Open access peer-reviewed chapter

Recent Advances in Plant: Arbuscular Mycorrhizal Fungi Associations and Their Application to Cassava Crops

Written By

Sarah Otun and Ikechukwu Achilonu

Submitted: 15 August 2022 Reviewed: 15 September 2022 Published: 22 March 2023

DOI: 10.5772/intechopen.108100

From the Edited Volume

Arbuscular Mycorrhizal Fungi in Agriculture - New Insights

Edited by Rodrigo Nogueira de Sousa

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Abstract

According to estimates, the world’s population is growing at 0.96% yearly, meaning that there will be approximately 7.3 billion people on earth by the year 2050. Consequently, the agricultural sector is demanded to boost production and provide food security for the rising world’s population. Unfortunately, almost 40% of the arable land has been damaged by several factors, such as industrialization, suburbanization, acidification, salinization, and erosion of the soil, environmental pollution, among others, resulting in a global agricultural and economical problem. However, several land recovery techniques have been developed over many years of research, such as the use of chemicals, cultural techniques, and Arbuscular Mycorrhizal Fungi (AMF). AMF forms a vital connection with the host plants and the soil nutrients and assists in the restoration of damaged agricultural lands. This reviews’ objective includes (i) providing a brief overview of AMF; (ii) highlighting AMF’s role in nutrient management; (iii) reviewing the roles of AMF in the regulation of plant (cassava) development; (iv) explaining the role of AMF in managing abiotic and biotic stressors; (vi) emphasizing the role of AMF in reducing greenhouse gas emissions, and (vi) highlighting significant areas within the study of AMF-cassava that has not yet been completely explored.

Keywords

  • arbuscular mycorrhizal fungi
  • beneficial soil microorganisms
  • cassava
  • soil nutrients
  • sustainable agriculture
  • nutrient-loss/uptake
  • abiotic/biotic stressors

1. Introduction

A symbiotic relationship between the root of the host plant and the fungi and spreads into the rhizosphere and the surrounding soil is known as mycorrhiza [1]. Mycorrhizal symbiosis has developed into a specialized area since Frank first used the term 136 years ago [2]. The scientific community aims to understand its characteristics and ramifications for both plants and fungi and other microorganisms like bacteria [1]. Mycorrhizae are now recognized as a type of “biological fertilization”, mostly because of their presence in almost all healthy plant roots [3]. Biofertilization has been presented as the bio-sustainable substitute for chemical fertilization, because of its variety of possible benefits, including fertilizing the host plants with nutrients, protection from biotic (pathogens), abiotic (drought, unfavorable temperature, amongst others) [1, 3].

Mycorrhiza, as a crucial functional group of the soil biota, provides various nutrients and aids the host plant in battling against unfavorable soil conditions (for instance, in drought settings it aids in increasing the surface of the roots) [4]. Endomycorrhizal fungi and ectomycorrhizal fungi are two different forms of mycorrhizal fungi that form a root biotrophic relationship [5]. In contrast to ectomycorrhizal fungi, which develop around the surrounding root cells, endomycorrhizal fungi colonize the root cells of plants [5].

AMF has been reported to be present in 80% of terrestrial plant species, including economically important crops like wheat, tomatoes, cassava, etc. [6]. Cassava is a crucial staple crop that produces starch-rich, tuberous roots, especially in sub-Saharan African smallholder farming systems [7]. Partly due to its resilience to drought, it has earned the title “drought’, ‘war’, and famine’ crop” [8]. Although cassava yield decreases noticeably during dry seasons, its relationship with and reliance on AMF may be partially responsible for its relatively strong resistance to drought [9]. Approximately 80% of angiosperms have connections made by AMF with their roots and exchange carbon molecules for water and vital nutrients including phosphorus and nitrogen for the host plant [10]. Additionally, they give increased resistance to both biotic and abiotic stressors [9, 10].

