Water-extractable humic substances alter root development and epidermal cell pattern in Arabidopsis

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The Use of Biostimulants for Enhancing Nutrient Uptake (2015, background): “…al., 1993; Zandonadi et al., 2007), municipal waste (Ayuso et al., 1996), vermicomposts and earthworm casts (Canellas et al., 2002; Russell et al., 2006), various coal deposits (Kulikova and Perminova, 2002), peat (Ayuso et al., 1996; Schmidt et al., 2007), and Leonardite (Nikbakht et al., 2008).”
Integrated Use of Humic Acid and Plant Growth Promoting Rhizobacteria to Ensure Higher Potato Productivity in Sustainable Agriculture (2019, background): “In addition, previous studies reported that humic substances directly affected plant growth by inducing an increase in the absorptive surface area of roots, especially lateral root emergence [52].”
Chemical properties of humic matter as related to induction of plant lateral roots (2011, background): “Many studies have confirmed the hypothesis of a direct effect of HS on plant physiology (Asli & Neumann, 2010; Mora et al., 2010), and in particular on the development of lateral roots (Canellas et al., 2002; Schmidt et al., 2007; Zandonadi et al., 2007; Canellas et al., 2008a).”
Rhizosphere humic acid interacts with root cell walls to reduce hydraulic conductivity and plant development (2010, background): “Interestingly, humic acids have been widely reported to stimulate root elongation, root hair and lateral root production (cf. Canellas et al. 2002; Zandonadi et al. 2007; Schmidt et al. 2007; Canellas et al. 2009).”