Mycorrhizal Fungi and Sustainable Agriculture

*Soibam Helena Devi, Ingudam Bhupenchandra, Soibam Sinyorita, S.K. Chongtham and E. Lamalakshmi Devi*

#### **Abstract**

The 20thcentury witnessed an augmentation in agricultural production, mainly through the progress and use of pesticides, fertilizers containing nitrogen and phosphorus, and developments in plant breeding and genetic skills. In the naturally existing ecology, rhizospheric soils have innumerable biological living beings to favor the plant development, nutrient assimilation, stress tolerance, disease deterrence, carbon seizing and others. These organisms include mycorrhizal fungi, bacteria, actinomycetes, etc. which solubilize nutrients and assist the plants in up taking by roots. Amongst them, arbuscular mycorrhizal (AM) fungi have key importance in natural ecosystem, but high rate of chemical fertilizer in agricultural fields is diminishing its importance. The majority of the terrestrial plants form association with Vesicular Arbuscular Mycorrhiza (VAM) or Arbuscular Mycorrhizal fungi (AMF). This symbiosis confers benefits directly to the host plant's growth and development through the acquisition of Phosphorus (P) and other mineral nutrients from the soil by the AMF. They may also enhance the protection of plants against pathogens and increases the plant diversity. This is achieved by the growth of AMF mycelium within the host root (intra radical) and out into the soil (extra radical) beyond. Proper management of Arbuscular Mycorrhizal fungi has the potential to improve the profitability and sustainability of agricultural systems. AM fungi are especially important for sustainable farming systems because AM fungi are efficient when nutrient availability is low and when nutrients are bound to organic matter and soil particles.

**Keywords:** Actinomycetes, Bacteria, Mycorrhizal fungi, Vesicular Arbuscular Mycorrhiza

#### **1. Introduction**

The productivity of agriculture has increased steadily since the middle of the last century in temperate regions and, more recently, in tropical areas [1] mainly due to improved varieties, machinery, fertilizers, and pesticides. However, recently agricultural yields have plateaued [2], largely because of land degradation often associated with unsustainable farming practices [3, 4]. To meet future demand, food production must rise significantly through this century, utilizing less cultivable land in a sustainable manner [5]. Many factors have contributed to making this increase in crop production essential, including population increases worldwide, food shortages in developing countries, the importance of agricultural products

for trade in many countries, the competition between urbanization and agriculture for land, and the competition that is likely to develop between food and non-food production [6]. However, the mechanism available for producers to increase yields tends to increase input costs, the number of operations in the field, and the financial risks of failure. At the same time environmental degradation linked to the use of chemical inputs (i.e., water pollution from nitrates, phosphates, and pesticides) is increasingly widespread and sometimes irreversible. Moreover, secondary effects on biogenesis and soil impoverishment have weakened cropping systems to make them increasingly dependent on chemicals [7]. Thus, there is growing demand for clean agriculture, high-quality food, and more information on how food is produced are finally having an effect on decreasing the level of chemical inputs used in developed countries [8]. However, in developing countries the great need for food implies a trend towards intensification, mainly through the use of more fertilizers [9] as plant nutrition has played a key role in the dramatic increase in meeting the demand for and supply of food. The consumption of Nitrogenous (N) fertilizer has increased almost nine-fold and that of Phosphorus (P) more than four-fold. The tremendous increase of N and P fertilizers in addition to the introduction of highly productive and agricultural systems has allowed these developments to occur at relatively low costs [10]. But the increasing use of fertilizers and highly productive system have also created environmental problems such as deterioration of soil quality, surface water and ground water as well as air pollution, reduced biodiversity and suppressed ecosystem function [10, 11]. Environmental pollution resulting from greater nutrient availability can be either direct or indirect. Directly, misuse and excessive or poorly managed use of fertilizers can result in leaching, volatilization, acidification and denitrification. Indirectly, the production (use of fossil fuel in Haber-Bosch process) and transport (combustion of fossil fuel) of fertilizer result in air- borne carbon dioxide and nitrogen pollution, which will be eventually deposited into terrestrial ecosystems.

The most limiting nutrients for plant growth are N and P. Most of N is tied into soil organic matter. Even after fertilization, plants have to compete with soil microbes for easily available soluble N but problem with P are different. In acidic soils, even when phosphorus fertilizer is added in substantial quantities, it becomes non available as fertilizers P precipitates with iron or aluminum whereas in alkaline soils P precipitates as calcium phosphates [12]. Accordingly, P limitation may be a difficult problem to overcome through the addition of P-containing fertilizers. So, the recent increase in crop yields and food production in developed countries have been achieved by intensive agricultural practices. This increase, however have not come without tremendous environmental costs. In developing countries, the problems are different. The lack of fertilizers and adequate agricultural practices do not allow intensive crop production and a vast segment of the population remains undernourished. Clearly, there is an urgent need for sustainable agricultural practices on a global level. In the developed world a reduction of energy and environmental costs is necessary. In developing countries, efficient, sustainable practices are needed to allow cost efficient production of adequate nutrition for the growing populations. To overcome the ecological problems resulting from the loss of plant nutrients and to increase crop yields in the absence of resources for obtaining costly fertilizers, microscopic organisms that allow more efficient nutrient use or increase nutrient availability can provide sustainable solutions for present and future agricultural practices. Therefore, a better knowledge of the processes and factors that govern the bioavailability of soil nutrients to plants, thus including the root-soil interactions understanding of microorganisms in the rhizosphere [12, 13] is necessary. As Samuil [14] mentioned, organic farming promotes sustainable production systems, diversified and balanced crop, to prevent pollution and the environment

**187**

plant health.

carbon in their fungal partners [34].

*Mycorrhizal Fungi and Sustainable Agriculture DOI: http://dx.doi.org/10.5772/intechopen.99262*

nents of agricultural 'sustainable intensification' [5].

**2. Arbuscular mycorrhizae symbiosis**

damages. Moreover, the importance of the mycorrhizal arbuscular fungi in organic farming and farmers' potential to increase the benefits of arbuscular mycorrhizae (AM) associations in such systems represented interesting subjects as it was synthesized by Gosling *et al.* [15]. Symbiotic soil organisms, such as mycorrhizal fungi, may be the source of many of these beneficial effects and thereby be key compo-

The term 'Mycorrhiza' was first introduced by Frank [16] and comprises of all symbiotic associations of soil-borne fungi with roots or rhizoids of higher plants. Allen [17] described the fungal-plant interaction from a more neutral or microbially oriented aspect stating that 'Mycorrhiza is a mutualistic symbiosis between plant and fungus localized in a root or root-like structure in which energy moves primarily from plant to fungus and inorganic resources move from fungus to plant'. The group of fungi and plants, which are involved in the interaction, determines the type of mycorrhiza they form [18]. Recently, there have been significant advances in the understanding of physiological processes and taxonomy of these fungi [19, 20]. They are obligate symbionts belonging to the phylum Glomeromycota [21]. Their activity in agricultural ecosystems is well documented [22–24]. The distribution of ectomycorrhizal (ECM) fungi is also widespread, but they form associations with only 3% of terrestrial plant families [25]. ECM fungi are members of the phyla Ascomycota and Basidiomycota [26, 27]. Unlike the ECM fungi, AM fungi are dependent on plants for their carbon (C) and when a symbiosis is formed, both ECM and AM fungi can demand 20–40% of photosynthetically fixed plant C [28] (**Figure 1a** and **b**). Soil microorganisms have significant impact on soil fertility and

Microbial symbionts including arbuscular mycorrhizal (AM) fungi form an essential component of the soil microbial community playing a key role in overall plant growth and development (**Figure 1c**). Arbuscular mycorrhizal (AM) fungi form symbiotic relationships with over 80% of terrestrial plant species [32]. Arbuscular mycorrhizas are ancient and ubiquitous symbioses formed between a relatively small group of soil fungi and higher plant roots which has been traced back 460 million years [21]. During AM symbiosis, the fungal hyphae penetrate the root cortical cell walls by formation of aspersoria leading to the development of intra-radical hyphal colonization and formation of arbuscules or coils that interface with the host cytoplasm [33]. The highly branched arbuscules aid in metabolic exchanges between the plant and the fungus. AM fungi also produce vesicles, which function as storage organs [33]. It has been estimated that in natural ecosystems plants colonized with AM fungi may invest 10–20% of the photo-synthetically fixed

AM fungi not only can promote via directs effects, but there are also a number of indirect effects such as a stimulation of soil quality and the suppression of organisms that reduce crop productivity (**Table 1**) [35]. AM fungi also interface directly with the soil by producing extra-radical hyphae that may extend several centimeters out into the soil thereby helping the host plants in uptake of nutrients especially P [36]. The extra-radical mycelium of AM fungi can also enhance mobilization of organically bound nitrogen (N) from plant litter [37]. Hyphae of AM fungi have been shown to play an important role in soil stabilization through formation of soil aggregates by secretion of glomalin [38]. Glomalin is a glycoprotein produced on hyphae of AM in the soil. It is discovered by [39] and termed as glomalin, after the source organism of phylum "Glomeromycota." It is apparently insoluble in water;

*Mycorrhizal Fungi and Sustainable Agriculture DOI: http://dx.doi.org/10.5772/intechopen.99262*