There are reports that AMF plays a significant role in improving cassava productivity [8, 9, 10]. Hence, cassava growers could considerably benefit from this technique because cassava is a crop that relies heavily on mycorrhizal fungi for the extraction of nutrients. On the symbiotic interaction between AMF and cassava, the most recent research trends were covered in this review. In addition, we examined several recent studies and extrapolated findings to improve the uses, advantages (Figure 1), and limitations of using AMF in cassava cultivation. The objective of this review is to highlight recent discoveries in a plant-AMF relationship that are associated with cassava crop production. This will contribute to developing a lasting solution to the global problem associated with poor quality and quantity in cassava production, hence solving the problem of hunger and poverty globally, especially in Southern Africa.

Figure 1.

An illustration of the potential for arbuscular mycorrhizal fungal colonization in managing soil–plant nutrients, biotic stressors, abiotic stressors, and reduction of greenhouse emissions.

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2. Database retrieval strategy

The literature review was done electronically utilizing ‘Google Scholar’, ‘PubMed’, ‘Scientific Electronic Library Online (SciELO)’, ‘cassavabase’, and the ‘Scopus databases. The articles to be examined were initially chosen using the following keyword combinations: Arbuscular mycorrhizal fungi, plant-pathogen relationship, cassava mycorrhizae; plant pathogen; abiotic factors; biotic factors, salinity; heavy metals. The majority of the chosen research articles (80% of all references) were published between 2018 and 2022. Given its relevance in the history of mycorrhizae, an old publication from 1885 was added.

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3. The significance of arbuscular mycorrhizal fungi (AMF)

AMF is the most common type of endophytic fungus to colonize the root of its host [6]. Their positive impacts on the survival of their host are widely documented, and these effects include but are not limited to nutrient uptake, assisting with soil aggregation, and resilience to biotic and abiotic stress [6, 7, 8, 9]. As a result, AMF symbioses have a major impact on the yield of the host and the health of the surrounding soil in both natural and laboratory settings [10]. According to Salomon et al. [11], certain AMF species can colonize a host plant and increase the number of its mycorrhizae in organic soils. The AMF’s hyphae graze on the soil’s water and mineral deposits before passing them on to the host plant, ensuring plant production and diversity even in soils low in micronutrients like phosphorus (P) [12]. The root system then passively refluxes these nutrients to the fungus [11]. Consequently, utilizing the soil microbial communities can result in the most sustainable and healthy crop production method. This will help to protect the biosphere by enhancing not just the soil’s fertility even in adverse weather, as well as the plants’ nutrition and health. When these potentials are combined, they would support agriculture and boost global food security [12, 13].

Studies on the interaction of AMF and cassava plants in responding to abiotic and biotic stressors are scarce, despite the extensive study on AMF symbiosis. However, due to the rising and unavoidable stress conditions (with biotic and abiotic factors), it is challenging to meet the global food needs. This study consequently discusses the role of AMF in controlling plant nutrition and growth in response to biotic stresses (plant diseases) as well as abiotic stresses including salinity, water scarcity, floods, high and low temperatures, soil acidity, and soil management, in addition to the involvement of AMF in the sustainable soil management nutrients.

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4. The role of AMF in nutrients management

The arbuscular mycorrhizal fungus can enhance the health of the soil and the plants by synthesizing metabolites and plant growth hormones, increasing the availability of essential vitamins to the plants in nutrient-deficient soils, and providing other ecological services. All these factors are further discussed.

4.1 The function of arbuscular mycorrhizal fungi in soil nutrient uptake

Numerous macro- and micronutrients are required by plants, including nitrogen (N), sodium chloride [NaCl], potassium (K), copper (C), calcium (Ca), iron (Fe), and zinc (Zn), amongst others [14]. Plants often take up these nutrients in an inorganic or fixed form from the soil or the air [6], before transferring them to the host plant. Conventionally, chemically produced fertilizer has been used to meet all the needs of plants because the quality of the soil’s nutrients had decreased owing to overuse and pollution [15]. Unfortunately, the continuous application of these chemicals on farms has led to issues with pollution and the degradation of soil quality [14].

