Alleviating excess boron stress in tomato calli by applying benzoic acid to various biochemical strategies

Benzoic acid (BA) represents vital roles in plant activity and response to diverse unfavorable conditions. However, its participation in mitigating excess boron (EB) stress in plants is elusive. Herein, we have examined the impacts of BA (1 μM) in controlling boron (B) uptake in tomato (Solanum lycopersicum L.) calli exposed to various EB levels (0, 1, 2, and 3 mM). The free, semi-bound, and bound B forms were stimulated by EB, while these forms were reduced in B-stressed calli by BA supplementation (40.37%, 36.08%, and 66.91%, respectively, less than 3 mM B-stressed calli alone). EB caused a reduction in the uptake of potassium (K⁺), calcium (Ca²⁺), magnesium (Mg²⁺), and nitrite (NO₂^-) while increasing the concentration of phosphorus (P), nitrate (NO₃^-), sulfur (S), and sulfate (SO₄^2-) in B-stressed calli. BA application induced the uptake of K⁺, Ca²⁺, Mg²⁺, NO₃^-, S, and SO₄^2-; however, it reduced P and NO₂^- concentrations in B-stressed calli. EB reduced nitrate reductase activity (NR), while BA application did not alleviate this reduction. EB treatments significantly, in most cases, increased sulfite oxidase (SO) activity. Supplementation of BA along with EB further enhanced SO activity. Cell wall components (cellulose, hemicellulose, and pectin) were decreased under EB treatments but considerably increased in B-stressed calli by BA application. Fourier Transform Infrared Spectrometer (FT-IR) output showed that EB treatments with/without BA led to alterations in cell wall functional groups of calli. Our findings indicated that BA application enabled tomato callus to counteract the harmful effect of EB, leading to improved callus growth.