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 Advances in Microbiology Research  Category:

Microbiology  Type: Review Article

Regulatory Role of Arbuscular Mycorrhiza

Fungi and Helper Bacteria Associations in
P and N Dynamics in Agriculture
Seema Sangwan1*, Garima Saxena1, Ram Swaroop Bana2
and Radha Prasanna1
1
Division of microbiology, ICAR Indian Agricultural Research
Institute, New Delhi, India
2 Division of agronomy, ICAR Indian Agricultural Research
Institute, New Delhi, India

*
Corresponding Author(s):
Seema Sangwan
Division Of Microbiology, ICAR Indian Agricultural Research
Institute, New Delhi, India
Tel:011-25847649,
Email:[email protected]

Received Date: Sep 12, 2023

Accepted Date: Oct 06, 2023
Published Date: Oct 13, 2023
DOI:10.24966/AMR-694X/100026
Abstract
Interactions among microflora and plants represent a major
pillar in rhizosphere biology for improving soil fertility and crop
productivity. As the most important macronutrients in soil, the
availability of Phosphorous (P) and Nitrogen (N) significantly
affects plant growth and yield, across crops, and around the
globe. In soil, bacteria and fungi constitute the major groups of
microbes, existing, both as free-living and in symbiotic/ loose
associations with other living forms, which find use as
biofertilizers in integrated nutrient management. Among various
types of fungi, Arbuscular Mycorrhiza Fungi (AMF) comprise
symbiotic fungi which form an extensively dense network of
mycelia around the plant roots, and improve the soil structure
and increase the uptake of water, as well as nutrients such as P,
N/ micronutrients by plants. Several bacteria capable of
solubilizing phosphorous mainly via releasing a wide range of
organic acids and chelating metabolites are also present in free
living form and in association with AMF. Such bacteria
associated with AMF hyphae and spores are called as Mycorrhiza
Helper Bacteria (MHB) as they help in regulating the activity and
functioning of AMF. Characterization of such bacteria and
developing promising combinations of AMF and MHB, can be
beneficial for improving the nutrient availability in soil and
stimulating plant growth. This review summarises and discusses
the current knowledge on the interactions among AMF and MHB
towards enhancing the availability of N and P availability, and its
uptake by plants, thereby, highlighting the research gaps that
need attention and in-depth research.

Keywords
Arbuscular Mycorrhiza Fungi; Mycorrhiza Helper Bacteria;
Nitrogen; Phosphorous; Synergism

Introduction
Replenishing the soil biological properties through the tools of
microbial inoculation and addition of requisite quantities of
nutrient inputs is the hall mark for keeping agricultural
revolution evergreen. For getting optimum yields in a
sustainable manner without any dire consequences to
environment, it becomes necessary to provide a regular supply
of these macronutrients, through fertilization (chemicals or
biofertilizers). Phosphorous (P) and Nitrogen (N) are the primary
nutrients required for the growth, development of plants and are
essential because they actively participate in various processes
e.g., carbon metabolism, energy generation, energy transfer,
enzyme activation, membrane fixation and nutrient cycling [1-4].
Phosphorous is involved in formation of ATP, nucleic acids and
phospholipids [5]. P deficiency is known to cause approximately
50% losses in all agricultural lands around the world [6,7]. Like P,
N is also a primary major nutrient required by plants as it forms
a part of the structure and functioning of some important macro
and micro building blocks in plants such as chlorophylls,
proteins and amino acids [5,8]. For improving the availability of P
and N to the plants, chemical fertilizers are often used in an
unbalanced manner, without knowing its long-term
consequences and its ill effects on the environment. To mitigate
this, the use of biological options to make available N and P to
the plants is the sustainable way forward. Arbuscular Mycorrhizal
Fungi (AMF) and their intimate relationship with Mycorrhiza
Helper Bacteria (MHB), are getting more attention among
various types of biofertilizers, as they effectively increase P, N
uptake and crop productivity [9,10].

