus with some of the fatty acids that
2009). They can be found in various ecosystems worldwide the microorganism needs to grow (Keymer et al., 2017). The
(Verbruggen et al., 2012). AMF is a key component of soil biosynthesis of fatty acids has not been observed in AMF in
microorganisms and belongs to the glomeromycota phylum. the absence of the plant. Moreover, the genes encoding for
This phylum is divided into three classes (Archaeosporomycetes, fatty acid biosynthesis have not been found in AMF, therefore,
Glomeromycetes, and Paraglomeromycetes), five orders these microorganisms depend on the lipid biosynthesis of the
(Archaeosporales, Diversisporales, Gigasporales, Glomerales, host plant. AMF is an extremely ancient symbiosis. Based on
and Paraglomerales), 14 families, 29 genera, and more than archeologic records, it dates to the appearance of terrestrial
240 species (Krüger et al., 2012; Redecker et al., 2013). Several plants million years ago and would have accompanied vascular
species of AMF have been studied in the world, however, the plants to colonize the terrestrial environment (Humphreys et
most species used as a model are as follows: Funneliformis al., 2010). AMF does not only have an impact on plant growth
mosseae (previously known as Glomus mosseae), Gigaspora and production but it has been also reported that they improve
rosea, Gigaspora margarita, Gigaspora gigantea, and Rhizophagus some soil characteristics, such as soil aggregation, soil nutrients
irregularis (previously known as Glomus intraradices and Glomus availability, water retention, microbial activities, nitrogen,
irregulare; Schüßler and Walker, 2010). AMF is not a parasite carbon, and phosphorus cycling, and soil acidity correction
but obligate symbionts that need a host plant to complete (Sadhana, 2014; Jamiołkowska et al., 2018; Parihar et al., 2020).
their life cycle. They improve crop productivity by increasing Several studies have reported that they play a crucial role in plant
water and nutrient uptake, such as nitrogen (N), phosphorus resistance against biotic and abiotic stresses. This review aims
(P), and potassium (K) (Anderson et al., 2018). The increase to summarize knowledge about AMF symbiosis, in particular,
of the host plant nutrient uptake is due to the characteristics the beneficial effects on soil (Figure 1). First, the role of AMF
of AMF mycelium. These mycelia or hyphae absorb nutrients in the physical, chemical, and biological properties of the soil
by osmotrophy and explore more surface area compared to is considered. The contribution of AMF in soil aggregation,
non-mycorrhizal roots (Duponnois et al., 2011). In return, AMF nutrient availability, and boosting beneficial soil microorganisms

FIGURE 1 | Effects of arbuscular mycorrhizal fungi on improving soil fertility.

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Fall et al. Mycorrhizal Fungi and Soil Fertility

