8Abbreviations: ABA, abscisic acid; APX, ascorbate peroxidase; AsA, ascorbate; Car, 9 carotenoid; CAT, catalase; Chl, chlorophyll; EA, enzymatic antioxidants; EL, 10 electrolyte leakage; Fv/Fm, maximal efficiency of PSII photochemistry; GR, 11 glutathione reductase; MDA, malondialdehyde; NEA, non-enzymatic antioxidants; 12 PMP, plasma membrane permeability; POD, peroxidase; PS, photosynthesis; qP, 13 photochemical quenching coefficient; ROS, reactive oxygen species; SD, standard 14 deviation; SOD, superoxide dismutase.
15One senence summary: The overproduction of ROS was the primary mechanism of 16 plants in response to water stress and that plants tend to acclimate to water stress over Abstract 29 Plants are key to the functionality of many ecosystem processes. The duration and 30 intensity of water stress are anticipated to increase in the future; however, an 31 elucidation of the responses of plants to water stress remains incomplete. For this 32 study, we present a global meta-analysis derived from 1301 paired observations from 33 84 studies to evaluate the response patterns and mechanisms of plants to water stress. 34 The results revealed that while water stress inhibited plant growth and photosynthesis, 35 reactive oxygen species (ROS), plasma membrane permeability, enzymatic 36 antioxidants, and non-enzymatic antioxidants increased. These responses generally 37 increased with the intensity of water stress but were mitigated with experimental 38 duration. Our findings suggested that the overproduction of ROS was the primary 39 mechanism of plants in response to water stress and that plants tend to acclimate to 40 water stress over time to some extent. Our synthesis provides a framework for 41 understanding the responses and mechanisms of plants under drought conditions. 42 KEYWORDS 43 Water stress, plants, meta-analysis, reactive oxygen species, drought adaption 44 48 required for photosynthesis due to stomatal closure and reduced internal water 49 transport (Breda et al., 2006). As such, water stress impairs normal plant functionality 50 and further induces morphological, physiological, and biochemical changes to 51 compensate for water limitations (Mitchell et al., 2013; Lee et al., 2016). 52 Understanding the patterns and mechanisms of responses by plants to water stress is 53 central to predicting future plant functionality and resilience to drought episodes. 54 The impacts of water stress on plant growth, physiology, and biochemistry are 55 well documented, and numerous individual studies have examined the roles of plant 56 physiological indexes as relates to their tolerance to water stress (van der Molen et al., 57 2011; Zwicke et al., 2015). For plants, water limitations lead to the overproduction of 58 reactive oxygen species (ROS), such as hydrogen peroxide (H 2 O 2 ), and superoxide 59 anion radical (O 2̄· ), which results in growth inhibition (Wallace et al., 2016), 60 decreases in photosynthetic functions (Deeba et al., 2012), lipid peroxidation, and 61 further programmed cell death pro...