AMF represent an obligate biotrophic association with plant

roots that establishes mutualistic symbiosis with 80% of the
terrestrial plants such as cereals, pulses, fruit trees, vegetable,
medicinal plants and other commercial crops such as sunflower,
cotton and sugarcane [11]. In exchange for plant
photosynthates, AMF facilitate the uptake and transfer of
mineral nutrients such as P, N, S, Ca, Cu and Zn from the soil to
the host plant through their Extra Radical Mycelium (ERM) [12].
ERM function as an efficient absorbing system that enables the
uptake of nutrients beyond the depletion zone [13-15]. Besides
providing nutrition to the plants, AMF also facilitates the
completion of several biochemical cycles, enhances tolerance to
biotic and abiotic stress [16], sequesters carbon, improves soil
aggregation [17] and plays an important role in synthesis of
health promoting phytochemicals [18]. In the mycorrhizosphere,
MHB which are associated with AMF spores and hyphae,
thereby, playing an important role in nutrient uptake and overall
growth of the plant, as they are potent bioenhancers of plant-
AMF associations [19]. They help in improving hyphal growth,
spore germination and establishment of mycorrhizal symbiosis.
MHB isolated and characterized till date are gram-positive
Actinobacteria and Firmicutes (e.g., Streptomyces, Brevibacillus,
Bacillus, Rhodococcus, Arthrobacter, Paenibacillus) and
Azospirillum, Pseudomonas, Rhizobium, Burkholderia,
Enterobacter, Agrobacterium, Azotobacter, Bradyrhizobium and
Klebsiella which come under the category of gram negative
Proteobacteria. Species of Streptomyces, Pseudomonas and
Corynebacterium have been shown to improve the germination
of F. mosseae, G. versiformae and G. margarita spores [19,20].
Actinobacteria are a group of bacteria frequently associated with
AMF spores able to hydrolyze chitin found in the spore wall
[21,22]. Other MHB such as Klebsiella pneumonia and
Paenibacillus validus have been reported to increase germling
hyphae growth [23] and Oxalobacteria enhance both spore
germination and germling growth along with root colonization.
ERM development is promoted by Penibacillus spp., Azospirillum
spp. and Pseudomonas spp. [24]. Additionally, MHB:

Promote mycorrhizal symbiosis by several direct and

indirect means including- improvement in root
receptiveness of fungi [25].
Modulate plant-fungi recognition and symbiosis
establishment.
Accelerate EMF propagule germination as well as spore
survival and mycelial growth.
Modify soil chemical properties e.g., for a better
connectivity with the soil fungus, protection against
pathogen and promote defence mechanism [26].

MHB are also known to help breakdown molecules to a usable

form as they can utilize both organic and inorganic nutrients in
soil through a process called “mineral weathering” which aids in
the recycling of nutrients in the rhizosphere environment. MHB
help to make available, P from soil [27,28] and phosphorus
solubilizing rhizobia are the most common MHB involved in P
uptake. They release compounds in soil to break down organic
inorganic P for further use by mycorrhizae. They are active even
under P limited conditions, helping mycorrhiza to establish and
grow [29]. Several MHB also have the capability to acquire
nitrogen and fix N in the soil, without plant modification, as
done by legumes to help in N fixation [28]. Researchers reported
the significant contributions of Bacillus spp. to N fixation [30]
(Table 1).