is discussed. Finally, the role diversity of interactions between of organic and inorganic phosphorus (Requejo and Eichler-
AMF and other soil microorganisms is examined. Löbermann, 2014). The accumulation of P in soils from fertilizers
is known as legacy P (Sattari et al., 2012). This legacy P has
the potential to play a key role in maintaining agricultural
ROLE OF AMF ON IMPROVING SOIL productivity (Condron et al., 2013; Rowe et al., 2016). It has
PHYSICAL PROPERTIES been revealed that the accumulated P in soils is sufficient to
sustain crop yields worldwide for about 100 years (Khan and
Importance of AMF on Soil Structure Zaidi, 2007). Unfortunately, only a few quantities of this soil’s P
Arbuscular Mycorrhizae Fungi have a beneficial effect on soil are available for the plants (Balemi and Negisho, 2012). The P
structure. The AMF mycelia are present in massive quantities is most often in the form of inorganic orthophosphate adsorbed
in soils (de Novais et al., 2019). These mycelia or hyphae have to soil cations. Thus, the availability of P in soil is affected
the property to create stable soil aggregations. Mycorrhizal fungi by the presence of iron (Fe), calcium (Ca), and aluminum
act as a long-term soil binding agent through the production of (Al) oxides, which fix phosphorus as iron phosphate (FePO4 ),
a glycoprotein (glomalin) by the extramatrical mycelia (Singh tri-calcium phosphate [Ca3 (PO4 )2 ], and aluminum phosphate
et al., 2020). This glomalin is a hydrophobic, thermo-tolerant, (AlPO4 ) (Amanullah and Zakirullah, 2010; Shen et al., 2011).
or heat-tolerant resistant to the hot temperature of the soil. Therefore, only a small proportion (<1%) of the legacy P is
The hydrophobic character of the glomalin confers resistance available to plants (Rodrigues et al., 2021). Phosphorus is taken
of soil aggregations to water, the production of this substance in the form of orthophosphates (inorganic phosphate Pi) by
reaches its maximum in senescent mycelia. The glycoprotein plants, but this mineral form of phosphorus is in limited quantity
is slowly biodegradable by bacteria and fungi in the soil. The in the soil and, under the action of root sampling, areas are
main function of glomalin is to stabilize soil aggregations quickly created depletion around the roots due to a slow supply
(Hu et al., 2019; Mubekaphi, 2019), act as a glue that binds of P, slow phase of the soil, and the low mobility of P in soils
together the soil micro-aggregations (diameter <250 µm) to (Javot et al., 2007). The reservoir of P must be hydrolyzed to
form stable macro-aggregations (Lehmann et al., 2020). These make it available in the soil for plants uptake. AMF plays a
soil macro-aggregations ensure better water infiltration, reduce key role in improving P availability in the soil. Indeed, it is
surface runoff, control soil erosion, reduce nutrients and organic a P activator that can accelerate the process to transform P
matter losses, increase gas exchange better retention of water into bio-available forms via a range of chemical reactions and
and minerals, especially potassium, therefore, improve crop biological interactions (Zhu et al., 2018). It was believed that
productivity (Demenois et al., 2018; Parihar et al., 2020). In AMF hydrolyzes the organic P into inorganic phosphorus (Shen
addition, the mycelia network constantly renews itself and the et al., 2011) through a mechanism linked to the production of
dead mycelia also preserve soil structure until decomposition enzymes named phosphatase (Tarafdar and Marschner, 1994).
(Gianinazzi et al., 2010). These dead mycelia contribute to the However, recent studies revealed that AMF lack the capacity
stocks of organic matter and physical binder involved in soil to release phosphatases into the soil (Zhang et al., 2016) but
aggregation (Hamel and Plenchette, 2017). All these mechanisms they recruit bacteria known as Phosphate Solubilizing Bacteria
reduce the risks of soil compaction and promote soil fertility (PSB) that produce phosphatase, which mineralizes organic P and
(Norton et al., 2020). It can be said that AMFs improve soil provides a function that is absent from the AMF (Zhang et al.,
structure through their chemical and biophysical mechanisms, 2018; Etesami and Jeong, 2021). PSB and AMF association is
such as enmeshment and alignment. However, there is a lack a beneficial feature that has the ability to mineralize insoluble
of information about the lifespan of glomalin in the soil and phosphate in the soil and release soluble P that can easily be
the effect of anthropology activities, such as bush burning, on assimilated by plants (Wei et al., 2017; Mahanta et al., 2018).
soil glomalin. In that association, the role of PSB is to produce organic acids,
such as gluconic acid, ketogluconic acids, siderophores, protons,
ROLE OF AMF ON IMPROVING SOIL and acid phosphatases that are involved in the mineralization of
organic P in soil (Dobbelaere et al., 2003; Lucy et al., 2004), while
CHEMICAL PROPERTIES AMF mycelia improve the absorption of soluble P in the plants
(Taktek et al., 2017). The phosphatase releases P from organic
Arbuscular Mycorrhizae Fungi symbionts are recognized as
P or inorganic orthophosphate by hydrolyzing phosphoric acid
being major microbial components in the development of the
monoesters into P ion and a molecule with a free hydroxyl group
main biogeochemical cycles of soils (P, N, and C). This results
(Othman and Panhwar, 2014). It was proven that the double
in an improvement in the growth of mycorrhizal plants.
inoculation of R. irregularis and Rahnella aquatilis improves
solubilization of inorganic P by the increased production of
Contribution of AMF on Soil Phosphorus phosphatase released by the bacteria that is also stimulated
Legacy Availability by AMF exuded fructose (Zhang et al., 2018). AMF can also
Phosphorus is an essential element for plants. It is a component solubilize inorganic phosphate into soluble forms through the
of many molecules, such as adenosine triphosphate (ATP), processes of acidification, chelation, exchange reactions, and
nucleotides, phospholipids, certain enzymes, and co-enzymes production of organic acids, H+ , and metabolites (Relwani et al.,
(Agledal et al., 2013). Most soils contain enormous amounts 2008; Behera et al., 2014). It is demonstrated that the metabolic