*Nitrogen in Agriculture - Physiological, Agricultural and Ecological Aspects*

deposited into terrestrial ecosystems.

for trade in many countries, the competition between urbanization and agriculture for land, and the competition that is likely to develop between food and non-food production [6]. However, the mechanism available for producers to increase yields tends to increase input costs, the number of operations in the field, and the financial risks of failure. At the same time environmental degradation linked to the use of chemical inputs (i.e., water pollution from nitrates, phosphates, and pesticides) is increasingly widespread and sometimes irreversible. Moreover, secondary effects on biogenesis and soil impoverishment have weakened cropping systems to make them increasingly dependent on chemicals [7]. Thus, there is growing demand for clean agriculture, high-quality food, and more information on how food is produced are finally having an effect on decreasing the level of chemical inputs used in developed countries [8]. However, in developing countries the great need for food implies a trend towards intensification, mainly through the use of more fertilizers [9] as plant nutrition has played a key role in the dramatic increase in meeting the demand for and supply of food. The consumption of Nitrogenous (N) fertilizer has increased almost nine-fold and that of Phosphorus (P) more than four-fold. The tremendous increase of N and P fertilizers in addition to the introduction of highly productive and agricultural systems has allowed these developments to occur at relatively low costs [10]. But the increasing use of fertilizers and highly productive system have also created environmental problems such as deterioration of soil quality, surface water and ground water as well as air pollution, reduced biodiversity and suppressed ecosystem function [10, 11]. Environmental pollution resulting from greater nutrient availability can be either direct or indirect. Directly, misuse and excessive or poorly managed use of fertilizers can result in leaching, volatilization, acidification and denitrification. Indirectly, the production (use of fossil fuel in Haber-Bosch process) and transport (combustion of fossil fuel) of fertilizer result in air- borne carbon dioxide and nitrogen pollution, which will be eventually

The most limiting nutrients for plant growth are N and P. Most of N is tied into soil organic matter. Even after fertilization, plants have to compete with soil microbes for easily available soluble N but problem with P are different. In acidic soils, even when phosphorus fertilizer is added in substantial quantities, it becomes non available as fertilizers P precipitates with iron or aluminum whereas in alkaline soils P precipitates as calcium phosphates [12]. Accordingly, P limitation may be a difficult problem to overcome through the addition of P-containing fertilizers. So, the recent increase in crop yields and food production in developed countries have been achieved by intensive agricultural practices. This increase, however have not come without tremendous environmental costs. In developing countries, the problems are different. The lack of fertilizers and adequate agricultural practices do not allow intensive crop production and a vast segment of the population remains undernourished. Clearly, there is an urgent need for sustainable agricultural practices on a global level. In the developed world a reduction of energy and environmental costs is necessary. In developing countries, efficient, sustainable practices are needed to allow cost efficient production of adequate nutrition for the growing populations. To overcome the ecological problems resulting from the loss of plant nutrients and to increase crop yields in the absence of resources for obtaining costly fertilizers, microscopic organisms that allow more efficient nutrient use or increase nutrient availability can provide sustainable solutions for present and future agricultural practices. Therefore, a better knowledge of the processes and factors that govern the bioavailability of soil nutrients to plants, thus including the root-soil interactions understanding of microorganisms in the rhizosphere [12, 13] is necessary. As Samuil [14] mentioned, organic farming promotes sustainable production systems, diversified and balanced crop, to prevent pollution and the environment

**186**

damages. Moreover, the importance of the mycorrhizal arbuscular fungi in organic farming and farmers' potential to increase the benefits of arbuscular mycorrhizae (AM) associations in such systems represented interesting subjects as it was synthesized by Gosling *et al.* [15]. Symbiotic soil organisms, such as mycorrhizal fungi, may be the source of many of these beneficial effects and thereby be key components of agricultural 'sustainable intensification' [5].

#### **2. Arbuscular mycorrhizae symbiosis**

The term 'Mycorrhiza' was first introduced by Frank [16] and comprises of all symbiotic associations of soil-borne fungi with roots or rhizoids of higher plants. Allen [17] described the fungal-plant interaction from a more neutral or microbially oriented aspect stating that 'Mycorrhiza is a mutualistic symbiosis between plant and fungus localized in a root or root-like structure in which energy moves primarily from plant to fungus and inorganic resources move from fungus to plant'. The group of fungi and plants, which are involved in the interaction, determines the type of mycorrhiza they form [18]. Recently, there have been significant advances in the understanding of physiological processes and taxonomy of these fungi [19, 20]. They are obligate symbionts belonging to the phylum Glomeromycota [21]. Their activity in agricultural ecosystems is well documented [22–24]. The distribution of ectomycorrhizal (ECM) fungi is also widespread, but they form associations with only 3% of terrestrial plant families [25]. ECM fungi are members of the phyla Ascomycota and Basidiomycota [26, 27]. Unlike the ECM fungi, AM fungi are dependent on plants for their carbon (C) and when a symbiosis is formed, both ECM and AM fungi can demand 20–40% of photosynthetically fixed plant C [28] (**Figure 1a** and **b**). Soil microorganisms have significant impact on soil fertility and plant health.

Microbial symbionts including arbuscular mycorrhizal (AM) fungi form an essential component of the soil microbial community playing a key role in overall plant growth and development (**Figure 1c**). Arbuscular mycorrhizal (AM) fungi form symbiotic relationships with over 80% of terrestrial plant species [32]. Arbuscular mycorrhizas are ancient and ubiquitous symbioses formed between a relatively small group of soil fungi and higher plant roots which has been traced back 460 million years [21]. During AM symbiosis, the fungal hyphae penetrate the root cortical cell walls by formation of aspersoria leading to the development of intra-radical hyphal colonization and formation of arbuscules or coils that interface with the host cytoplasm [33]. The highly branched arbuscules aid in metabolic exchanges between the plant and the fungus. AM fungi also produce vesicles, which function as storage organs [33]. It has been estimated that in natural ecosystems plants colonized with AM fungi may invest 10–20% of the photo-synthetically fixed carbon in their fungal partners [34].

AM fungi not only can promote via directs effects, but there are also a number of indirect effects such as a stimulation of soil quality and the suppression of organisms that reduce crop productivity (**Table 1**) [35]. AM fungi also interface directly with the soil by producing extra-radical hyphae that may extend several centimeters out into the soil thereby helping the host plants in uptake of nutrients especially P [36]. The extra-radical mycelium of AM fungi can also enhance mobilization of organically bound nitrogen (N) from plant litter [37]. Hyphae of AM fungi have been shown to play an important role in soil stabilization through formation of soil aggregates by secretion of glomalin [38]. Glomalin is a glycoprotein produced on hyphae of AM in the soil. It is discovered by [39] and termed as glomalin, after the source organism of phylum "Glomeromycota." It is apparently insoluble in water;

#### **Figure 1.**

*(a) Life cycle of an AM fungus and the different steps during AM development (Adated from Bücking et al. [29]). (b). A-F: The structural colonization of AMF in roots [a]: Vesicle and mycelium occurrence of the AMF-colonized root sections (arrow), [B]: Spore morphology of AMF (arrow), [C]: Intact mycorrhizal spores and subtending hyphae (arrow), [D]: Vesicles and subtending hyphae of AMF (arrow), [E]: Vesicles and subtending hyphae (arrow), [F]: Trunk, arbuscle and hyphae of AMF (arrow). (adapted from Abeer [30]. (c) Diagram of a root colonized by AM fungi (adapted from Habte, M., and R.L. Fox. [31].*


#### **Table 1.**

*Direct and indirect effects of mycorrhizal fungi on crop productivity in organic farming systems [35].*

extremely persistent glycoproteinaceous compound [40] thus remains in soil after the death of the hyphae [41]. It improves soil physical properties, carbon sequestration, mineral elements, microbes' activities, stabilizes pollutants, and ultimately helping in restoration of soil ecosystem. This kind of beneficial effect of glomalin may be through its existing impact on soil; by serving as a substrate for microbial population, a gluing mediator for aggregate formation, chelation of heavy metals

**189**

*Mycorrhizal Fungi and Sustainable Agriculture DOI: http://dx.doi.org/10.5772/intechopen.99262*

(SYM genes) [45].