Ecological AMF Crop Reference

function

N, P, K, Fe Glomus mosseae Rice [31]

and Zn

C, N, P, K Glomus spp. Finger millet [32]

peanut, pigeon
pea

N, P Rhizophagusintraradices, Tomato [33]

Glomus versiformae,

Claroideoglomusetunicatum,

Claroidioglomusclaroideum

C, N, P Funneliformismosseae, Apple [34]

Rhizophagusirregularis

N Funneliformismosseae, Chrysanthemum [35]

morifolium
Diversisporaversiformis

Table 1: Ecological role of AMF to facilitate P and N uptake

Plant Response to Deficiency of P and N in Soil: Plants exhibit

several biochemical, physiological and morphological
adaptations to deal with N and P deficiency, which work in a
linked manner. To increase P and N acquisition capacity, plants
show “P and N starvation response” as shown in Figures 1&2.
Recent scientific advances have helped to provide a better
understanding of the signalling pathway of plant response to N
limitation by microarray and sequencing-based transcription
profiling, e.g. genome scale expression in many plants e.g., rice,
corn [36-38]. Arabidopsis adapts itself to limited nitrogen
through NLA gene (nitrogen limitation adaptation) by inducing
the anthocyanin synthesis [39].

Figure 1: Response of plant to P starvation

Figure 2: Response of plant to N starvation

Mechanism of AMF for P Mobilisation and uptake: The

benefits of AMF in various plants are documented and its
significance in P uptake is the most noteworthy. AMF being
obligate biotrophs, they form a symbiotic relationship with the
roots of plant for their survival [11,40]. These fungi penetrate the
root cortical cells, establish arbuscules, and mediate the
exchange of nutrients such as P [41]. AMF expands the extent of
the mycorrhizal hyphae network up to 25 cm around the root in
the mycorrhizosphere and creates a niche for other microbes, to
facilitate better availability of plant nutrients like P and N [11]. It
has been established that 50% of P uptake in the mycorrhizal
plants is supported by the AMF [42] and the P uptake
mechanisms include-

i. P uptake rate (influx) per unit of arbuscular mycorrhizal root

increases due to the greater absorption surface area of
mycorrhizal roots compared to the cylindrical root surface
of plant systems [43].
ii. AMF enhances the root zone absorption by 10-100% and
improves the plant’s ability to reach more soil resources due
to ERM by increasing absorption area of root through ER
hyphae which extend beyond the P depleted zone, leading
to greater bioavailability of Pi [43].
iii. These hyphae with small diameter allow the ERM to access
small cores to achieve greater P intake, improve P
acquisition plant possess the ability have to overcome the
depletion zone and improve root surface area [11].
iv. AMF store P in polyphosphate form so that they can keep
their Pi level low to efficiently transfer Pi from the soil to
extra radical hyphae and then to intra radical hyphae
[28,44].
v. In addition, organic exudates (citrate, malate) which chelate
Ca, Al ions and dissolve aluminum and calcium phosphate
are secreted in soil [45].
vi. AMF also release some chemicals such as glomalin (a
glycoprotein) which are reservoirs of C in soil and aid in
uptake of Fe, and P [46,19].
vii. Improve the functioning of cell-specific Pi transporters
(Figure 3) which transfer Pi to the host plant also known as
plant Phosphate Transporters (PTs) [47,48]. They facilitate
the acquisition of inorganic Phosphate (Pi) and maintain its
homeostasis within the plant.
 Figure 3: Phosphate and Nitrogen transport between the plant
cell and fungus

(Phosphate and nitrogen acquisition by fungus and transport to

the plant cell through Pht1 i.e. phosphate transporters; nitrogen
through AMT1 i.e. nitrogen transporters)

Mechanism of AMF for N Availability: It is also indicated that

AM fungi can influence the uptake of other nutrients e.g.,
nitrogen which is necessary for plant growth and yield [49].
Results from the root organ culture studies suggest that up to
50% of the root N is acquired through AMF route [50]. A “Trade
balance model” was proposed to explain the apparent nitrogen
paradox [51]. This model suggested that fertilization with N will
be only beneficial, if plant is limited by P and will therefore
benefit from providing C to the roots and mycorrhizal fungi.
AMF can acquire N in various ways:-