Therefore, the use of AMF in connection with other nutrients-solubilizing or -fixing microorganisms has recently been taken into consideration as a sustainable alternative to soil nourishment [9, 10, 16]. The soil’s nutrients were mobilized to the host plant by AMF, through the mycorrhizal hyphae, which connect the plant roots to the soil. Several investigations into the mechanism of action of nutrient absorption and translocation with AMF have found that:

  1. The mycorrhizal hyphae can stretch and explore a larger region of the ground soil than the roots of the host plant can achieve, thereby enabling it to gain access to both micro and macronutrients that the plant alone is unable to absorb [17]. Hence, AMF can cross the depletion zone caused by the plant’s quick uptake of nutrients in the region around its root system and provide the appropriate nutrient elements to the plant [12].

  2. The fungal hyphae are favorable for nutrient intake because of their small (<10 μm diameter) sizes, which enable them to enter tiny pores that are inaccessible to plant roots, having significant effects on water and micronutrient absorption [12]. Similarly, Püschel [18], reported that the ability of mycorrhizal to modify their hyphal diameter by the size of the soil pore enables them to provide nourishment for plants regardless of soil texture.

Similarly, several researchers documented the beneficial impact of mycorrhizae on cassava plants with nutrient intake [8, 19, 20, 21, 22]. For instance, AMF’s contribution to Phosphate uptake was examined by Ndeko et al. [23]. They discovered that using AMF is suitable for enhancing cassava’s phosphate nutrition in various soil types. The results of their experiment demonstrated that the root abundance and dry weight were increased by the inoculation of an unusual fungal strain (Rhizophagus iregularis). However, following AMF inoculation in unsterilized soil, the root dry weight dropped. They concluded that the Rhizophagus iregularis strain, particularly when the soil is not treated with phosphorus, increases the Phosphate uptake of the cassava plants [23].

Furthermore, the advantages of phosphorus nutrition were revealed using cassava and Rhizophagus irregularis inoculum. To determine whether the paradigm holds in tropical field settings, field tests were carried out in three areas utilizing varied AMF and cassava cultivars in both Kenya and Tanzania at varied Phosphorus fertilization levels. It was discovered that contrary to what the paradigm would have us believe, Cassava’s ability to colonize AMF and respond to inoculation does not necessarily decrease as phosphorus availability rises. The obtained results showed that cassava genotypes and fungal availability play a role in maximizing inoculation responsiveness, which is not always the case in low Phosphorus availability settings [24].

Also, Poku [21], investigated how AMF could help cassava absorb more phosphorus from the soil. Phosphorus-fertilizer, AMF, Phosphorus + AMF, and Untreated (Control) were the four treatments used. In comparison to Phosphorus-fertilizer treatment plots, the Phosphorus + AMF were significantly (p0.05) taller, and the Control and AMF-treated plots were significantly (p0.05) identical. At all four locations, the percentage of leaf Phosphorus was statistically comparable, with a grand mean was 0.4%. The content of Phosphorus in the leaves was significantly raised to 0.5% by adding AMF and Phosphorus + AMF to the soil. In comparison to control plots, tubers taken from Phosphorus + AMF-treated plot lines were meaningfully longer (p 0.05). when compared to the control samples, tuber length rose in plots treated with Phosphorus and AMF. The tuber yields on all soil treatments were higher than on control-treatment plots by a substantial amount (p 0.05). Phosphorus+AMF treated plots and AMF treated plots, however, had considerably higher values than P than the control plots. According to this study, cassava yield can be increased by utilizing AMF or Phosphorus+AMF in comparison to Phosphorus alone or untreated control plants and this can be used to maximize tuber yield [21].

Furthermore, in the work of Lopes et al. [25], they sought to ascertain whether co-inoculating micro-propagated cassava with AMF (Glomus clarum) and PGPBs (Plant growth promoting bacteria) improved greenhouse growth. Inoculated PGPB strains in the cassava variety “BRA Pretinha III” affected the number of glomerospores and mycorrhizal colonization, while Glomus clarum and PGPBs had synergistic interactions, the Glomus clarum and PGPBs combined inoculation promoted higher performance in cassava development with time like all the variants examined. Hence, Co-inoculating PGPBs and AMF can meet cassava’s need for nitrogen, therefore, minimizing the need for nitrogen fertilizer [25].