**3. Application of AMF in agriculture**

**3.1 AMF enhances soil fertility**

*3.1.1 Uptake of phosphorous*

cal region of the roots [53].

relationship and the growth environment [43, 44].

and toxic pollutants, and enhancing carbon sequestration via long-term persistence in soil [42]. The indirect or 'secondary' impacts of glomalin on the formation and stabilization of soil aggregates further improved the efficiency of the symbiotic

In addition to increasing the absorptive surface area of their host plant root systems, the extra-radical hyphae of AM fungi provide an increased area for interactions with other microorganisms, and an important pathway for the translocation of energy-rich plant assimilates to the soil. The symbiosis is primarily characterized by its association with phosphorus (P) uptake by host plants and the enhancement of water uptake through the extra radical fungal hyphal networks. This symbiosis can also trigger physiological and molecular signals at subcellular levels, alter plant community structure and increase plant tolerance to various abiotic and biotic stresses. Mycorrhizal hyphal networks link plants of the same and different species below ground and are able to transfer resources between plants and release signal molecule defense-related proteins, lipochito oligosaccharides and strigolactones. Molecules like strigolactones secreted by the roots help fungi identify their host plants. Strigolactones also stimulate AM fungal growth and its branching. The fungi reciprocate to this signal by secreting a set of hypothetical factors known as Mycorrhizal Factors (Myc). These factors also play a major role in communication between AM fungi and nitrogen-fixing bacteria. The AM interactions are established further with the induction of seven genes

Microbes inhabiting in the rhizospheric region interact with the plant root system and generate a nutrient-rich situation improving crop development [46]. At the same time such microorganisms also protect the plant root from harmful chemicals either by repelling them or by diluting their impacts in rhizosphere. Symbiotic association between a particular group of fungus with the plant root system enhances the availability of essential mineral elements principally those elements which are less mobile (phosphorus, copper, zinc etc.) in soil thus allowing crop roots system to access them at ease, which are otherwise difficult to uptake because of their less mobility [47, 48]. Such association also helps crops to establish even in soil with poor fertility, as the microbes will help in uptake of essential nutrients, both major and minor [49]. Studies reported that the amount of fertilizer (particularly phos-

phatic) used can be decreased with the appliance of mycorrhizae [50, 51].

P is a critical and macro essential element (**Figure 2**). It performs an important function in all biological system as it is involved in all energy transfer processes in the form of ATP. Moreover, it also acts as an indispensable constituent for various biological molecules like nucleotides, phospholipids and sugar phosphates [52]. One of the significant advantages of mycorrhizal application is the augment in the P nutrition to the crop. The normal mechanism of P uptake may be discussed in these following ways-; (i) AMF hyphae will absorb it (P) from the rhizosphere, (ii) the absorbed P will be translocated along the hyphae from outside to inner (root cortex) mycelium, (iii) ultimately the absorbed phosphorous is transferred to corti*Mycorrhizal Fungi and Sustainable Agriculture DOI: http://dx.doi.org/10.5772/intechopen.99262*

*Nitrogen in Agriculture - Physiological, Agricultural and Ecological Aspects*

extremely persistent glycoproteinaceous compound [40] thus remains in soil after the death of the hyphae [41]. It improves soil physical properties, carbon sequestration, mineral elements, microbes' activities, stabilizes pollutants, and ultimately helping in restoration of soil ecosystem. This kind of beneficial effect of glomalin may be through its existing impact on soil; by serving as a substrate for microbial population, a gluing mediator for aggregate formation, chelation of heavy metals

*Direct and indirect effects of mycorrhizal fungi on crop productivity in organic farming systems [35].*

Weed suppression

(green manure)

Stimulation of nitrogen fixation by legumes

Suppression of some soil pathogens Stimulation of soil biological activity Increased soil carbon storage Reduction of nutrient leaching

Stimulation of soil aggregation and soil structure

*(a) Life cycle of an AM fungus and the different steps during AM development (Adated from Bücking et al. [29]). (b). A-F: The structural colonization of AMF in roots [a]: Vesicle and mycelium occurrence of the AMF-colonized root sections (arrow), [B]: Spore morphology of AMF (arrow), [C]: Intact mycorrhizal spores and subtending hyphae (arrow), [D]: Vesicles and subtending hyphae of AMF (arrow), [E]: Vesicles and subtending hyphae (arrow), [F]: Trunk, arbuscle and hyphae of AMF (arrow). (adapted from Abeer [30].* 

*(c) Diagram of a root colonized by AM fungi (adapted from Habte, M., and R.L. Fox. [31].*

*Direct effects on crops Indirect effects*

• Stimulation of plant productivity of various crops

• Nutrient acquisition (P, N, Cu, Fe, Zn) • Enhanced seedling establishment

• Drought resistance • Heavy metal resistance

**188**

**Table 1.**

**Figure 1.**

and toxic pollutants, and enhancing carbon sequestration via long-term persistence in soil [42]. The indirect or 'secondary' impacts of glomalin on the formation and stabilization of soil aggregates further improved the efficiency of the symbiotic relationship and the growth environment [43, 44].

In addition to increasing the absorptive surface area of their host plant root systems, the extra-radical hyphae of AM fungi provide an increased area for interactions with other microorganisms, and an important pathway for the translocation of energy-rich plant assimilates to the soil. The symbiosis is primarily characterized by its association with phosphorus (P) uptake by host plants and the enhancement of water uptake through the extra radical fungal hyphal networks. This symbiosis can also trigger physiological and molecular signals at subcellular levels, alter plant community structure and increase plant tolerance to various abiotic and biotic stresses. Mycorrhizal hyphal networks link plants of the same and different species below ground and are able to transfer resources between plants and release signal molecule defense-related proteins, lipochito oligosaccharides and strigolactones. Molecules like strigolactones secreted by the roots help fungi identify their host plants. Strigolactones also stimulate AM fungal growth and its branching. The fungi reciprocate to this signal by secreting a set of hypothetical factors known as Mycorrhizal Factors (Myc). These factors also play a major role in communication between AM fungi and nitrogen-fixing bacteria. The AM interactions are established further with the induction of seven genes (SYM genes) [45].

#### **3. Application of AMF in agriculture**

#### **3.1 AMF enhances soil fertility**

Microbes inhabiting in the rhizospheric region interact with the plant root system and generate a nutrient-rich situation improving crop development [46]. At the same time such microorganisms also protect the plant root from harmful chemicals either by repelling them or by diluting their impacts in rhizosphere. Symbiotic association between a particular group of fungus with the plant root system enhances the availability of essential mineral elements principally those elements which are less mobile (phosphorus, copper, zinc etc.) in soil thus allowing crop roots system to access them at ease, which are otherwise difficult to uptake because of their less mobility [47, 48]. Such association also helps crops to establish even in soil with poor fertility, as the microbes will help in uptake of essential nutrients, both major and minor [49]. Studies reported that the amount of fertilizer (particularly phosphatic) used can be decreased with the appliance of mycorrhizae [50, 51].

#### *3.1.1 Uptake of phosphorous*

P is a critical and macro essential element (**Figure 2**). It performs an important function in all biological system as it is involved in all energy transfer processes in the form of ATP. Moreover, it also acts as an indispensable constituent for various biological molecules like nucleotides, phospholipids and sugar phosphates [52]. One of the significant advantages of mycorrhizal application is the augment in the P nutrition to the crop. The normal mechanism of P uptake may be discussed in these following ways-; (i) AMF hyphae will absorb it (P) from the rhizosphere, (ii) the absorbed P will be translocated along the hyphae from outside to inner (root cortex) mycelium, (iii) ultimately the absorbed phosphorous is transferred to cortical region of the roots [53].

#### **Figure 2.**

*Mycorrhizal and non-mycorrhizal phosphorous uptake pathways. In mycorrhizal roots, the P-depletion zone is extended by extraradical fungal hyphae beyond that of the non-mycorrhizal root and root hairs (adapted from smith and smith [52]).*

A variety of techniques were anticipated in order to increase the accessibility to essential elements such as (i) enhanced assessment of soil; (ii) better movement of phosphate into roots via arbuscules; (iii) alteration of root growing region; (iv) proficient exploitation of phosphate inside crops; (v) proficient transportation of Phosphorous to crop root systems; and (vi) improved storing of captivated Phosphorous. Bhat and Kaveriappa [54] reported that the uptake of PO4 −3 by root systems occur more rapidly as compared to the rate at which the ions diffuse to absorbing the site the root system. Thus, it may result in a PO4 −3 exhaustion region in the roots surroundings. However, the vast expanded mycorrhizal hyphae lengthen into the surroundings region of plant roots scaffoldingPO4 −3 exhaustion region. Consequently, improves the ability of the crop to venture their growing region farther than the nutrient exhaustion region where rootlets and root hair cannot grow [55]. An enzyme known as phosphatase is responsible for converting organic Phosphorous into absorbable form and the action of this enzyme is used by mycorrhizae in order to metabolized PO4 −3 in the soil. This enzyme is found inside polyphosphate granules residing in the vacuoles of the fungus. The action of this enzyme will break the polyphosphate granules in fine branches of ultimately the mineralized are released in the protoplasm.

The AMF possibly will give a P uptake route; however, AMF colonization can lower the crop acquisition of PO4 −3 via their root PO4 −3 transporter [56]. If this reduction is not compensated via AMF pathway of PO4 −3 acquisition then it may result in lowering the ultimate cropPO4 −3 uptake in AMF-colonized crops [56]. Dissimilarity in the proportionate contributions of these PO4 −3 delivery routes can be accountable for the differences in P absorption amongst crop cultivars. In some crop families the AMF has lately been exposed to give a large enhancement on their development example *Alliaceae, Fabaceae* and the *Solanaceae* [57], however the response of other plants, particularly grass family, are ambiguous [58]. Additionally, dissimilarity in growth due to AMF colonization too occurs amongst cultivars within species [59].

#### *3.1.2 Uptake of nitrogen*

N is an essential component of amino acid and nitrogenous bases therefore, it is necessary though in direct for protein and nucleic acid bio-synthesis. AMF

**191**

form NO3

−

hormones [67, 68].