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Fall et al. Mycorrhizal Fungi and Soil Fertility

activities of AMF produce alkaline phosphatases, which cleave AMF plays also a critical in the formation and maintenance
substrates present in the soil and make the phosphate accessible of soil aggregations through the production of Glomalin. This
(Liu et al., 2013). Moreover, the organic acid produced by glomalin protects organic matter from microbial degradation,
AMF solubilizes insoluble mineral phosphate into a soluble form increases the hydrophobicity and stability of macro-aggregations,
(Lapeyrie, 1988). In addition, AMFs help to release P from which control soil carbon loss and increase soil carbon stocks (C
rock phosphate (RP) fertilizer. RP has low effectiveness. This is sequestration; Wilson et al., 2009; Rillig et al., 2010). More studies
due to when added as fertilizer only one part is accessible to are needed to distinguish the role of AMF in the dynamics of soils
the plants and the remaining part is converted into insoluble carbon sequestration. This involves in particular determining
fixed forms (Billah et al., 2019). Thus, AMF can solubilize the quantity of carbon fixed by the AMF because this lack of
insoluble phosphate from RP to make it available in the soil knowledge means that AMF cannot currently be included in the
(Andrino et al., 2021). AMF converts the insoluble P into soluble models of reducing the rate of atmospheric carbon. In addition,
forms through their production of acids during their metabolic limited information is available on the regulation of carbon to
activities (Kalayu, 2019). However, little is known about whether nutrient exchange across the mycorrhizal interface.
there is an activator dose of P that allows AMF to initiate root
infection. Because it is known that a rate of 50 kg N ha−1 is the Contribution of AMF on Soil Trace
starter dose to activate rhizobium symbiosis. Elements Transfer
Contribution of AMF on Soil Nitrogen Trace elements play roles in enzymatic activities involved in
photosynthesis, oxidative respiration, protection against free
Availability radicals, or even lipid biosynthesis (Dominguez-Nuñez et al.,
Like phosphorus, nitrogen (N) is a vital part of plants. It is 2016). It is known that AMF allows better absorption of low
a constituent of phospholipids, coenzymes, and amino acids mobile trace elements in soils, such as potassium (K), calcium
(Hawkesford et al., 2012). In the soil, N is present in organic (Ca), magnesium (Mg), copper (Cu), zinc (Zn), iron (Fe),
and mineral forms (nitrites, nitrates, and ammonium ions). manganese (Mn), and cobalt (Co) (Garcia et al., 2016; Hashem
The ammonium form is weakly absorbed by plants that prefer et al., 2018). For instance, according to Krishna and Bagyaraj
nitrogen in the form of nitrate (NO− 3 ). AMF helps to mobilize (1984), the level of Zn, Fe, and Mn is twice in mycorrhizal
the inorganic form of nitrogen (NH4+ ) from the soil (Casieri peanut plants compared to non-mycorrhizal plants. It has also
et al., 2013). The AMF mycelium is able to absorb nitrogen in the been revealed that mycorrhizal inoculation improved Zn and
form of ammonium ions (NH4+ ), in the form of nitrates (NO− 3 ), Cu nutrition in soybeans and clover (Schoeneberger et al.,
and in the form of amino acids (Chen et al., 2018; Drechsler et 1989). However, when some of these elements are present in
al., 2018; Jansa et al., 2019). Nitrogen availability requires the high quantities and therefore possess a toxic character, the
activity of local transporters in the AMF hyphae. It has also mycorrhization can play a role in the protection of the plant, by
been demonstrated that mycorrhizal associations could play a strong retention of these elements (Liu et al., 2000). Besides trace
significant role in the decomposition and mineralization of plant elements, more research studies are needed on the role of AMF
organic matter and mobilize nutrients, particularly nitrogen, for to synthesize or transport phyto-hormones (auxin, cytokinins,
the benefit of the host plant (Lambers et al., 2008). However, gibberellic acid, etc.) and antibiotics from plant to plant and from
more research should be conducted in a controlled environment plant to soil microorganisms.
to determine the quantity of nitrogen that transits through the
AMF mycelia network. Moreover, a study should be carried out
to determine whether AMFs use nitrogen from the soil or from ROLE OF AMF ON IMPROVING SOIL
the host plant. BIOLOGICAL PROPERTIES
Contribution of AMF on the Soil Carbon Microorganisms are one of the most important soil components.
Cycle and C Sequestration These microorganisms interact between them and with their
Arbuscular Mycorrhizae Fungi play an essential role in the global environment to contribute to the functioning of the soil and
C cycle. AMF hyphae are involved in C translocation into the thus participate in the provision of ecosystem services necessary
soil and provide a key link in the terrestrial C cycle (Finlay, for our survival (plant production, purification of pollutants,
2008). Indeed, AMF is an efficient agent to improve carbon etc.; Nielsen et al., 2011). Soil is therefore a continually active
sequestration in a mechanism of translocation C away from biological reactor where diverse biochemical reactions and
the high respiratory activity around the root and into the soil essential ecological processes happen (solubilization of organic
aggregations (Zhu and Miller, 2003). It has been demonstrated matter, the biogeochemical cycles of the elements, etc.; Gessner
that mycorrhizal roots create a sink demand for carbon. When et al., 2010). The microbial activities in the soils contribute to its
the atmospheric CO2 increases, the allocation of C from the fertility through synergy between microorganisms, competition,
plants to AMF also increases and stimulates the growth of AMF and parasitism (Topalović and Vestergård, 2021). Within the soil,
(Drigo et al., 2010). This C demand is provided by the host AMFs interact with a wide range of microorganisms to better
plant from the C fixed through photosynthesis (Parihar et al., improve soil fertility. It has been demonstrated that the secretions
2020). In addition, AMF extramatrical hyphae represent 20–80% of AMF influence the composition and activity of microbial
soil microbial biomass which consists of 15% of soil organic communities in the rhizosphere (Veresoglou and Rillig, 2012).
C (Kabir et al., 1997; Leake et al., 2004). As discussed above, The biological activities of AMF lead to the appearance of a