4.2 How arbuscular mycorrhizal fungi contribute to soil aggregation

Sand, silt, and clay particles are bonded together to form aggregates of different sizes, and this arrangement is referred to as the soil structure [26]. Soil aggregation is essential to the health of the entire ecosystem because it serves as a major site for the exchange of water, gaseous, and nutrient flows as well as a significant source of carbon storage [27]. It is believed that fungal hyphae are one of the key binding agents involved in maintaining micro aggregates. However, intensive agricultural practices used today have significantly impacted soil structure by lowering aggregation stability [26, 27], via the following steps.

The first step is that the extraradical hyphae compress the soil physically as they ramify around plant roots, causing clay particles to reorient and ramification in macroaggregate pores [28], thereby affecting the plant water status, and contributing to the soils’ cohesion and strength, particularly in drought conditions [29]. Also, the production of glomalin, which is a hydrophobic glycoprotein generated by AMF hyphae enables the hydrophilic fungal wall to stick to hydrophobic surfaces found on soil particles [30]. Glomalin production also increases carbon storage and availability, which has an impact on the microbial community, aggregate stability, and soil structure. Based on the design of the plant’s roots and how they are connected to the fungus, glomalin promotes aggregation to varying degrees; the greatest impact on macro aggregation was observed with thin host plants’ roots (0.2–1 mm in diameter) [28].

Furthermore, numerous authors have positively confirmed AMF’s ability to lessen soil aggregation’s detrimental impact on plant growth [28, 29, 30, 31], but few studies have explicitly focused on cassava crops. One of these uncommon investigations was conducted by Morris et al. [31]. They were able to assess how AMF altered aggregate turnover durations. They demonstrated how AMF accelerated the production of large macroaggregates and slowed the dissolution of both big and small macroaggregates. In the presence of AMF, macroaggregates turnover increased. The internal aggregate organization suggested that although the accretion of soil to organic materials in the form of micro aggregates is a prevalent process, it is not the only one at work [31].

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5. How arbuscular mycorrhizal fungi manages abiotic stressors

Numerous abiotic stress studies have demonstrated how human activities associated with agriculture (such as irrigation, overuse of chemical fertilizers and pesticides deforestation, and waste material diffusion) have hurt the plant’s growth, health, and output, leading to major production losses [12, 26, 27]. A general route is involved in how plants respond to stress; it begins with the membrane receptor acquiring the stress signal that culminates in the creation of genes, whose byproducts may defend the plant either directly or indirectly [32]. However, numerous investigations on AMF symbiosis have demonstrated that the contributing fungus typically uses several strategies to help the plant resist some abiotic stressors, including but not limited to salinity, heavy metal pollution, and drought [33]. Abiotic stressors are the main obstacle to achieving global food security since they significantly reduce crop production quality and quantity [30, 32, 33].

5.1 Salinity stress

High salt concentrations in the soil make it difficult for roots to draw water from the ground and can be damaging to crops, with adverse effects such as ethylene production, plasmolysis, an unbalanced diet, the inhibition of photosynthesis, and the creation of reactive oxygen species (ROS) [34]. However, one of the ways by which plants manage salinity stress is via osmotic adaptation. A physiological technique employed by crops to sustain a variety of water movements between cells without experiencing turgor or growth decreases is called osmotic adaptation [35]. The buildup of appropriate solutes in plant cells, including proline and glycine betaine, serves as an illustration of this [34]. Furthermore, the existence of salt-tolerant AMF species was demonstrated by several recent scientific studies to alleviate the plant’s salinity stress. The following four AMF properties are specifically mentioned by the researchers as having the capacity to reduce salt stress:

  1. Increased water intake: Mycorrhizal hyphae possesses a stronger capability to penetrate the ground soil, allowing it to absorb more water and lessen the two primary consequences of salinity that jeopardize a plant’s water status—turgor loss and dehydration [36].

  2. Increased mineral uptake: Under osmotic stress, sodium (Na+) levels in the soil are frequently quite high, which has a detrimental effect on some other transporters present in the roots, like the potassium ion (K+) selective channels [37]. However, it was noted that plants connected to AMF showed a rise in total nutrient availability and a large K+ build-up, which assisted the plants in preserving a low Na+/K+ ratio and avoiding harm to their biological processes [37].