**3.3 Uptake of water**

**3.2 Plant growth hormones**

*Mycorrhizal Fungi and Sustainable Agriculture DOI: http://dx.doi.org/10.5772/intechopen.99262*

colonized crops shown to have improved N content in their above ground part. Various processes may be implied for this improvement, viz. (i) enhancement of symbiotic fixing of atmospheric N; (ii) direct acquisition of combined N by mycorrhiza; (iii) crops without nodule are also benefited as a portion of N fixed by the plants with nodule is transferred towards the former; (iv) activity enzymes viz. pectinase, xyloglucanase and cellulose which are necessary for N metabolism as well as decomposing soil organic matter where enhanced [53]. AMF hyphae have the capacity to extort N from the soil and transporting it to crops. They possess the enzymes to facilitate breaking of organic nitrogen and have N reductase that transform the N present in the rhizosphere. AM not only promotes growth, nodulation and N fixing process in legume-*Rhizobium* symbiosis but also enhances acquisition of ammonia easily from soil which represent the major contribution of accessible N in numerous innate environments. Bacteria which can fix N thus distinctly have the capacity to impact AM fungi. Minerdi *et al.* [60] reported the occurrence of genes responsible for fixing of N in endosymbiotic *Burkholderia* sp., however, significant expression of this genes so that it can impact the growth of the fungal association has not been confirmed. *Rhizobium spp*. might perform collegially with AM fungi on their plant hosts. Nodulation as well as fixing of N are normally augmented in legumes subsequently after AM colonization, possibly for the reason that the fungus provides access of P to the plant and the rhizobacteria, which is necessary for the enzymes essential in the fixing of N. Supplementary fixing of N also improves growth of mycorrhizae [61]. In soils where major fraction of N is available in the

, AMF have a mere impact in uptake of N by legumes [62]. AMF hyphae

enhance translocation of N in crop colony, as the connection of AM mycelia joins varied crops species thriving close by and helps pooling and enhancing availability of nutrients for these crops. McFarland *et al.* [63] reported that above 50% of N

Fungi associated with a pants produced crop growth regulators [64]. Arbuscular

AMF may also have a vital position in the improving water economy. The AMF symbiosis either augments the hydraulic conductivity of the roots thereby improving water absorption by the crops or modifies the crop physiology so as to decrease the stress response to soil drought [69]. In severe arid situation, crops associated with mycorrhizae have superior endurance than those crops without the association. It was revealed that mycelial complex may lengthen beneath and broader the soil profile in explore of water and essential mineral elements. Mycorrhizal

mycorrhizae develop a reciprocal and advantageous symbiosis with the majority land plants as their association is not species specific. The existence of host plant at the time of beginning and during the progress of symbiosis is essential, as their symbiotic relation is obligatory in nature. However, AM fungal spores have the capacity to germinate even if the host plant is not present [65]. The fungal counterpart can sense the incidence of the host plant via some signal transduction. Amongst the signaling compounds, that can influence mycorrhizal associations are plant hormones, which might definitely or negatively influence the associations [66]. Plant hormones can manage the intensity and therefore specificity of mycorrhizal associations. Consequently, it might be probable to improve the effectiveness of mycorrhizal association during stresses by regulating the amount of stress

required by the crop is by associating with mycorrhizal.

#### *Mycorrhizal Fungi and Sustainable Agriculture DOI: http://dx.doi.org/10.5772/intechopen.99262*

*Nitrogen in Agriculture - Physiological, Agricultural and Ecological Aspects*

A variety of techniques were anticipated in order to increase the accessibility to essential elements such as (i) enhanced assessment of soil; (ii) better movement of phosphate into roots via arbuscules; (iii) alteration of root growing region; (iv) proficient exploitation of phosphate inside crops; (v) proficient transportation of Phosphorous to crop root systems; and (vi) improved storing of captivated

*Mycorrhizal and non-mycorrhizal phosphorous uptake pathways. In mycorrhizal roots, the P-depletion zone is extended by extraradical fungal hyphae beyond that of the non-mycorrhizal root and root hairs (adapted from* 

systems occur more rapidly as compared to the rate at which the ions diffuse to absorb-

roots surroundings. However, the vast expanded mycorrhizal hyphae lengthen into the

improves the ability of the crop to venture their growing region farther than the nutrient exhaustion region where rootlets and root hair cannot grow [55]. An enzyme known as phosphatase is responsible for converting organic Phosphorous into absorbable form and the action of this enzyme is used by mycorrhizae in order to metabolized PO4

the soil. This enzyme is found inside polyphosphate granules residing in the vacuoles of the fungus. The action of this enzyme will break the polyphosphate granules in fine

The AMF possibly will give a P uptake route; however, AMF colonization can

for the differences in P absorption amongst crop cultivars. In some crop families the AMF has lately been exposed to give a large enhancement on their development example *Alliaceae, Fabaceae* and the *Solanaceae* [57], however the response of other plants, particularly grass family, are ambiguous [58]. Additionally, dissimilarity in growth due to AMF colonization too occurs amongst cultivars within species [59].

N is an essential component of amino acid and nitrogenous bases therefore, it is necessary though in direct for protein and nucleic acid bio-synthesis. AMF

−3 via their root PO4

−3 by root

−3 in

−3 exhaustion region in the

−3 exhaustion region. Consequently,

−3 transporter [56]. If this reduc-

−3 acquisition then it may result in

−3 delivery routes can be accountable

−3 uptake in AMF-colonized crops [56]. Dissimilarity

Phosphorous. Bhat and Kaveriappa [54] reported that the uptake of PO4

branches of ultimately the mineralized are released in the protoplasm.

ing the site the root system. Thus, it may result in a PO4

surroundings region of plant roots scaffoldingPO4

tion is not compensated via AMF pathway of PO4

in the proportionate contributions of these PO4

lower the crop acquisition of PO4

lowering the ultimate cropPO4

*3.1.2 Uptake of nitrogen*

**190**

**Figure 2.**

*smith and smith [52]).*

colonized crops shown to have improved N content in their above ground part. Various processes may be implied for this improvement, viz. (i) enhancement of symbiotic fixing of atmospheric N; (ii) direct acquisition of combined N by mycorrhiza; (iii) crops without nodule are also benefited as a portion of N fixed by the plants with nodule is transferred towards the former; (iv) activity enzymes viz. pectinase, xyloglucanase and cellulose which are necessary for N metabolism as well as decomposing soil organic matter where enhanced [53]. AMF hyphae have the capacity to extort N from the soil and transporting it to crops. They possess the enzymes to facilitate breaking of organic nitrogen and have N reductase that transform the N present in the rhizosphere. AM not only promotes growth, nodulation and N fixing process in legume-*Rhizobium* symbiosis but also enhances acquisition of ammonia easily from soil which represent the major contribution of accessible N in numerous innate environments. Bacteria which can fix N thus distinctly have the capacity to impact AM fungi. Minerdi *et al.* [60] reported the occurrence of genes responsible for fixing of N in endosymbiotic *Burkholderia* sp., however, significant expression of this genes so that it can impact the growth of the fungal association has not been confirmed. *Rhizobium spp*. might perform collegially with AM fungi on their plant hosts. Nodulation as well as fixing of N are normally augmented in legumes subsequently after AM colonization, possibly for the reason that the fungus provides access of P to the plant and the rhizobacteria, which is necessary for the enzymes essential in the fixing of N. Supplementary fixing of N also improves growth of mycorrhizae [61]. In soils where major fraction of N is available in the form NO3 − , AMF have a mere impact in uptake of N by legumes [62]. AMF hyphae enhance translocation of N in crop colony, as the connection of AM mycelia joins varied crops species thriving close by and helps pooling and enhancing availability of nutrients for these crops. McFarland *et al.* [63] reported that above 50% of N required by the crop is by associating with mycorrhizal.

#### **3.2 Plant growth hormones**

Fungi associated with a pants produced crop growth regulators [64]. Arbuscular mycorrhizae develop a reciprocal and advantageous symbiosis with the majority land plants as their association is not species specific. The existence of host plant at the time of beginning and during the progress of symbiosis is essential, as their symbiotic relation is obligatory in nature. However, AM fungal spores have the capacity to germinate even if the host plant is not present [65]. The fungal counterpart can sense the incidence of the host plant via some signal transduction. Amongst the signaling compounds, that can influence mycorrhizal associations are plant hormones, which might definitely or negatively influence the associations [66]. Plant hormones can manage the intensity and therefore specificity of mycorrhizal associations. Consequently, it might be probable to improve the effectiveness of mycorrhizal association during stresses by regulating the amount of stress hormones [67, 68].

#### **3.3 Uptake of water**

AMF may also have a vital position in the improving water economy. The AMF symbiosis either augments the hydraulic conductivity of the roots thereby improving water absorption by the crops or modifies the crop physiology so as to decrease the stress response to soil drought [69]. In severe arid situation, crops associated with mycorrhizae have superior endurance than those crops without the association. It was revealed that mycelial complex may lengthen beneath and broader the soil profile in explore of water and essential mineral elements. Mycorrhizal

association might also adjust the selectivity of plasma membrane to water, with enhanced P nourishment thus these AMF association could enhance the drought resistance of crops [70, 71]. During the situation of water stress, AMF apply their impact by enhancing the transpirational rate and enhancing stomatal conductance or by modifying the equilibrium of plant hormones [72]. The alteration in leaf flexibility owing to AMF association improve H2O and turgor pressure of leaves and also enhance root growth and extension [73] thus they might affect H2O relations and consequently, the water stress resistance of the crops. The possible basis for the improved moisture and nutrient absorption by mycorrhiza-crops association may be owed to improved allocation of absorbing hyphal system, more constructive ideotype of hyphae as compared to roots, superior absorbing area and rapid expansion rate, improved serviceable longevity, chemical modification in soil rhizosphere thus modifying microorganism inhabiting in the sphere, uptake kinetics, superior hydraulic conductivities, reduce transportation rates per unit leaf area, withdrawal of moisture from soil towards less ᴪw and more swift revival form moisture stress.