i. Inorganic nitrogen in the form of ammonia and nitrate

[52,53].
ii. Directly facilitate the uptake of glycine in Glomus mosseae
through a mechanism, as depicted in Figure 3, in which AMF
acquires organic N directly from the soil substrate.
Researchers have provided evidence for the uptake of
organic N by dipeptide transporters in the intraradical
mycelium of Rhizophagus irregularis [54].
iii. In arid regions, where N mobility/nitrate mobility is low,
AMF can be effective under such conditions to absorb
inorganic N [55,56] indicating that even in such stress
conditions, AMF can help to increase the uptake of N.
 iv. Plant ammonium transporter activation in the presence of
AMF has been reported similar to P [57]. Using labelled N, it
has been demonstrated that the hyphae of AMF are able to
utilise inorganic N efficiently and transfer it to 10-30 cm
[58,59,25].
v. AMF utilise the inorganic N reused from organic source
such as like amino acids [60,61].
vi. To mineralise N, it also produces hydrolytic enzymes such as
pectinase, xyloglucanases, cellulases which are able to
decompose SOM, under humid conditions. This also helps
to reduce the leaching of nitrate in the soil [62].

Under low N availability, competition for N between AMF and

plant has been documented resulting in non-mycorrhizal plants
becoming more efficient than mycorrhizal plant [63]. AMF-
mediated P uptake could also have a significant positive effect
on BNF [64]. There are various modes of N uptake in the
rhizosphere, so it is difficult to distinguish the most important
process- biotrophic, nutrient cycling or transfer through root
exudates [61,65]. There is limited evidence from experiments
involving quantum dot technology that the organic N fragment
could have been taken up by the AMF and uptake of N in the
form of amino acid has also been reported to be as high as
compared to non-mycorrhizal plants (Figure 4).

Figure 4: Mechanisms employed by AMF for P and N uptake

Organic N uptake by AMF: AMF development favours the

release of nutrients from soil through mineralisation (specifically
NH4+ ions) [66]. AMF is found to possess a weak exoenzyme
repertoire [67], such as exophosphatase activities. However, they
are very unlikely to mineralise on their own. AMF rely on other
saprophytic /hyper symbiotic microbes for the N mineralisation
[68]. There are some microbial grazers who excrete a large
amount of NH4+ ions in the soil when they digest other
microbes in soil, thereby returning the N to soil-free ammonium
pool. It can be easily utilised by AMF [61,66]. As of now, there is
very scanty information regarding ERM to be able to directly
take organic N molecules such as amino acids, peptides and
nucleosides from the soil solution [53].

Chitin as a Source of Organic N for AMF: Chitin is a polymer

rich in N (>6%) by weight which is present abundantly in the soil
micro and microfauna. It has been reported that the addition of
crab shell chitin during plant cultivation as substrate can
promote AMF species sporulation [69]. The genes responsible
for the breakdown of NAG (N acetyl glucosamine); a subunit of
chitin polymer, are present in the membrane in Rhizophagus
irregularis [70]. The transporter genes have also been identified
in intra-radical hyphae only. After the chitinolytic degradation of
chitin by various fungi and microbes, it gets transported by the
AMF and while the decomposers are eaten up by grazers and
the N returns to the ammonium pool. But there is a chance of
competition between AMF and ammonium oxidising bacteria in
the soil solution [66].