  3. Abundant synthesis of suitable organic solutes: AMF-plants produce more proline, glycine, betaine, and sugars, and these chemicals appear to be positively associated with the invasion of fungi [38]. Because they are necessary for AMF’s function in ROS detoxification, regulating membrane structure, and sustaining enzymes and proteins, their synthesis can aid in cellular osmotic adjustment [38].

  4. Increased antioxidant enzyme activity: Research has demonstrated that AMF symbiosis enhances enzyme system performance in ROS detoxification, particularly that of peroxides, superoxide, hydroxyl radicals, and alpha oxygen, whose generation in plants is significantly influenced by stress conditions like salt. In AMF plants, which normally show less oxidative damage, the effects of these chemicals on cell metabolisms, such as DNA damage, the oxidation of polyunsaturated fatty acids in lipids and amino acids in proteins, and the inhibition of enzymes, appear to be less severe [36, 39].

The benefits of arbuscular mycorrhiza fungi (AMF) on cassava plant growth under salinity have rarely been studied in field settings but in controlled settings. An example is a study conducted by Carretero et al. [20]. This study examined the effects of G. intraradices colonization on three cassava cultivars’ biomass and salt tolerance (measured as growth) (SOM-1, 05, and 50). In both AMF-inoculated and non-inoculated cassava cultivars, the survival rate of the root, stem, and leaf development, as well as nutrient accumulation, were assessed in the presence of different sodium chloride concentrations (0, 68.4, or 136.8 mM) in the medium. It was reported that at 136.8 mM of salt, the AMF colonization boosted plant survival and encouraged growth. The SOM-1 cultivars outperformed the other two in terms of salt tolerance. In addition to promoting growth, the G. intraradices-inoculation proved essential for protecting salt-sensitive cassava cultivars (especially in salt-sensitive cultivars). When compared to non-mycorrhizal cultivars growing in the absence of salt, the AMF cultivars grew in 136.8 mM of NaCl2 and produced more dry weight. The results show that AMF-colonization offers a biological process whereby cassava cultivars can increase their salt tolerance and biomass, which, in both low- and high-stress settings, is essential for optimal cassava development [20].

5.2 Drought stress

Drought is another frequent abiotic factor that negatively influences plant growth, survival, and development [40]. The main symptoms of drought stress include wilting as well as a decline in the net rate of photosynthesis, transpiration rate, water usage effectiveness, relative moisture content, and overall chlorophyll content [41]. Also, drought compromises the electron transport system, which results in the generation of activated oxygen and the shutting of plant stomata, consequently reducing CO2 absorption [42].

AMF play the following crucial functions in the response to drought stress; several studies have demonstrated how crucial plant symbiosis is for reducing the detrimental effects of drought:

  1. Regulation of water uptake: To assist in keeping the plant moist, AMF hyphae delve deeper into the soil and scour a large area for water [43].

  2. Osmotic adjustment: The activity of the AMF sustains several activities, including stomatal opening, cellular expansion, and development, which enable the cells to maintain their turgor [44].

  3. Trehalose biosynthesis: Trehalose is a sugar produced by AMF that assists the plant to resist drought stress, maintaining biological nitrogen fixation, and protecting it from a shortage of water [45].

  4. An increase in antioxidant levels: Catalase, peroxidase, and superoxide dismutase, which decrease ROS, and hydroxyl radicals, appear to be present in higher amounts in plants colonized by AMF [46].

  5. Gene expression: AMF typically encourages the expression of some drought-resistant genes such as 1-pyrroline-5-carboxylate synthetase, and 9-cis-epoxycarotenoid dioxygenase genes. Consequently, the stomata are close to stopping water loss, and water movement either within or outside the cell [47].

It was reported that cassava growth could be stimulated under drought stress by inoculating with an AMF. For instance, according to the experiment by Ekanayake et al. [48] the inoculation with G. clarum, and G. mosseae significantly increased the photosynthetic photochemical efficiency of the photosystem 11 light reactions in cassava. The PS11 photochemistry’s maximal quantum yield (Fv/Fm) was similarly significantly decreased (p 0.05) by water stress, whereas AMF inoculation significantly (p 0.05) decreased the negative effects of water stress on cassava cultivars grown under a water deficit regime.