#### **3.4 Improvement of soil texture and structure**

Interruption in ecology may influence the physical and bio-chemical activities in the soil. AMF assist in the fastening of soil particles and improving aggregation of soil and thus aiding conservation of soil [74]. Mycorrhizal fungi also enhance soil quality, because they generate glomalin subsequently when this molecule get accumulated in soil, then with fungal hyphae they produce micro aggregate and ultimately macro aggregates thereby, acting directly as a skeleton for soil aggregation and stabilization. It also produces discharge in the soil consequently improves and stabilizes soil and boosting the growth of microbes [34].

#### **3.5 AMF in plant defense**

AMF association can improve host plant endurance to pests and pathogens both above- and below-ground (**Figure 3**) [75–77]. Although the clear-cut biological and chemical reasons are uncertain, their colonization is identified to stimulate subtle modifications in plant metabolic process, like jasmonate and salicylic acid signaling routes, which are vital constituents of crop defense mechanism [78]. Mycorrhizal colonization of a plant before they are attacked by pathogen or pest may have a systemic priming effect via defense molecules/signal re-allocation [79]. So, crop defense related DNA can be expressed more rapidly and extensively than in non-colonized plants [80]. Mycorrhizal colonization may promote crop nutrients uptake and growth; however, this improvement may result in making the plant more attractive, enhance quantity and more nutritive to herbivores [81], thus promoting herbivore action [82]. In fact, phloem-feeding insect normally have enhanced performance on AMF-colonized crops than non-AM crops [81, 83]. Therefore, stabilizing these compromises ought to represent a key module of every farming management tactic associating AMF. Some plants may perhaps display a nonspecific method of defense against phloem-feeding herbivore, facilitated by AMF symbionts. Aphid-infested crops can interact signals via common mycelial networks (CMN) to an uninfected plant (**Figure 3**) [84], therefore a quick flux can be elicited for aphid-repellent volatiles compounds prior to the attack of the plant, this can check the widening or severity of invasion and consequent reduction in productivity [85].

Organic management of crop diseases is one of the important mechanisms for improving crop productivity either by repressing or killing the disease-causing microbes, thus augmenting the capability of crops to endure pathogens or by defending crops against disease causing microbes. In this context mycorrhizal

**193**

plant pathogens:

*Thirkell et al. [73]).*

**Figure 3.**

pathogen [88].

of *Douglas fir* [90].

*Mycorrhizal Fungi and Sustainable Agriculture DOI: http://dx.doi.org/10.5772/intechopen.99262*

association proves to be the organisms which may serve as bio protectors of crops [86]. Several mechanisms might be involved in mycorrhizal symbiosis in regulating

*Mechanisms by which arbuscular mycorrhizal fungi (AMF) may influence agroecosystem physiological ecology. (1) Common mycelia network transmits signal, altering volatile organic compounds (VOCs) in neighboring plants, repelling herbivores and attracting parasitoids (blue arrows), (2) AMF induce VOCs, repelling herbivores (yellow arrows), and (3) AMF elicit systemic plant defense priming (red arrows). (Adapted from* 

expansion of disease-causing microbes [87].

courage the infestation of microbes [91].

against disease causing microbes [92].

root zone than the microbes [93].

I.Generating a physical fence and obstructing the infiltration and successive

II.Making the cell wall thicker via deposition of lignin and by producing many complex carbohydrates which consecutively hamper the penetration of root

III.Activating host plants specially in the roots to synthesize and concentrate adequate amount of biochemical compounds (alkaloids etc.) which improve

the host ability to resist against microbes invasion in their tissue [89].

V.Augmenting the amount of orthodihydroxy phenols in roots, which dis-

VI.Synthesizing antifungal and antibacterial antibiotics and toxins function

VII.Augmenting competitive advantage to host root for nutrients element in the

consequently forbidding the pathogens to acquire entrance to the roots [94].

VIII.Increasing microbial functioning and competition in the rhizosphere

preventing lesion development by the pathogen *Fusarium oxgsporum* in roots

IV.Promoting flavonoid wall infusions eg. *Laccaria bicolor*, subsequently

*Mycorrhizal Fungi and Sustainable Agriculture DOI: http://dx.doi.org/10.5772/intechopen.99262*

#### **Figure 3.**

*Nitrogen in Agriculture - Physiological, Agricultural and Ecological Aspects*

**3.4 Improvement of soil texture and structure**

**3.5 AMF in plant defense**

and stabilizes soil and boosting the growth of microbes [34].

severity of invasion and consequent reduction in productivity [85].

Organic management of crop diseases is one of the important mechanisms for improving crop productivity either by repressing or killing the disease-causing microbes, thus augmenting the capability of crops to endure pathogens or by defending crops against disease causing microbes. In this context mycorrhizal

association might also adjust the selectivity of plasma membrane to water, with enhanced P nourishment thus these AMF association could enhance the drought resistance of crops [70, 71]. During the situation of water stress, AMF apply their impact by enhancing the transpirational rate and enhancing stomatal conductance or by modifying the equilibrium of plant hormones [72]. The alteration in leaf flexibility owing to AMF association improve H2O and turgor pressure of leaves and also enhance root growth and extension [73] thus they might affect H2O relations and consequently, the water stress resistance of the crops. The possible basis for the improved moisture and nutrient absorption by mycorrhiza-crops association may be owed to improved allocation of absorbing hyphal system, more constructive ideotype of hyphae as compared to roots, superior absorbing area and rapid expansion rate, improved serviceable longevity, chemical modification in soil rhizosphere thus modifying microorganism inhabiting in the sphere, uptake kinetics, superior hydraulic conductivities, reduce transportation rates per unit leaf area, withdrawal of moisture from soil towards less ᴪw and more swift revival form moisture stress.

Interruption in ecology may influence the physical and bio-chemical activities in the soil. AMF assist in the fastening of soil particles and improving aggregation of soil and thus aiding conservation of soil [74]. Mycorrhizal fungi also enhance soil quality, because they generate glomalin subsequently when this molecule get accumulated in soil, then with fungal hyphae they produce micro aggregate and ultimately macro aggregates thereby, acting directly as a skeleton for soil aggregation and stabilization. It also produces discharge in the soil consequently improves

AMF association can improve host plant endurance to pests and pathogens both above- and below-ground (**Figure 3**) [75–77]. Although the clear-cut biological and chemical reasons are uncertain, their colonization is identified to stimulate subtle modifications in plant metabolic process, like jasmonate and salicylic acid signaling routes, which are vital constituents of crop defense mechanism [78]. Mycorrhizal colonization of a plant before they are attacked by pathogen or pest may have a systemic priming effect via defense molecules/signal re-allocation [79]. So, crop defense related DNA can be expressed more rapidly and extensively than in non-colonized plants [80]. Mycorrhizal colonization may promote crop nutrients uptake and growth; however, this improvement may result in making the plant more attractive, enhance quantity and more nutritive to herbivores [81], thus promoting herbivore action [82]. In fact, phloem-feeding insect normally have enhanced performance on AMF-colonized crops than non-AM crops [81, 83]. Therefore, stabilizing these compromises ought to represent a key module of every farming management tactic associating AMF. Some plants may perhaps display a nonspecific method of defense against phloem-feeding herbivore, facilitated by AMF symbionts. Aphid-infested crops can interact signals via common mycelial networks (CMN) to an uninfected plant (**Figure 3**) [84], therefore a quick flux can be elicited for aphid-repellent volatiles compounds prior to the attack of the plant, this can check the widening or

**192**

*Mechanisms by which arbuscular mycorrhizal fungi (AMF) may influence agroecosystem physiological ecology. (1) Common mycelia network transmits signal, altering volatile organic compounds (VOCs) in neighboring plants, repelling herbivores and attracting parasitoids (blue arrows), (2) AMF induce VOCs, repelling herbivores (yellow arrows), and (3) AMF elicit systemic plant defense priming (red arrows). (Adapted from Thirkell et al. [73]).*

association proves to be the organisms which may serve as bio protectors of crops [86]. Several mechanisms might be involved in mycorrhizal symbiosis in regulating plant pathogens:


Roots associating with VAM/AM fungi might serve as a site for actinomycetes accumulation thus opposing root pathogens [95].


Reduced pathogen growth in mycorrhizal and non-mycorrhizal portions of infected roots is connected with aggregation of phenolic and plant cell resistance mechanism. It has been observed that mycorrhizal infected crops were shown to be more lenient to fungal root pathogen in the presence of the root modulating symbiont viz. *Rhizobium leguminosarum*. Thus, indicating that interaction between mycorrhizae and other microorganism thriving in the rhizoplane is more effective in warding off the soil-borne pathogens than the mycorrhizae alone. Therefore, it is essential to understand of the mechanisms of plant disease resistance in mycorrhizae inoculated plants so as to get improved guidelines for developing an efficient plant production and sustainable agriculture. The compositions with mycorrhizae are extremely essential to produce novel groups of biocides which can also give a reduction in risk to both humane healthiness and the ecology [98].