Mechanism of MHB for P uptake: In order to develop

strategies to promote mycorrhization of AMF and improve
uptake of P from soil, there is a need to elucidate the aspects of
metabolic signalling pathway of MHB and illustrate their
functional significance. MHB are known to help break down
molecules to a more usable form [28]. They obtain both organic
and inorganic nutrients from the soil through a direct process
called “mineral weathering” which aids in recycling the nutrients
e.g., Pseudomonas, Burkholderia. Phosphate Solubilizing
Rhizobacteria (PSRB) are the most common MHB which aid in
the phosphate uptake. They release the phosphate degrading
compounds to break down organic and inorganic phosphate,
thus creates a phosphate pool that mycorrhiza can further use.
PSB are also called as PGPR [28,71,72] as the improved
availability of P improves plant growth. PSB solubilise P by
producing phenolic compounds, organic acids, siderophores and
hormones [73]. The acids destroy the P-bound structure, such as
those of dicalcium phosphate, hydroxyl phosphate, tricalcium
phosphate, rock phosphate, and make P available to plant in the
form of dihydrogen phosphate or hydrogen phosphate [28]. As a
result of all these mechanisms, it has been found that PSB can
increase the growth of cereals, legumes, fibre crops, horticultural
and oil seed crops [22]. The details of these mechanisms are
given as:

i. Production of Organic Acids: The dissolution of P bearing

compounds is pH dependent i.e., P uptake occurs at a pH of
6.5-7 so PSB produce organic and inorganic acids by
mineralization [74,75]. Organic acids like malic, aspartic,
tartaric, oxalic, gluconic are produced by bacterial
metabolism, due to oxidative respiration. Organic acids can
solubilize P from mineral surfaces by ligand exchange or
ligand promoted dissociation. PSB indirectly reduce the pH
and increase the P level by enhancing root exudates that
increase the P availability by maintaining electro-neutrality
in the soil [75]. Such organic acids sometimes also compete
with phosphate at the fixation site but the siderophores
(chelating agent) present in root exudates facilitate the
soluble complex formation and improve Pi precipitation
[76]. Root exudates attract the MHB and lead to the
production of more organic acids and reduction in pH.
ii. Production of Inorganic Acids: MHB also include Sulfur
Oxidizing Bacteria (SOB) and Nitrifying Bacteria (NB)
belonging to the genera Thiobacillus and Nitrobacter which
participate in the production of inorganic acids (carbonic
acid, hydrochloric acid, nitric acid, sulphuric acid) reported
to solubilize P by reducing the pH and increasing the P
availability in the soil [77,78]. SOB oxidizes sulphur
compounds in the presence of oxygen to produce sulphuric
acid and NB oxidizes inorganic nitrogen to produce nitric
acid.
iii. Siderophore Production: Plant and microbes both produce
siderophores under low iron conditions. Siderophores are
the organic compounds having iron chelating ability which
makes iron available to both plants and microbes [79].
These siderophores also bind to various metals like Cd, Pb,
Zn, Al, Ca, Mn, and Mo. Several PSB also reported to
produce siderophore [80] and mediate solublising /
dissolution of the insoluble P bearing minerals [77].
iv. IAA and ACC Deaminase Production: PGPB and PSB and
other MHB can improve P uptake by plant, by promoting
 phytostimulation i.e., stimulating root growth through
increased branching as a result of hormonal stimulation, or
root hair development by the production of IAA and ACC
deaminase [81-83]. The IAA and ACC deaminase promote
effective root architecture, branching, increase root
exudation to lower the rhizospheric pH and improve P
solubilization [84,85]. Many PSB were reported to produce
IAA and ACC deaminase, as also IAA producing PSB similarly
solublising Pi by increasing surface area and root exudates
[82,86]. This synergy of IAA production and P solubilization
being intricately linked is beneficial for plant growth [87].
v. P Hydrolyzing Enzymes: Organic P contributes 40-80% of
soil total P. To hydrolyze organic P compounds, plant and
microbes both produce enzymes. PSB mineralize organic P
by secreting phosphatase enzymes [28,77] while microbes
derive phosphatase mainly combine with phosphate and
help in releasing orthophosphates from soil organic P
[28,77,88]. Phytases are enzymes secreted by microbes
which convert phytate into esters, followed by action of
phosphatase to break down to Pi [5,89]. According to Le-
Chateliers principle, lowering the concentration of Pi in soil
solution promotes the production of Pi by indirect
dissolution of K-apatite or calcium phosphate [90].

