PHENOLICS IN RHIZOSPHERE ACT AS A DEFENSE MECHANISM

Phenolic compounds enter soil via plant root secreting root exudates or as leachates or as plant residue decomposition. Generally phenolic compounds released from seeds, roots or residue decomposition can act against soil borne pathogens and root feeding insect (Ndakidemi and Dakora 2003; Mandal et al., 2010). Phenolics usually accumulate in the central vacuoles of the guard cells and epidermal cells as well as in subepidermal cells of leaves and shoot (Lattanzio et al., 2006). Few phenolics are covalently linked to the plant cell wall, while others occur in waxes or on the external surfaces of the plant organ. Past studies also suggest deposition of flavonoids in nuclei of certain tree species, suggesting that flavonoid-DNA complex provides mutual protection against oxidative damage (Sharma and Sharma 1999; Lattanzio et al., 2006). Phenolic storing cells are distributed within most tissues. It is proposed that these cells by decompartmentation, rapid oxidation of their phenolic content, further lead to lignification and suberization of cells, cell death, seal off infection or injuries at the infection site as a defense response against the pathogen attack (Beckman 2000).

Plant phenolics may be divided into two forms i) Constitutive phenolics function as preformed inhibitors associated with nonhost resistance and ii) Induced phenolics that are synthesized in response to the abiotic and biotic stress (Hammerschmidt et al., 2001; Nicholson and Hammerschmidt 1992; Dixon et al., 2002). Phenolics in soil exists in three forms (Min et al., 2015):

  1. as a dissolved form, which moves freely in soil solution
  2. a sorbed form which is reversibly bounded to the soil particles or proteins and
  3. a polymerized form, consisting of humic substances

Plants produce phenolic allelochemicals including simple phenolics, however phenolic allelochemicals from dead plants and crops are lignin-related phenolics acids (Mushtaq and Fauconnier 2024). Cinnamic acid derivatives, notably t-cinnamic, p-coumaric, ferulic and vanillic acids   possess antifungal activity (Tawata et al., 1996; Kim et al., 2004; Bisogno et al., 2007). Phenolic allelochemicals can increase cell membrane permeability. Consequently, leakage of cell content and increased lipid peroxidation leading to slow growth or death of plant tissue. In addition, phenolic allelochemicals can inhibit plant from absorbing nutrients from the surroundings and affect the normal growth of the plants (Li et al., 2010). Phenol protocatechuic acid and catechol from onion are toxic substance that provides resistance against the fungus Colletotrichum circinans causing onion smudge disease (Link and Walker 1933). Allelochemical phenolics such as ferulic acid and cinnamic acid can inhibit protein synthesis in Latuca sativa L. (Cameron and Julian 1980). Phenolics exert allelopathic effects on various physiological processes in plants such as inhibition of cell division, elongation and submicroscopic structures, various enzymes functions and activities, synthesis of plant endogenous hormones and protein synthesis (John and Sarda 2012). The production, release and addition of several phenolics act as defense mechanisms against herbivores, pest and pathogens (Misra et al., 2023).

Many of the phenolic root exudate serve as a chemotactic signal for several soil microorganisms that recognize them and move towards the plant roots in a carbon-rich rhizosphere (Taylor and Grotewold 2005). Different organisms are repelled from or attracted towards the same chemical, which then elicit different responses in different organisms. Example, isoflavones from soybean roots serve as an attractant for both the symbiotic Bradyrhizobium japonicum and the pathogenic Phytophthora sojae (Morris et al., 1998). The zoospores of soybean pathogen Phytophthora sojae respond chemotactically to the soybean isoflavones daidzein and genistein exuded from the roots of soybean in the rhizosphere, suggesting the hyphal tips from zoospores that have encysted adjacent to root may use specific host isoflavones to locate their host (Morris et al., 1998).

Beneficial bacteria and fungi as well as bacterial pathogen can induced phenolic production (Wallis and Galarneau 2020). Phenolic production is associated with salicylic acid resistance response, and interaction between jasmonic acid and salicylic acid may have a role in regulating and fine tuning the induced defense which is activated in response to the pathogen attack or insect attack or wounding (Yang et al., 2011; Wallis and Galarneau 2020). Phenolic compounds are often increased following attack during hypersensitive response that leads to cell death. Insects need living tissues for successful feeding would inhibit host defense response (Wallis and Galarneau 2020). Solanaceous plant release endogenous phenolic substances in response to pathogen attack. Whereas the exogenous phenolic substances function to prevent and control pest and diseases during the production of solanaceous plants (Wang et al., 2024). Different type of resistance strategy produce cost, especially cost associated with inducible defenses (Purrington 2000). Plants that suffer frequent and serious damage may invest mainly in constitutive defense, while plants that are attacked rarely may rely on induced defense (Wittstock and Gershenzon 2002; Lattanzio et al., 2006).

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