Similarly, Pea et al. [49] investigated whether intraspecific variation in R. irregularis affects the physiological responses of cassava to water stress because it frequently experiences seasonal drought where it is cultivated. Two genetically distinct R. irregularis isolates were inoculated into cassava to promote recovery, which was then subjected to a drought situation before being re-watered. Cassava samples treated with the two distinct fungi had considerably different physiological stress reactions to drought. However, after being re-watered, both plants recovered but at different rates. They concluded that intraspecific genetic variation in AMF considerably affects the physiological reactions of cassava under water stress. This shows the opportunity to increase cassava resistance to water stress by utilizing naturally occurring polymorphism in AMF [49].

5.3 Heavy metal stress

For several enzyme-catalyzed or redox reactions, as well as the metabolism of nucleic acids, plants require specific mineral elements, such as copper (Cu), iron (Fe), nickel (Ni), and zinc (Zn) amongst others [50]. However, high concentrations of these heavy metals, can alter the protein structure or cause mutations in the plant’s genetic makeup [51]. This may cause indications of deficiency such as chlorosis, diminished germination, and slow growth. Diffusion of these heavy metals into the plant’s roots after a groundwater and soil surface deposition is mostly caused by anthropogenic activities including fertilizer application and the diffusion of industrial waste, which pose a significant stressor for these plants [50, 51].

However, AMF had been reported in supporting plants in heavy metal-contaminated environments using these mechanisms.

  1. The AMF hyphae have strong soil exploration abilities and function as a great adsorbent position for the buildup of cations that inhibit the buildup of dangerous metals [52]. Heavy metals are typically precipitated in the extraradical hyphae by AMF-produced proteins termed ‘glomalin’ [51].

  2. The symbiosis also has a localized benefit in the soil where AMF generates exudates that contain citric acid, lactic acid, etc. By forming compounds with the metals, these organic acids lower their intensity in the soil [53].

Recently, the benefits of AMF as reviewed by Riaz et al. [52], highlighted their potential as plant-based remediation techniques for extremely heavy metal-contaminated soils [52]. Additionally, according to Dushenkove et al. [54], certain plants, like cassava, can remove heavy metals from soils that have been contaminated by crude oil by using high biomass crops in tandem with a system of soil amendments using the rhizofiltration approach. Rhizofiltration is a technique that uses plants to remove toxins from aqueous streams [54].

For the first time, the capacity of cassava to phytoextract mercury (Hg) and gold (Au) from biosolids and mine tailings that contain these metals was successfully proven [55]. Pre-rooted cassava cuttings with 5–7 nodes were grown in a blend of 25% mine tailings and 75% biosolids. Plant cuttings were additionally grown in hydroponics samples containing Hg and/or Au to gauge the two metals’ root uptake. Up to 12.59 g/kg of Hg and 1.89 g/kg of Au were discovered in the cassava’s fibrous roots. Hence, due to the cultivation simplicity, cassava provides a sustainable choice for Hg removal and Au recuperation.

5.4 Temperatures stress

Increased temperature due to several factors including global warming has a significant impact on the cycling of nutrients because temperature regulates both the decomposition of organic materials and soil microbial activity [56]. In AMF, reduced spore diameter, root colonization, species variety, and soil glomalin content have all been linked to higher temperatures, resulting in varying water and N levels impacting the plants’ growth [57]. Reduced mycorrhizal reliance had an impact on the growth of mycobionts, particularly in AMF even while plants could obtain the necessary nutrients. AMF has been reported to act as a buffer for plants that were affected by climate change by widening their “niche,” [58].

Also, low soil temperature inhibits AMF root growth much like high soil temperature does [33]. Even though the ideal temperature for AMF development varies according to the fungal isolates, most species prefer temperatures between 18 and 30OC [59]. At 15°C, mycorrhization of roots declines, and a further drop in temperature (10°C or 5°C) prevents AMF production [33]. AMF sporulation was impacted by low temperatures [12]. For instance, R. intraradices’ sporulation slows down at 15°C, although spores’ metabolic processes are not affected until temperatures of 10°C or lower [60]. Organic additions may improve AMF’s effectiveness in reducing cold stress. An example is the development of Lolium perenne L. on salty soils at low temperatures was enhanced by the synergistic inoculation of R. intraradices and biochar [61].