Mechanism of MHB for N uptake: MHB in the rhizosphere

often have the capability to acquire nitrogen that the plant can
use. Such MHB are able to fix N in the soil and create pools of
available nitrogen [28,91]. However, MHB do not bring about
any plant modification, as legumes do, to help with nitrogen
fixation [61,91]. Nitrogen fixation is done in the surrounding soil
in relation to mycorrhiza. A Bacillus functioning as MHB
contributed to the nitrogen fixation and among other factors
helped the plant grow when inoculated with fungus and
tripartite interaction with AMF [92], host plant and soil microbes
enhancing the nitrogen fixation capacity in Rhizobium. The P
uptake pathway positively affects the rhizobium N fixation by
influencing the energy-producing pathway. Inoculation of
diazotrophic bacteria such as Gluconaacetobacter, Diazotrphicus,
Burkholdera tropica, Azospirillum amazonese, Herbaspirillum
rubisubalbicans, and Herbaspirillum seropeduca has been show
to distinctly influence and promote N fixation in sugarcane [93].
Except for sugarcane, the use of non-rhizobial N fixing MHB in
other non-leguminous crops has met with limited success [94]
(Table 2).

Crop MHB Nutrient Mechanism Reference

uptake
involved

Finger Bacillus spp. P P [95]

millet, solubilisation
Maize,
Amaranth,
Buckwheat,
French
bean

Ficus Bacillus coagulans, P Positively [96]

benjamina affecting
Trichoderma harizanum
mycorrhizal
fungi

Wheat Azotobacter chroococcum P P [97]

solubilisation
and
hormone
production

Wheat Bacillus P P [98]

circulans,Cladosporiumherbarrum solubilisation
and interact
with AMF

Tomato Enterobacter N, P P [99]

solubilisation
agglomenans
also
positively
affects
mycorrhizal
fungi
 Maize and Pseudomonas alcaligenes, N, P, K Hormonal [100]
sunflower effect on
Bacillus polymyxa
root growth

Chickpea Rhizobia N N fixation [10]

Table 2: MHB functions in P and N uptake

Improving P And N Availability Through

Synergism Between AMF And MHB
“MHB are known to influence AMF fitness” i.e., the synergistic
relationship between MHB and AMF can be described as one in
which MHB leads to increase in plant growth parameters by
stimulating the native AMF association, spore germination,
hyphal growth, rate of colonisation and effectively access soluble
P and N sources [101-103]. The presence of MHB on the surface
of the spores belonging to Rhizophagus irregularis, is illustrated
using Scanning Electron Microscopy (SEM), wherein they are
visible as members of the sheet forming biofilms, and shown to
enhance growth in such associations (Figure 5). Glomus mosseae
spores were reported to be stimulated in terms of mycelial
growth by PGP rhizobacterium [104]. Such bacteria also help to
enhance root colonization of indigenous and inoculated AMF
[92,105]. Bacterial IAA is known to loosen plant cell walls to
release root exudates for providing additional nutrition to
support microbial growth. These exudates contain several
enzymes such as amylase, urease, phosphatases and plant
hormones like IAA, ABA, and gas [106], which help in
mineralisation of P and N for AMF [107,108] and flavonoids to
facilitate root colonization [109]. MHB sometimes release
exudates and create a biofilm; this biofilm represents a means of
long-distance migration and acquisition of nutrients from distant
locations. In onion crop, MHB strains Enterobacter spp. and
Bacillus subtilis were seen to enhance the uptake P from rock
phosphate when inoculated alone with Rhizophagus irregularis
[105]. A mixed biofilm was developed constituting the
Bradyrhizobium eklanii along with phosphate solubilizing fungi
that could solubilise the rock phosphate. AMF also establish
extensive network creating a dedicated niche for bacteria and its
hyphae are the C-rich source to MHBs [21,72]. PSB can grow
alongside AMF hyphae in and out of root in sterile conditions as
well as with an indigenous microbial community [110,111]. AMF
release myc factors similar to nod factors of Rhizobium that
activate nodulation factors inducible gene i.e., MtEnod 11 [61].
This gene enhances lateral root formation and mycorrhization
leading to increased nodulation and symbiotic nitrogen fixation
in mycorrhizal legumes [112]. It also lowers down the root zone
pH by uptake of ammonium ions and release hydrogen ions into
the soil solution which increase the solubility of P and also
increases the nitrogen flow to the plant. PSB mobilise
orthophosphate and also decomposes organic matter in turn to
improve N and P availability for AMF and plants [113]. Co-
inoculation of AMF-MHB by different formulations is efficient for
P and N uptake and needs to be advocated across crop and
ecologies.