#### **4. AM fungi and sustainable agriculture**

For sustainability in agriculture sector, it is essential to exploit the innate mechanisms so as to realize an adequate level of yield and food quality at the same time reducing chemicals inputs, reducing input overheads and obviate ecological contamination and its impact [27, 99]. However, it must be viable in the existing ecosystem and socially accountable. Edaphic factors have numerous contributions towards achieving sustainability in agricultural system such as controlling of pathogen in rhizosphere and augmenting the growth of soil microbes and their activeness which enhanced antagonistic and parasitic interaction in the crop root growing region [100, 101]. Plant and microbe interactions were the intriguing events contributing in agricultural sustainability. Mycorrhizal fungi are native to soil and crop rhizoplane thus making them as an essential mechanism for sustainability in agricultural. In sustainable agricultural systems specially when there are shorts supply for essential elements the mycorrhizae association turn out to play a vital role. In these conditions AM extra-radical mycelium have a vital position for mobilizing the nutrient components into usable form. Hamel and Strullu [102] stated analyzing an AM fungi have been a tough job, however nowadays they are identified as vital ingredients of edaphic environments rather than only crop root constituent. AM fungi can impact plant development, nutrient supply and production even in P-riched soils [103, 104] but the positive response of AM fungi will be more pronounced when the crop is grown in less fertile soil agro ecosystem and also resulting in reduced environmental nutrient loss.

Therefore, attention in AM fungal proliferation for sustainable agricultural system is escalating owing to its function in the encouragement of plant vigor, and enhancement in soil quality and structural stability. These fungal associations could be used efficiently for escalating productivity whilst decreasing the utilization of chemical inputs such as insecticides, fertilizers etc. To enhance crop productivity in

**195**

*Mycorrhizal Fungi and Sustainable Agriculture DOI: http://dx.doi.org/10.5772/intechopen.99262*

and may lessen both biotic and abiotic stresses [33, 71, 110].

as per the climate, soil and existing markets.

**5. Conclusion**

soil with low fertility, inorganic fertilizers have been extensively employed, organic matter is included and techniques like keeping the soil fallow or incorporation leguminous crops have been practiced to improve soil ecosystem, boost soil microbial growth and functioning and augment nutrient re-cycling in order to decrease external inputs and make the most of their utility [105]. This mechanism has been initiated for managing soil ecosystem using earthworms and micro-symbionts [106, 107]. These soil organisms might contribute above 90% of soil organic activity consequently aiding the courses like nutrient cycling, soil fertility and symbiotic association in the plant root zone. Soil fungal multiplicity and functions have not been sufficiently investigated and inferred [108]. Mycorrhizae serve as a significant faction since they have an extensive dissemination and might have significant contribution to biomass of soil microbes and to soil nutrient cycling processes in crops. For sustainable improvement in crop yield more dependence should be on natural and organic mechanism by acclimating on germplasm favorable to plants [109]. They advance absorption of essential elements, particularly phosphorous, and minor nutrients like Zn, Cu; they influence the production of growth material

In order to have sustainability agricultural system, administration of appropriate nutrient supply represents necessary condition and in this situation mycorrhizae involvement cannot be ignored. Native mycorrhizae spores attack the plant roots and contribute in sustainability nutrients management, moisture, soil properties and productivity [48]. With increasing concern and demand on the requirement to augment sustainability in agricultural development, mycorrhizae fungi possess a significant part to perform in decreasing the detrimental consequent of chemical inputs of agriculture viz. pesticides, synthetic nutrients for promoting plant growth and regulating numerous pathogens. It is an economic and non-detrimental way of acquiring higher yield which may leads to development of a feasible, minimuminput cropping system. It is expected that, in the coming days, limitation in crop cultivation in the globe may be bypassed by using the techniques based on organic processes like mycorrhizae fungi. In the circumstance of existing limitation associated to large population, food requirement and ecological changes, it is obvious that the recent development and progress in arbuscular mycorrhiza field research and those in the future should be considered on maximizing production, improving its quality, enhancing cost effectiveness and profit return, conserving biodiversity and protection of environment. These limitations can be attained by interacting different discipline in particular ways to generate an interdisciplinary connection and association of all researchers in the field [111]. Mattoo and Teasdale [112] stated that sustainable agriculture systems be likely to meet the common necessities in terms of, productivity efficiency and management by confined appliance of the principles

Presently, it is probable the beneficial effect of AMF colonization is not nutritional, through their effect on soil aggregates and activity and on plant defenses. Therefore, research in the coming days must focus on improving AM effects on nutrient acquisition so that productivity is continued at the same time as optimizing the sustainability of production. By recognizing and improving those character linked with AMF accessibility, functionality and climate resilience (e.g. water stress tolerance) in new cultivars, considerable development can be made towards acquir-

ing food supply in coming days in more sustainable agricultural system.

#### *Mycorrhizal Fungi and Sustainable Agriculture DOI: http://dx.doi.org/10.5772/intechopen.99262*

soil with low fertility, inorganic fertilizers have been extensively employed, organic matter is included and techniques like keeping the soil fallow or incorporation leguminous crops have been practiced to improve soil ecosystem, boost soil microbial growth and functioning and augment nutrient re-cycling in order to decrease external inputs and make the most of their utility [105]. This mechanism has been initiated for managing soil ecosystem using earthworms and micro-symbionts [106, 107]. These soil organisms might contribute above 90% of soil organic activity consequently aiding the courses like nutrient cycling, soil fertility and symbiotic association in the plant root zone. Soil fungal multiplicity and functions have not been sufficiently investigated and inferred [108]. Mycorrhizae serve as a significant faction since they have an extensive dissemination and might have significant contribution to biomass of soil microbes and to soil nutrient cycling processes in crops. For sustainable improvement in crop yield more dependence should be on natural and organic mechanism by acclimating on germplasm favorable to plants [109]. They advance absorption of essential elements, particularly phosphorous, and minor nutrients like Zn, Cu; they influence the production of growth material and may lessen both biotic and abiotic stresses [33, 71, 110].

### **5. Conclusion**

*Nitrogen in Agriculture - Physiological, Agricultural and Ecological Aspects*

to damage by microbes attacked [96].

plant roots exudates [97].

**4. AM fungi and sustainable agriculture**

resulting in reduced environmental nutrient loss.

cetes accumulation thus opposing root pathogens [95].

reduction in risk to both humane healthiness and the ecology [98].

For sustainability in agriculture sector, it is essential to exploit the innate mechanisms so as to realize an adequate level of yield and food quality at the same time reducing chemicals inputs, reducing input overheads and obviate ecological contamination and its impact [27, 99]. However, it must be viable in the existing ecosystem and socially accountable. Edaphic factors have numerous contributions towards achieving sustainability in agricultural system such as controlling of pathogen in rhizosphere and augmenting the growth of soil microbes and their activeness which enhanced antagonistic and parasitic interaction in the crop root growing region [100, 101]. Plant and microbe interactions were the intriguing events contributing in agricultural sustainability. Mycorrhizal fungi are native to soil and crop rhizoplane thus making them as an essential mechanism for sustainability in agricultural. In sustainable agricultural systems specially when there are shorts supply for essential elements the mycorrhizae association turn out to play a vital role. In these conditions AM extra-radical mycelium have a vital position for mobilizing the nutrient components into usable form. Hamel and Strullu [102] stated analyzing an AM fungi have been a tough job, however nowadays they are identified as vital ingredients of edaphic environments rather than only crop root constituent. AM fungi can impact plant development, nutrient supply and production even in P-riched soils [103, 104] but the positive response of AM fungi will be more pronounced when the crop is grown in less fertile soil agro ecosystem and also

Therefore, attention in AM fungal proliferation for sustainable agricultural system is escalating owing to its function in the encouragement of plant vigor, and enhancement in soil quality and structural stability. These fungal associations could be used efficiently for escalating productivity whilst decreasing the utilization of chemical inputs such as insecticides, fertilizers etc. To enhance crop productivity in

Roots associating with VAM/AM fungi might serve as a site for actinomy-

IX.Balancing the trade-off in nutrient uptake mechanism of roots caused due

X.Altering the quantity and nature of chemical produces by plant root and this would be difficulty for that pathogen which depends on specific type

Reduced pathogen growth in mycorrhizal and non-mycorrhizal portions of infected roots is connected with aggregation of phenolic and plant cell resistance mechanism. It has been observed that mycorrhizal infected crops were shown to be more lenient to fungal root pathogen in the presence of the root modulating symbiont viz. *Rhizobium leguminosarum*. Thus, indicating that interaction between mycorrhizae and other microorganism thriving in the rhizoplane is more effective in warding off the soil-borne pathogens than the mycorrhizae alone. Therefore, it is essential to understand of the mechanisms of plant disease resistance in mycorrhizae inoculated plants so as to get improved guidelines for developing an efficient plant production and sustainable agriculture. The compositions with mycorrhizae are extremely essential to produce novel groups of biocides which can also give a

**194**

In order to have sustainability agricultural system, administration of appropriate nutrient supply represents necessary condition and in this situation mycorrhizae involvement cannot be ignored. Native mycorrhizae spores attack the plant roots and contribute in sustainability nutrients management, moisture, soil properties and productivity [48]. With increasing concern and demand on the requirement to augment sustainability in agricultural development, mycorrhizae fungi possess a significant part to perform in decreasing the detrimental consequent of chemical inputs of agriculture viz. pesticides, synthetic nutrients for promoting plant growth and regulating numerous pathogens. It is an economic and non-detrimental way of acquiring higher yield which may leads to development of a feasible, minimuminput cropping system. It is expected that, in the coming days, limitation in crop cultivation in the globe may be bypassed by using the techniques based on organic processes like mycorrhizae fungi. In the circumstance of existing limitation associated to large population, food requirement and ecological changes, it is obvious that the recent development and progress in arbuscular mycorrhiza field research and those in the future should be considered on maximizing production, improving its quality, enhancing cost effectiveness and profit return, conserving biodiversity and protection of environment. These limitations can be attained by interacting different discipline in particular ways to generate an interdisciplinary connection and association of all researchers in the field [111]. Mattoo and Teasdale [112] stated that sustainable agriculture systems be likely to meet the common necessities in terms of, productivity efficiency and management by confined appliance of the principles as per the climate, soil and existing markets.