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Fall et al. Mycorrhizal Fungi and Soil Fertility

TABLE 1 | Examples of some interactions between AMF and soil microorganisms.

Interactions AMF + microbes Mechanisms Effects References

AMF and Pseudomonas Glomus intraradice (AMF) stimulates the The antibiotic protects the host plants against Ma et al., 2019
fluorescens production of antibiotic Gaeumannomyces graminis.
(2,4-diacetylphloroglucinol) by Pseudomonas
fluorescens.
AMF and saprotrophic fungi AMF increase the biomass of saprotrophic Dissolution of soil organic matter into mineral Albertsen et al., 2006;
fungi. matter. Carteron et al., 2021
AMF and Gram-positive/negative AMF has a deletion effect on certain This interaction affects the production of Welc et al., 2010
bacteria Gram-positive and Gram-negative bacteria. bioactive metabolites and the decomposition
of organic matter.
AMF and Rhizobia AMF work in synergy with Rhizobia Provide legumes woody and crop legumes Chatarpaul et al., 1989;
(Faba bean) with essential soil nutrients Xavier and Germida, 2002
AMF, Rhizobia, and phosphorus Tripartite relationship. Solubilize P by Improve host plant phosphorus uptake. Afkhami and Stinchcombe,
solubilizing microorganisms mineralization, low soil pH, chelation and 2016; Kalayu, 2019;
(PSM) production of phosphatase, organic acid and Nacoon et al., 2020
proton.
AMF and Mycorrhization Helper MHB help: in the receptivity of the root to the Beneficial effect of bacteria on mycorrhizae. Rigamonte et al., 2010
Bacteria (MHB) AMF, in root-AMF recognition, in AMF growth, Improve soil fertility and nutrients uptake by
in the modification of the rhizospheric soil, and the host plants.
in the germination of AMF propagules.
AMF and Plant Growth AMF work in synergy with PGPR to stimulate Soil fertility and plant growth. Increase the Linderman, 2000; Ahemad
Promoting Rhizobacteria (PGPR) Ammonia production, N fixation, solubilization diversity and abundance of soil parasite and Kibret, 2014; Vafadar
of mineral phosphate, and other essential antagonists. Mitigate water deficit damage et al., 2014; Battini et al.,
nutrients, production of plant hormones. and improve water stress tolerance (i.e., 2016; Moreira et al., 2020
Accumulate ascorbate peroxidase and Cupressus arizonica).
glutathione peroxidase. Secrete organic acids
responsible for dissolving phosphorus phytate
mineralization and inorganic P solubilisation.
PGPR found in AMF mycelia produce
siderophore and indol acetic acid production.
AMF and Frankia Synergistic interaction between AMF and Improve actinorhizal plants’ height, the Oliveira et al., 2005
Frankia (nitrogen-fixing actinobacteria). numbers and dry weight of root nodules, leaf
area, shoot height, total biomass, and N and
P leaf contents (i.e., Alnus glutinosa).
AMF and Bacillus subtilis AMF stimulate the production of nitrate and Increase shoot and root dry weight, nodule Hashem et al., 2017
nitrite reductase and nitrogenase activities and number, and leghemoglobin content.
osmoprotectants such as glycine, betaine, and
proline by Bacillus subtilis.