Figure 5: SEM images demonstrating the association of MHB

with Rhizophagus irregularis. (a) Bacterial cells present in form of
sheet like biofilms (yellow arrow) on AMF spore surface (scale
bar: 50 µm); (b) and (c) presence of bacterial isolates (white
arrow) on AMF spore surface, (scale bar: 20 µm and 2 µm)

Improvement in Product Quality and Yield by AMF and

MHB: The quality of plant products and their derived foods and
ingredients represent the main area of focus in sustainable
agriculture, which is also in the best interests of farmers,
consumers, and producers. Recently, the concept of food quality,
traditionally based on nutritional and sensory properties, has
acquired an additional meaning, referring to the health-
promoting properties of plant foods, thereby representing an
important societal issue, highly demanded by consumers. The
potential of AMF in enhancing the crop yield and grain quality
have been very widely studied since decades. Their inoculation
causes improvement in plant nutrition, photosynthesis, and
stress resistance which results in 25-30% increase in overall
shoot and root biomass [114]. The potential effects of AMF
alone and in synergism with beneficial microbes have been
attested in some crops yield including wheat [115], barley [116],
soybean [117], and chickpea [118-120]. AMF and associated
bacteria enhance plant growth and health, and affect the
production of polyphenols and carotenoids, and the activity of
antioxidant enzymes. The diversity and content of
phytochemicals in plant products are affected by different
variables, such as plant genotype, agronomic factors, and
Arbuscular Mycorrhizal Fungi (AMF), which establish mycorrhizal
symbioses with most crops, including cereals, legumes,
vegetables, fruit trees, sunflower, cotton, and sugarcane. The
production of health-promoting phytochemicals was shown to
be differentially modulated by different AMF isolates and
bacterial strains, in several food plants, i.e., tomato, lettuce,
strawberry, artichoke, maize, grapevine, sunflower. Here, we
provide an overview of recent studies concerning the multiple
roles played by AMF and associated bacteria in the modulation
of the biosynthesis of plant secondary metabolites with health-
promoting activity, and discuss the development of designed
multifunctional consortia to be used in sustainable agriculture
[121]. However, the beneficial effect of AMF whether alone or in
synergism with MHB is always regulated by the functional group
of crops, across agroecologies.