Presently, it is probable the beneficial effect of AMF colonization is not nutritional, through their effect on soil aggregates and activity and on plant defenses. Therefore, research in the coming days must focus on improving AM effects on nutrient acquisition so that productivity is continued at the same time as optimizing the sustainability of production. By recognizing and improving those character linked with AMF accessibility, functionality and climate resilience (e.g. water stress tolerance) in new cultivars, considerable development can be made towards acquiring food supply in coming days in more sustainable agricultural system.

#### **Author details**

Soibam Helena Devi1 , Ingudam Bhupenchandra<sup>2</sup> \*, Soibam Sinyorita2 , S.K. Chongtham3 and E. Lamalakshmi Devi4

1 Department of Crop Physiology,Assam Agricultural University, Jorhat, India

2 ICAR-KVK Tamenglong, ICAR-RC for NEH Region, Manipur Centre, India

3 Multi Technology Testing Centre and Vocational Training Centre, CAEPHT, CAU, Gangtok, Sikkim, India

4 ICAR-RC for NEH Region, Sikkim Centre, Gangtok, Sikkim, India

\*Address all correspondence to: ibhupenj@gmail.com

© 2021 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

**197**

*Mycorrhizal Fungi and Sustainable Agriculture DOI: http://dx.doi.org/10.5772/intechopen.99262*

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*Mycorrhizal Fungi and Sustainable Agriculture DOI: http://dx.doi.org/10.5772/intechopen.99262*

#### **References**

*Nitrogen in Agriculture - Physiological, Agricultural and Ecological Aspects*

, Ingudam Bhupenchandra<sup>2</sup>

4 ICAR-RC for NEH Region, Sikkim Centre, Gangtok, Sikkim, India

\*Address all correspondence to: ibhupenj@gmail.com

provided the original work is properly cited.

1 Department of Crop Physiology,Assam Agricultural University, Jorhat, India

2 ICAR-KVK Tamenglong, ICAR-RC for NEH Region, Manipur Centre, India

3 Multi Technology Testing Centre and Vocational Training Centre, CAEPHT, CAU,

© 2021 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium,

and E. Lamalakshmi Devi4

\*, Soibam Sinyorita2

,

**196**

**Author details**

S.K. Chongtham3

Soibam Helena Devi1

Gangtok, Sikkim, India

[1] Craswell ET, Karjalainen U. Recent research on fertilizer problems in Asian agriculture. Fertilizer Research. 1990; **26**: 243-248.

[2] Grassini P, Eskridge KM, Cassman KG. Distinguishing between yield advances and yield plateaus in historical crop production trends. Nature Communications. 2013;**4**: 2918.

[3] Bindraban PS, van der Velde M, Ye L,van den Berg M, Materechera S, Kiba DI, Tamene L, Ragnarsdottir KV, Jongschaap REE, Hoogmoed M, Hoogmoed WB, Beek CL, van Lynden GWJ. Assessing the impact of soil degradation on food production. Current Opinion in Environmental Sustainability. 2012;**4**: 478-488.

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conference.html

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[25] Smith SE, Read DJ. Mycorrhizal symbiosis. Academic Press, London.

Donoghue MJ. Evolutionary instability of ectomycorrhizal symbioses in basidiomycetes. Nature. 2000; **407**:

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Akhter MS, Futai K (Eds.) Mycorrhizae: sustainable agriculture and forestry, Springer, Dordrecht, 2008. 2008; 1-35.

rhizosphere – roots, soil and everything

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An Overview. In: Siddiqui ZA,

[28] McNear DH Jr (2013) The

in between. Nature Education Knowledge. 2013;**4**(3):1.

[29] Bücking H, Liepold E and Ambilwade P. The role of the mycorrhizal symbiosis in nutrient uptake of plants and the regulatory mechanisms underlying these transport processes. 2012. IntechOpen. https:// www.intechopen.com/books/

the-role-of-the-mycorrhizal-

Abdallah EF, Egamberdieva D.

symbiosisin-nutrient-uptake-of-plantsand-the-regulatorymechanisms-und

[30] Abeer H, Salwa AA, Alqarawi AA,

Arbuscular mycorrhizal fungi enhance basil tolerance to salt stress through improved physiological and nutritional status. Pakistan Journal Botany. 2016;

[31] Habte, M, Fox RL. Effectiveness of VAM fungi in nonsterile soils before and after optimization of P in soil solution. Plant and Soil. 1993; **151**: 219-226.

plantscience/

**48**(1): 37-45

2014; **68**: 429-439

2008; 800.

506-508.

Bäume durchunterir dische Pilze. https://doi.org/10.1111/j.1438-8677.1885.

[17] Allen MF. The ecology of mycorrhiza. Cambridge: Cambridge

University Press, 1991; 184.

[18] Molina R, Massicotte H, Trappe JM. Specificity phenomena in mycorrhizal symbiosis: Community-ecological consequences and practical implications. In: Allen, M.F. (Ed.), Mycorrhizal Functioning: An integrative plant-fungal process. Chapman and Hall, New York. 1992; 357-423.

[19] Kohout P, Sudova´ R, Janouskova´ M, Ctvrtlı'kova´ M, Hejda M, Pa'nkova´ H, Slavı'kova´ R, Stajerova´ K, Vosa'tka M, Sy'korova´ Z. Comparison of commonly used primer sets for evaluating arbuscular mycorrhizal fungal communities: Is there a universal solution? Soil Biology and Biochemistry.

[20] Saito M. Symbiotic exchange of nutrients in arbuscular mycorrhizas: transport and transfer of phosphorus. In: Douds DD, Kapunik Y (Eds.)

[21] Redecker D. Molecular identification and phylogeny of

view of the formation of VA mycorrhizas. Plant and Soil. 1994;

[23] Bedini S, Avio L, Sbrana C, Turrini A, Migliorini P, Vazzana C, Giovannetti M. Mycorrhizal activity and diversity in a long-term organic Mediterranean agroecosystem. Biology

and Fertility of Soils. 2013; **49**:

Soil. 2002; **244**:67-73.

**159**:69-78.

Arbuscular mycorrhizas: physiology and function. Kluwer, Dordrecht, The Netherlands. 2000; 85-106.

arbuscular mycorrhizal fungi. Plant and

[22] Abbott LK, Gazey C. An ecological

2014; **68**: 482-493

tb04240.x

**198**

781-790.

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[33] Smith SE, Read DJ. Mycorrhizal Symbiosis. Academic Press, San Diego. 1997; 605.

[34] Johnson D, Leake JR, Ostle N, Ineson P, Read DJ. *In situ* 13CO2 pulselabelling of upland grasslands demonstrates a rapid pathway of carbon flux from arbuscular mycorrhizal mycelia to the soil. New Phytologist. 2002*;***153**: 327- 334*.*

[35] Van der Heijden MGA, Rinaudo V, Verbruggen E, Scherrer C, Bàrberi P, Giovannetti M. The significance of mycorrhizal fungi for crop productivity and ecosystem sustainability in organic farming systems. 16thIFOAM Organic World Congress, Modena, Italy, June 16-20, 2008. Archived at http:// orgprints.org/view/projects/ conference.html

[36] Read DJ, Perez-Moreno J. 2003. Mycorrhizas and nutrient cycling in ecosystems – a journey towards relevance? New Phytologist. 2003; **157**: 475-492.

[37] Hodge A, Campbell CD, Fitter AH. An arbuscular mycorrhizal fungus accelerates decomposition and acquires nitrogen directly from organic matter. Nature. 2001; **413**: 297-299.

[38] Tisdall JM,Oades JM. Stabilization of soil aggregates by the root systems of rye grass. Australian Journal of Soil Research. 1979; **17**: 429-441.

[39] Wright S F, Upadhyaya A 1996. Extraction of an abundant and unusual protein from soil and comparison with hyphal protein from arbuscular mycorrhizal fungi. Soil Science. 161: 575-586.