AMF, Arbuscular Mycorrhizae Fungi.

positive, neutral, or negative relationship between AMFs and rhizobia) and fungal communities depends on N and P status
other soil microorganisms. in the soil. However, there is a lack of information and pending
Many microbial components of the soil work synergistically questions which need to be answered. How soil microorganisms
with AMF, promoting the growth and protection of plants may hamper or totally inhibit the activities and functioning
(Gryndler, 2000; Barea et al., 2002). The positive interactions of AMF? What is the role of AMF in the trophic chain? In
involve the nutrient acquisition, biological control of root another word, can AMFs subject to any kind of predation or
pathogens, improvement of plant tolerance to abiotic parasitism from soil microorganisms? In addition, a study on
stresses, and soil fertility. AMF communities influence AMF and free native nematode interactions and their impact on
the physicochemical environment of the rhizosphere and the development of cereal crops under water stress conditions
control various soil microbial interactions (Alimi et al., 2021). are also needed.
Mycorrhization directly affects the quantity and quality of
root exudates. These exudates influence the composition of
the microflora of the rhizosphere (Baltrus, 2017). Table 1
INFLUENCE OF CROP AND SOIL
shows some examples of interactions between AMFs and MANAGEMENT PRACTICES ON THE AMF
other microorganisms. However, these interactions depend FUNCTIONING AND PERFORMING
on several factors, such as the amounts of phosphorus and VARIOUS SOIL FUNCTIONS
nitrogen available (Larimer et al., 2014). This is confirmed by
Wang et al. (2011) and Xu et al. (2018) who demonstrated a Arbuscular Mycorrhizae Fungi improve soil health by improving
synergistic relationship between AMF and the bacterial (i.e., its physical, chemical, and biological health. The previous

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Fall et al. Mycorrhizal Fungi and Soil Fertility