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6. How arbuscular mycorrhizal Fungi manages biotic stressors

6.1 Resistance to pathogens

In the agricultural industry, AMF can be utilized as a biological agent against many plant diseases, providing an effective substitute for chemical pesticides by fostering sustainability and lowering the hazards to the general populace’s health [62]. It is known that AMF symbiosis helps to lessen the harm done by a variety of soil-borne diseases, such as nematodes and fungi that are responsible for significant production losses [63]. The following are the biological control mechanisms of AMF against pathogens:

  1. Modifications in root development and morphological characteristics: AMF colonization alters the dynamics of pathogens and microbial populations and may stimulate microbiota elements that have antagonistic activity toward specific root illnesses [64].

  2. Modifications in the nutrition of the plant host: The pathogens’ reduction of root biomass or function is made up for by the plant’s increased vigor and the ensuing rise in resilience brought on by the AMF symbiosis [33].

  3. Competition for colonization spots and photosynthates: Root diseases and AMF compete for the carbon molecules that get to the root. Both are dependent on host photosynthates. The increasing carbon demand, though, might prevent pathogen growth since AMF have main access to photosynthates [63].

  4. The activation of the host defense mechanisms: AMF colonization causes the activation of the host defense mechanisms. Examples of these defense mechanisms are the synthesis of Phytoalexins, phenylpropanoid pathway enzymes, chitinases, b-1,3-glucanases, peroxidases, pathogenesis-related (PR) proteins, callose, hydroxyproline-rich glycoproteins (HRGP), and phenolics are amongst the biological controls that the host plant produces in response to AMF [65].

For instance, the capacity of two local AMF species (A. Colombiana and A. appendicula) to promote growth and enhance resistance to root-knot nematode and water stresses in the cassava Yayo cultivar was examined using both single and multiple inoculations in a greenhouse setting. It was determined that of the two AMF species, A. colombiana greatly boosted cassava growth and drought tolerance. However, A. colombiana and A. appendicula both gave cassava plants bioprotective properties against the nematode, including tolerance or resistance [22]. Finally, Investigations into the effects of the ACMV (African Cassava Mosaic Virus) on AMF root colonization and leaf symptoms were conducted. The results revealed that both mycorrhized and non-mycorrhized plants had comparable colonization parameters, and ACMV infection after mycorrhization establishment, had no effect on the AMF root colonization [66].

6.2 The advantages of arbuscular mycorrhizal Fungi interaction with other beneficial soil microorganisms

Several soil microorganisms engage in interactions with AMF [67]. The interactions of AMF with other microorganisms could either be positive, neutral, or negative [68]. They could help plants acquire nutrients, biologically regulate pathogens that cause root infections, and increase soil quality and plant resistance to abiotic stress [65, 68]. Similar to how AMF and plant growth-promoting Rhizobacteria (PGPR) collaborate to aid the growth of the host plant [67]. While AMF and nitrogen-fixing bacteria provide plants with essential soil nutrients, More et al.’s [69] hypothesis states that co-inoculating PGPR and AMF will have the greatest synergistic effects. This notion was confirmed by the interactions in their experiment between AMF and modifying rhizobial bacteria [69, 70].

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7. The function of arbuscular mycorrhizal fungi in reducing greenhouse gas emissions

Reduced emissions of greenhouse gases and boosted carbon sequestration are the two primary goals the Food and Agriculture Organization (FAO) has developed for climate-smart agriculture [71]. N2O is a potent greenhouse gas with a larger global warming potential (280–310) than CO2 and longer persistence (118–131 years) in the atmosphere and sadly, agriculture is a significant source of N2O emissions [72]. Dissimilatory nitrate reduction to ammonium and ammonia oxidation are amongst the main sources of N2O emissions in farms through a variety of denitrification processes. Slow-growing nitrifiers are outcompeted by AMF hyphae for ammonium, which hinders nitrification and, ultimately, the production of N2O [73].

