Halophytes display distinctive physiological mechanisms that enable their survival and growth under extreme saline conditions. This makes them potential candidates for their use in saline agriculture. In this research, tomato (Solanum lycopersium Mill.) was cultivated in moderately saline conditions under two different managements involving Arthrocaulon macrostachyum L., a salt accumulator shrub: intercropping, i.e., co-cultivation of tomato/halophyte; and crop rotation, in which tomato is grown where the halophyte was previously cultivated. The effect of these crop managements was evaluated in tomato plants in comparison with tomato in monoculture, with regards to physiological and biochemical variables and metabolomic and proteomic profiles. Both halophyte-based managements reduced soil salinity. Crop rotation enhanced photosynthesis and protective mechanisms at the photosynthetic level. In addition, both crop managements altered the hormone profile and the antioxidant capacity, whereas a reactive oxygen species over-accumulation in leaf tissues indicated the establishment of a controlled mild oxidative stress. However, tomato production remained unchanged. Metabolomic and proteomic approaches suggest complex interactions at the leaf level, driven by the influence of the halophyte. In this regard, an interplay of ROS/lipid-based signalling pathways is proposed. Moreover, improved photosynthesis under crop rotation was associated with accumulation of sugar metabolism-related compounds and photosynthesis-related proteins. Likewise, acylamino acid-releasing enzymes, a class of serine-proteases, remarkably increased under both halophyte-based managements, which may act to modulate the antioxidant capacity of tomato plants. In summary, this work reveals common and distinctive patterns in tomato under intercropping and crop rotation conditions with the halophyte, supporting the use of A. macrostachyum in farming systems.

Department of Molecular Biology and Radiobiology Faculty of AgriSciences Mendel University Brno Czech Republic

Department of Plant Breeding CEBAS CSIC Group of Fruit Biotechnology Murcia Spain

Research School of Biology The Australian National University Acton Australia

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