sections have described its role in nutrient cycling and interaction pesticides, and herbicides applications are essential for crop
with other soil microorganisms. However, agriculture practices nutrition, and they replenish the soil nutrients pool removed
significantly impact AMF communities and their performance, or harvested by crops, weed control, and pest management
influencing various soil functions. This section will discuss some (Rana et al., 2019). However, high- and long-term inorganic
crop and soil management practices and their influence on AMF fertilizer application reduces the plant’s dependency on AMF,
functioning and performing activities in the soil. subsequently, mycorrhizal diversity, and abundance (Kour et al.,
2020). This phenomenon can be explained by the fact that
Impact of Crop Management on AMF the symbiotic relationship between the plant and AMF is
To maintain and improve nutrient availability and soil health and energetically costly for the plant; therefore, when the soil is
minimize pest and disease incidence, farmers use management rich in nutrients, plants allocate fewer carbohydrates to AMF,
strategies, such as crop rotation and intercropping, to achieve which ultimately affects the spore development and hyphae
sustainable agriculture production. These two practices influence production (Tian et al., 2013). Soil phosphorus plays the
the diversity, abundance, and functioning of AMF in the soil. most significant role in regulating plant mycorrhizal symbiosis
For instance, crop rotation increases the number of AMF spore (Kowalska et al., 2015). Thus, high P application negatively affects
density and root colonization in maize, and it also increases root colonization and AMF diversity (Cheng et al., 2013). On
wheat yield when the preceding crop is soybean or chickpea the other side, low fertilizer application optimizes the plant
(Higo et al., 2013; Bakhshandeh et al., 2017). These positive effects mycorrhizal symbiosis (Rana et al., 2020). For instance, Liu
are not the effects of leguminous nitrogen fixation but rather due et al. (2016) found that P application in nutrient-deprived soil
to higher AMF activities in the soil (Bakhshandeh et al., 2017). improves the mycorrhizal-mediated benefits to the plant. In
However, crop rotation with non-mycorrhizal crops, such as the addition, organic fertilizer application also has both positive
Brassica family, reduces the abundance of AMF in the soil and the and negative impacts on AMF diversity (Liu J. et al., 2020).
symbiotic benefits conferred to the crops through the production According to Zhu et al. (2016), organic matter improves the
of antimicrobial isothiocyanates (Valetti et al., 2016). Moreover, AMF community composition in the rhizosphere of maize. In
some plants are more mycorrhizal-dependent than others. For the same vein, inorganic pesticide and herbicide applications also
instance, maize crop is more mycotrope than soybean (Troeh and have both positive and negative effects on AMF. These effects are
Loynachan, 2003; Wang et al., 2016), while wheat is considered a mediated by the secretion of active substances up taken by plants
non-mycorrhizal or mycorrhizal plant species depending on the via root or hyphal from the rhizosphere (Hage-Ahmed et al.,
cultivar (Hetrick et al., 1993; Stefani et al., 2020). Wheat cultivars 2019). For instance, azoxystrobin and glyphosate, respectively,
respond differently to AMF inoculation in terms of growth, root fungicide and herbicide, inhibit the spore germination of
colonization, and carbon for nutrient exchange; depending on some AMF species (Buysens et al., 2015). In addition to
the age of a cultivar, old cultivars benefit more consistently from the chemical application, extensive tillage greatly influences
AMF than new cultivars that effectively exploit highly fertilized the community composition of AMF by reducing mycelium
systems with less reliance on symbiosis (García de León et al., extension, colonization rate, and diversity structure (Säle et al.,
2020). This mechanism is known as a mycorrhizal dependency, 2015; Zhao et al., 2015). This is due to the effects of tillage on
whereby AMF presence in the soil affects the growth response of the soil’s permeability, texture, and microbial food substrates,
plant species differentially (Kandhasamy et al., 2020). In addition, which ultimately affect the soil microbiota activity and their
intercropping various plant species in similar conditions impacts habitat type (Wang et al., 2020). Several studies demonstrated
the composition of the community and diversity of AMF and that conservation or zero tillage improves AMF diversity and
the plant diversity. Indeed, AMF is involved in transporting abundance, resulting in better plant growth (Qin et al., 2017; Gu
plant assimilates from the dominant species to plant species et al., 2020). However, a study carried out in a Mediterranean
subordinated through the AMF mycelium network (Egerton- agroecosystem found that AMF spore density or extraradical
Warburton et al., 2007). Therefore, the presence of AMF impacts mycelium density is not affected by conventional tillage practices
the type of vegetation, the relative abundance of plant species, (Curaqueo et al., 2011). These contradicting results indicate that
and their diversity (Yadav et al., 2020). This mechanism operates the effects of different chemical applications and tillage practices
through a change in the soil microorganisms and soil properties on AMF still need further studies to understand better the effect
(soil structure; Liu L. et al., 2020). AMF influences soil microbiota of different soil management practices on soil AMF.
through mycelium products and biophysical mechanisms, such
as enmeshment and alignment. All these mechanisms occur in
a complex interaction process that involves various factors in a CONCLUSION AND FUTURE AREAS OF
cycling way. However, we need to carry more research studies at RESEARCH
the field level to really understand the impact of plant biodiversity
on AMF diversity and functioning. Arbuscular Mycorrhizal Fungi appear to be one of the most
important soil organisms to take into account. AMFs are involved
Impact of Soil Management Practice on in plant mineral nutrition, water absorption, and protection
AMF against biotic and abiotic stresses in plants. Despite the fact
Soil management practices have a significant impact on soil that the importance of AMF in improving soil fertility is well-
properties and microbial diversity. Mineral fertilizer, chemical established, our understanding of the underlying mechanisms