Under varying moisture conditions, the emissions of greenhouse gases from soil can also be controlled by AMF (as well as the water relationship between plants) [11]. The impact of AMF in controlling N2O emissions has not been studied in cassava, but it has been studied in legume systems. A field experiment with AMF on legumes revealed a significant reduction in yield-scaled N2O emissions due to greater biological nitrification inhibition [74]. The term “biological nitrification inhibition” (BNI) refers to the ability of the host plant’s roots to lower the activity of soil nitrifiers by generating and releasing nitrogen removal inhibitors. To manage the soil nitrifier activities, help reduce emissions of greenhouse gases and generally make agriculture more ecologically friendly and effective, BNI-enabled plants and pastures have been suggested for use in agriculture [72, 73].

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8. Limitations

Given the projected rise in population, it will be necessary to enhance the production of economically important crops like cassava. Hence, increasing soil fertility in an environmentally sustainable approach may be the greatest strategy to accomplish the expected crop yield. Given the effects of climate change, nutrient depletion, drought, salt, and metal toxicity, it has been well established that AMF may sustainably improve plant growth, production, and crop nutritional quality for this purpose, thereby ensuring food security for both present and future generations.

However, most studies on AMF-mediated advances in plant health, nutrient uptake, and regulation of the impacts of biotic and abiotic stress conditions have thus far used in vitro studies on laboratory scales, and under controlled temperature and low nutrients profiles trials studies. However, if more farm field experiments are done, the results may offer a more in-depth understanding of the intricate mechanisms underpinning Plant-nutrient connections that are mediated by AMF and AMF interactions with both the biotic and abiotic stressors. These understandings may have an advantage to the effective application of AMF in boosting the production of more important commercial crops, such as cassava, regardless of the region.

Also, the manipulation of AMF in both artificial and natural systems has proven to be a particularly difficult barrier to overcome. It is anticipated that future tests would use cutting-edge techniques to understand the mystery surrounding plants-AMF community ecology, what patterns do plants and AMF communities follow? what does this mean in terms of ecosystem management, recovery, and restoration? To get answers to these issues, we both hope and anticipate that we will not have to wait another 13 years.

Additionally, the glycoproteins (glomalin) produced by AMF are not currently considered in soil health assessment programs even though they are linked to several ecosystem processes and act as a low-cost proxy for soil quality, the efficiency of agricultural management and restoration methods, and AMF biomass [75]. Controlled studies using these methods (ELISA and advanced spectroscopic classification) and AMF quantification (quantitative PCR assays, AMF-signature lipids, microscopic measurements) are necessary to inform researchers and farmers about the potential of these glycoproteins as a C-sequestration and quality indicator, along with field demonstrations.

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9. Conclusions

Mycorrhizae and their usage in the agronomic field have gained popularity in numerous scientific studies conducted all over the world in recent years. Most of the research has been on the advantages that arbuscular fungus provides for host plants regarding its productivity and resistance to all biotic and abiotic challenges. Numerous studies of the fungus’ metabolic processes and pathways have been conducted to enhance the fungus’ capacity for nutrient and water absorption as well as its defense against infections, salt, and heavy metals.

The cassava plant, one of the most significant food crops in the world, has been the centre of numerous studies on mycorrhizal inoculation. However, choosing cassava varieties with very effective mycorrhizal symbiosis may serve as the cornerstone to produce foods in low-impact agricultural systems. The useful rhizosphere microbial inoculants are now widely used in agricultural techniques for many crops than cassava. However, there might also be negative interactions, including rivalry and hostility, between various rhizosphere microbe kinds. It is counterproductive when specific indigenous soil microbes interact negatively with externally applied microorganisms. Hence, choosing the most contagious and effective mycorrhizal fungi and cassava together will make it easier to employ them as biofertilizers to replace the lost biological fertility of the soil, reduction of chemical usage and mitigate the effects of biotic and abiotic stress.

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Acknowledgments

Dr. Otun acknowledges the University of Witwatersrand, the SARChi programme of the Department of Science and Technology and the South African National Research Foundation for her postdoctoral fellowship funding.

Conflict of interest

The authors declare no conflict of interest.

References

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Written By

Sarah Otun and Ikechukwu Achilonu

Submitted: 15 August 2022 Reviewed: 15 September 2022 Published: 22 March 2023