11 Endophytes as Plant Nutrient Uptake-Promoter in Plants
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endophyte Beauveria bassiana (Bals.-Criv.) Vuill. On the other hand, Lledó et al.
(2015, 2016) observed genus Penicillium and Chaetosphaeronema increased the
total root biomass in Poa pratensis, and Chaetosphaeronema, Sordaria fimicola, and
Epicoccum nigrum exhibited a similar effect in Trifolium subterraneum.
Studies performed by Bartholdy et al. (2001) and Johnson et al. (2013) stated the
enhacement of Fe uptake in plants growth and development. One of the mechanisms
of such an effect might be due to the production of siderophores, which strongly bind
Fe
3+ . This is supported by the fact that several endophytic fungi, such as species of
Epichloë/Neotyphodium, contain a non-ribosomal peptide synthetase gene (sidN)
encoding a siderophore synthetase (Johnson et al. 2013). Earlier, Altomare et al.
(1999) reported the secretion of siderophores by endophyte Trichoderma harzianum
Rifai, increasing Fe uptake by their host plants.
In general, nutrient uptake in plants through roots is mediated, among others,
by mass flow, and/or root interception (Jungk 2002). Mass flow takes place when
nutrients are transported to root by the movement of water in the soil, hence any modification in the above or below ground biomass in a plant could affect the nutrient
acquisition (White et al. 1997). In plants with a more proliferation of roots, mass
flow tends to increase corresponding to increased nutrients uptake due to the presence
of endophytic fungi. This fact was reported by Soto-Barajas et al. (2016) in a study
conducted with Epichloë endophytes, wherein N, Ca, Mg, S, Mn, and Mo increased in
endophyte-infected plant, Lolium perenne. Root interception occurs when nutrients
play physical contact with the root surface and thus when a plant-endophyte interaction stimulates larger root biomass, root interception might be positively altered. This
interaction can also modify rhizosphere conditions, affecting the presence, survival,
or development of different rhizospheric organisms, which in turn may facilitate
nutrient uptake (Antunes et al. 2008; Liu et al. 2011; Omacini et al. 2006).
Nutrient transfer between many plant-fungal symbioses is a common strategy,
where fungal symbionts facilitate the uptake of soil nutrients by plant hosts, and the
plant may supply plant-derived carbohydrates to the developing fungus (Kiers et al.
2011). Under this nutrient-transfer perspective, root-associated endophytes were able
to mobilize different nutrients via fungal hyphae, thus helping plants in their nutrient
uptake (Behie and Bidochka 2014). Usuki and Narisawa (2007) found that the endophyte Heteroconium chaetospira (Grove) Ellis, transfered N to Chinese cabbage
plants, and Newsham (2011) stated that Phialocephala fortinii transfered both, N and
P from soil to plant roots via fungal hyphae, thus an increase in minerals concentration
occurred in plant roots and shoots. Similarly, studies conducted with the ascomycete
root-inhabiting endophyte, Colletotrichum tofieldiae (Pat.), demonstrated their role
to facilitate the transfer of P to non-mycorrhizal plant hosts via their hyphae (Almario
et al. 2017; Hiruma et al. 2016). This fact is especially interesting for plants growing
in nutrient-limited conditions. Studies conducted with the fungal endophyte Piriformospora indica Sav. showed its capacity to induce a normal growth of maize
plants cultivated under nutrient-stress conditions, by facilitating the transfer of P
from soil to plant roots (Yadav et al. 2010). Chen et al. (2013) observed Phomopsis
liquidambari causing growth promotion, nitrification, and NH
4+ –N release. Further,
the fungal endophyte P. indica has been reported stimulating the expression of the
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