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Fall et al. Mycorrhizal Fungi and Soil Fertility

is still limited. There are a few studies that simultaneously acid, and antibiotics from plant to plant and from plant to soil
investigated the effects of AMF on the physical, chemical, microorganisms, is also poorly understood. Therefore, studies
and biological properties of the soil. Therefore, the current targeting the identification and characterization of such AMF
review provides a holistic overview of the existing information function are paramount.
regarding the role of AMF symbiotic relationships with crops • Moreover, the role of AMF in soil basal respiration is an
in improving the physical, chemical, and biological properties interesting field to investigate.
of the soil. Regarding the impact of AMF on soil fertility, • Advances in our knowledge of the functions played by AMFs
we highlighted several mechanisms, such as the production in the soil are partly hindered by the obligate biotrophic
of glomalin, which is beneficial to the accumulation and nature of these fungal microorganisms. Therefore, more field
circulation of soil carbon and enhances soil stability. In experiments on the impact of plant biodiversity on AMF
addition, the beneficial interaction between AMF with other diversity and functioning are necessary.
soil microorganisms, such as PSB, which produce phosphatase • Moreover, future studies on the effects of different
and mineralize organic P, was highlighted. However, some soil management practices (i.e., tillage) and chemical
of the functions involved in this symbiosis that determine applications on AMF functioning and performance in the soil
the performance of AMF in the soil should be addressed in are important.
future studies.
All these research topics should be based on new approaches,
• Future studies are required to characterize the soil P and N such as recent methodological advances in physiology, molecular
critical threshold below which AMF establishes symbiosis and biotechnology, and agroecology integrated into both laboratory
above which AMFs are not active on a broader range of plant- and field conditions. Such interventions are paramount to our
AMF species combinations, soil types, and edaphic conditions. ability to establish a new “green revolution” aligned to the
• Furthermore, future research should investigate the regulation requirements for achieving a sustainable development ingrained
of N and its uptake from the soil during AMF symbiosis by in agricultural production.
using molecular tools, such as transcriptomic, gnomonic, and
the development of fungal mutants. AUTHOR CONTRIBUTIONS
• Further attention is needed on the role of glomalin in
improving carbon sequestration efficiency from various AFF contributed on inception of the paper, research, and writing.
climate and soil types to expedite its use in solving soil GN contributed on inception and reviews of the paper. JS
degradation problems that will be worsened by prevailing contributed on inception and reviews. HFM contributed on
climate disturbances. Assessing the accumulation and lifespan inception and reviewed the work. SOA contributed on write up.
of glomalin in soil fertility parameters under different climate, AB contributed on write up and the revision of the manuscript.
land use, and management conditions is of critical importance. AN and KN reviewed the work. All authors contributed to the
• It is well-established that the interactions between AMF and article and approved the submitted version.
certain soil microorganisms are beneficial to soil fertility;
however, the interaction between AMF and free native FUNDING
nematode and their impact on soil structure under drought
stress calls further research. This work has been funded by Regional Academic Exchange
• The role of AMF in the synthesis or transport of for Enhanced Skills in Fragile Ecosystem Management in Africa
phytohormones, such as auxin, cytokinins, gibberellic (REFORM) grant number: 2017–2861.

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doi: 10.1016/S0038-0717(01)00165-1 Publisher’s Note: All claims expressed in this article are solely those of the authors
Xu, J., Liu, S., Song, S., Guo, H., Tang, J., Yong, J. W., et al. (2018). Arbuscular and do not necessarily represent those of their affiliated organizations, or those of
mycorrhizal fungi influence decomposition and the associated soil microbial the publisher, the editors and the reviewers. Any product that may be evaluated in
community under different soil phosphorus availability. Soil Biol. Biochem. 120,
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181–190. doi: 10.1016/j.soilbio.2018.02.010
endorsed by the publisher.
Yadav, A. N., Rastegari, A. A., Yadav, N., and Kour, D. (2020). Advances
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Zhang, L., Shi, N., Fan, J., Wang, F., George, T. S., and Feng, G. (2018). Arbuscular Badji and Ngom. This is an open-access article distributed under the terms of
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