[40] Wright S F, Upadhyaya A. A survey of soils for aggregate stability and

glomalin, a glycoprotein produced by hyphae of arbuscular mycorrhizal fungi. Plant and Soil. 1998; 198:97-107.

[41] Driver, James D, Holben, William E, Rillig, Matthias C. Characterization of glomalin as a hyphal wall component of arbuscular mycorrhizal fungi. Soil Biology and Biochemistry. 2005; **37**(1): 101-106. https://doi.org/10.1016/j. soilbio.2004.06.011.

[42] Singh AK, Zhu X,Chen C, Wu J ,Yang B , Zakari S , Jiang XJ , Singh N, Liu W The role of glomalin in mitigation of multiple soil degradation problems. Critical Reviews in Environmental Science and Technology; 2020. doi=10.1 080/10643389.2020.1862561

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[45] Bonfante P, Genre A. Mechanism underlying beneficial plant fungus interactions in mycorrhizal symbiosis. Nature Communications. 2010; **1**(8). https://doi.org/10.1038/ncomms1046

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[57] Elbon A, Whalen JK. Phosphorus supply to vegetable crops from

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Delhi. 2007; 295-309.

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calcium spiking responses to the fungus. Frontier of Plant Science. 2013; **4:** 426.

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Science. 2006; **86**: 941-950.

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doi: 10.3389/fpls.2013.00426

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[105] Sanchez PA. Properties

York: Wiley- Interscience. 1994.

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science. 2004; **60**: 149-157.

**102**: 144-153.

*Mycorrhizal Fungi and Sustainable Agriculture DOI: http://dx.doi.org/10.5772/intechopen.99262*

*Nitrogen in Agriculture - Physiological, Agricultural and Ecological Aspects*

[89] Sampangi RK. Some recent advances in the study of fungal root diseases. Indian Phytopathology. 1989;

[90] Strobel NE, Sinclair WA. Role of flavonilic wall inclusions in the

[91] Krishna KR, Bagyaraj DJ.

and Sclerotium *rolfsii* in peanut. Canadian Journal of Botany. 1984;

[92] Marx DH. Ectomycorrhizae as biological deterents to pathogenic root

infections. Annual Review of Phytopathology. 1972; **10**: 429-454.

[93] Reid CPP. Mycorrhizas. In:

Wiley and Sons.1990; 281-315.

as determinant of rhizospheric microbial diversity. In: Mukerji KG, Manoharachary C, Singh J (Eds.), Microbial activity in the rhizosphere. Springer Verlag, Berlin, Heidelberg.

Lynch JM. (Ed.), The Rhizosphere, John

[94] Singh G, Mukerji KG. Root exudates

[95] Mukerji KG. Rhizosphere Biology. In: Mukerji KG, Manoharachary C, Chamola BP (Eds.), Techniques in Mycorrhizal Studies, Kluwer Academic Publishers, Dordrecht, The Netherlands.

[96] Smith SE, Reid DJ. Mycorrhizal Symbiosis. Academic Press. 1997; 605.

[97] Matsubara Y, Tamura H, Harada T. Growth enhancement and Verticillium wilt control vesicular-arbuscular mycorrhiza fungus inoculation in eggplant. Journal of the Japanese Society for Horticultural Science. 1995; **64**(3):

resistance induced by *Laccaria bicolor* to *Fusarium oxysporum* in primary roots of *Donglas* fir. Phytopathology. 1991; **81**:

Interaction between *Glomus fasciculatum*

**22**: 1-17.

420-425.

**61**(9): 2349-2351.

2006; 39-55.

2002; 87-101.

555-561.

mycorrhizal fungus. Frontiers in Plant

[81] Hartley SE, Gange AC. Impacts of plant symbiotic fungi on insect

herbivores: mutualism in a multitrophic context. Annual Review of Entomology.

[82] Kempel A, Schmidt AK, Brandl R,

underground: induced plant resistance depends on arbuscular mycorrhizal fungi. Functional Ecology. 2010; **24**:

[83] Koricheva J, Gange AC, Jones T. Effects of mycorrhizal fungi on insect herbivores: a meta-analysis. Ecology.

[84] Babikova Z, Gilbert L, Bruce TJA, Birkett M, Caulfield JC, Woodcock C, Pickett, JA, Johnson D. Underground signals carried through common mycelial networks warn neighbouring plants of aphid attack. Ecology Letters.

[85] Babikova Z, Gilbert L, Bruce T, Dewhirst SY, Pickett JA, Johnson D. Arbuscular mycorrhizal fungi and aphids interact by changing host plant

Functional Ecology. 2014; **28**: 375-385.

[86] Zcon AC, Barea JM. Interactions between mycorrhizal fungi and other rhizosphere microorganisms. In: Allen MF (ed.), Mycorrhizal Functioning: An integrative plant-fungal process. New York: Chapman and Hall. 1992;

[87] Ingham ER. The soil food web. In: Soil biology primer. Rev. edition. Ankeny, USA, Soil and Water Conservation Society. 2000.

[88] Dehne HW. Interaction between vesicular arbuscular mycorrhizal fungi and plant pathogens. Phytopathology.

1982; **72**(8): 1115-1119.

quality and volatile emission.

Schadler M. Support from the

Science. 2015; **6**: 786.

2009; **54**: 323-342.

2009; **90**: 2088-2097.

2013; **16**: 835-843.

293-300.

**202**

163-198.

[98] Tarraf W, Ruta C, Tagarelli A, De Cillis F, De Mastro G. Influence of arbuscular mycorrhizae on plant growth, essential oil production and phosphorus uptake of *Salvia officinalis L.* Industrial Crops and Products. 2017; **102**: 144-153.

[99] Harrier LA, Watson CA. The potential role of arbuscular mycorrhizal (AM) fungi in the bioprotection of plants against soil-borne pathogens in organic and/or other sustainable farming systems. Pest management science. 2004; **60**: 149-157.

[100] Jawson MD, Franzlubbers AJ, Galusha DK, Aiken RM. Soil fumigation within monoculture and rotations response of corn and mycorrhizae. Agronomy Journal. 1993; **85**: 1174-1180.

[101] Knudsen IMB, Debosz K, Hockenhull J, Jensen DF, Elmholt S. Suppressiveness of organically and conventionally managed soils towards brown foot rot of barley. Applied Soil Ecology. 1995; **12**: 61-72.

[102] Hamel C, Strullu D-G. Arbuscular mycorrhizal fungi in field crop production: potential and new direction. Canadian Journal of Plant Science. 2006; **86**: 941-950.

[103] Balzergue C, Chabaud M, Barker DG, Becard G, Rochange SF. High phosphate reduces host ability to develop arbuscular mycorrhizal symbiosis without affecting root calcium spiking responses to the fungus. Frontier of Plant Science. 2013; **4:** 426. doi: 10.3389/fpls.2013.00426

[104] Solaiman ZM, Hirata H. Effect of arbuscular mycorrhizal fungi inoculation of rice seedlings at the nursery stage upon performance in the paddy field and greenhouse. Plant and Soil. 1997; **191**: 1-12.

[105] Sanchez PA. Properties management of soils in the tropics. New York: Wiley- Interscience. 1994.

[106] Woomer PL, Swift, MJ. The Biological Management of Tropical Soil Fertility. Chichester, UK: Wiley/UK: TSBF and Sayce. 1994.

[107] Swift MJ. Toward the second paradigm: integrated biological management of soil. Paper presented for the FERTBIO Conference, Brazil. 1998.

[108] Hawksworth DL. The fungal dimension of biodiversity: magnitude significance and conservation. Mycological Research. 1991; **95**: 641-655.

[109] Gianinazzi-Pearson V, Diem HG. Endomycorrhizae in the tropics. In: Dommergues YR,Diem HG (Eds.), Microbiology of Tropical Soil and Plant Productivity*.* The Hague: MartinusNijhoff/Dr Junk W. Publishers. 1982.

[110] Davet P. Vie microbienne du sol et production végétale. Paris: INRA. 1996.

[111] Ike-Izundu NE. Interaction between arbuscular mycorrhizal fungi and soil microbial population in the rhizosphere. Doctoral Thesis. 2007.

[112] Mattoo AK, Teasdale JR. Ecological and genetic systems underlying sustainable horticulture. Horticultural Reviews 37. Jules Janick (Ed.), Wiley-Blackwell. 2010

## *Edited by Takuji Ohyama and Kazuyuki Inubushi*

Nitrogen is the most important nutrient in agricultural practice because the availability of nitrogen from the soil is generally not enough to support crop yields. To maintain soil fertility, the application of organic matters and crop rotation have been practiced. Farmers can use convenient chemical nitrogen fertilizers to obtain high crop yields. However, the inappropriate use of nitrogen fertilizers causes environmental problems such as nitrate leaching, contamination in groundwater, and the emission of N2O gas. This book is divided into the following four sections: "Ecology and Environmental Aspects of Nitrogen in Agriculture", "Nitrogen Fertilizers and Nitrogen Management in Agriculture", "N Utilization and Metabolism in Crops", "Plant-Microbe Interactions".

Published in London, UK © 2021 IntechOpen © subjob / iStock

Nitrogen in Agriculture - Physiological, Agricultural and Ecological Aspects

Nitrogen in Agriculture

Physiological, Agricultural

and Ecological Aspects

*Edited by Takuji Ohyama and Kazuyuki Inubushi*