Future Perspectives
The multiple beneficial activities of AMF and their associated
bacteria with complex networking in the mycorrhizosphere are
functionally important for plant growth, nutrition and overall
health. Very little is known about their phylogenetic interactions
and intraspecific diversity of AMF and MHB. Recent studies
confirmed the occurrence of diverse beneficial taxa in a
commercial AMF inoculum in which 14 isolates showed the best
combination of PGP traits, while 6 of them were able to
solubilise P i.e., Bacillus spp., Enterobacter and Streptomyces [21].
This highlights the need for more such studies using different
AMF species and associated bacteria, both singly and in various
combinations to identify the best performing inoculants. The
areas requiring focussed attention include:
 In-depth studies on elucidating the mechanisms underlying
the resilience to diverse environmental conditions such as
drought, salt and metal stress, pesticides [122] and etc.
Transcriptomic or genomic studies to reveal the expression
level of P transporter genes and ammonium transporter
genes in fungal hyphae. Molecular level studies are needed
to understand the transporter mechanisms using tools such
as Real Time PCR (RT-PCR), Phospholipid derived Fatty Acid
(PLFA) / Neutral derived Fatty Acid (NLFA) ratio.
Commercial production of newly designed multifunctional
microbial consortia for use as a bioenhancer for a
sustainable production system
Rapid and automated methods for identification and
screening criteria for quick and efficient selection of
performing bacteria and fungi.
Deciphering the signaling mechanism(s) involved in the
interaction of AMF and the metabolites that benefit the
microorganisms. Although previous research has suggested
the significance of physical interaction of AMF leading to
competition for nutrients [123,124], and AMF hyphal
exudation directly or indirectly manipulating the
communities.
Targeted research to identify the combinations and PGP
characters that are critically affecting fungal secretion and
growth along with the synergistic interaction between AMF
and PSB. It is not clear whether PSB is able to attach to
extraradical AMF hyphal or not. Nitrogen Limitation
Adaptation (NLA) genes and their function at molecular
level in nitrogen deficiency conditions and the
mineralization of organic N and their uptake mechanism
need to be elucidated, which underpin their role as the
future hope of organic farming.
Efforts to characterize unculturable bacteria in the
mycorrhizosphere and develop synthetic microbial-AMF as
an option to enhance the AMF-PSB combinations efficacy.

It is difficult to distinguish the transfer of the N, as it is

biotrophic and mediated through root exudates and nutrient
cycling. This can be traced by monoxenic cultures or tissue
culture-based experiments. By using isotopically labelled
compounds, uptake of simple amino acids has been tested.
There has been no specific information about the identity of any
primary organic N decomposers associated with AMF. Also, an
interesting question that needs to be explained in the future is -
can AMF utilize N from exogenous chitin, without the
contribution of any helper bacteria and if there is any
significance of N uptake depending upon rate of N release. In
addition to this, future experimentation should focus on
complex N/C-rich compounds such as plant biomass or litter for
deeper understanding of mechanism(s) underlying the utilisation
of organic sources by AMF and the associated plant.

Conclusions
A number of macromolecules that support life on this earth
contain Phosphorous (P) and Nitrogen (N) as integral
components e.g., in polysaccharides, proteins, nucleic acids and
many secondary metabolites. It is very important to think about
the roles played by the omnipresent, yet still broadly under-
appreciated Arbuscular Mycorrhizal Fungi (AMF) and associated
helper bacteria which need to be highlighted such that the tri-
partite association between plant and microbes can be utilized
for accelerating agricultural sustainability and promoting human
welfare. To improve environmentally sustainable agricultural
practices, basic findings about the efficiency of organic P and N
recycling in soils using microbes can be translated in the form of
applied research by improving our understanding the
mechanism underlying the interactions of AMF with its
microbiome. In order to ensure enough food on this planet for
every human being, there is an urgent need to improve the
efficient and sustainable production of agriculture, without any
decrease in productivity, which is a challenge itself. To ensure
sustained food availability on this planet for every human being
now and in the future, it is vitally important to take up this
challenge and come up with novel options, including
broadening the scope of use of AMF and its multifunctional
attributes which provide immense benefits to agriculture.

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Citation: Sangwan S, Saxena G, Bana RS, Prasanna R (2023)

Regulatory Role of Arbuscular Mycorrhiza Fungi and Helper
Bacteria Associations in P and N Dynamics in Agriculture. J Adv
Microbiol Res 6: 026

Copyright: © 2023 Seema Sangwan, et al. This is an open-

access article distributed under the terms of the Creative
Commons Attribution License, which permits unrestricted use,
distribution, and reproduction in any medium, provided the
original author and source are credited.
Journal Highlights
 Microbiology

 Micobes

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 Environmental Microbiology

 Food Microbiology
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