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Plants increase their freezing resistance upon exposure to low temperatureThe freezing resistance (LT50 values) was found to vary ranging from − 6.9 °C (14-August-2017) to − 31.7 °C (04-November-2018) over the course of study period. The freezing resistance of leaves recorded during the 12 sampling time-points has been provided in Table 1 (also see39). The overlap of confidence intervals around the mean was examined for comparison of LT50 values for the different sampling time-points. Significant differences in freezing resistance were observed across the sampling time-points (Table 1). Leaves of R. anthopogon collected during summer [July and August (Air temperature and photoperiod was about 9.6 °C and 13 h day−1 respectively)] showed marginal resistance to freezing (LT50: − 7 °C) and thus, are more susceptible to freezing damage. Further, as the ambient air temperature and photoperiod decreased towards the end of growing season (i.e., October and November 2017 with air temperature and photoperiod of about − 1.1 °C and 10.5 h day−1 respectively), the plants acquired the highest freezing resistance (LT50: − 30 °C). Interestingly, a sharp increase in freezing resistance (− 29.4 °C) was observed in September 2018, when the daily mean air temperature decreased below 0 °C due to sudden snowfall (Supplementary Fig. S2). Comparison of LT50 values of all the leaf samples of R. anthopogon showed that cold de-acclimation occurred after the snowmelt during early spring in June (LT50: − 13.4 °C) with an increase in air temperature and photoperiod. These results demonstrated that R. anthopogon plants exhibit lowered freezing resistance during the warmer months [hence, these time-periods were referred as non-acclimation (NA)], progressively develop greater freezing resistance during the onset of winter season (hence, referred as cold acclimation) followed by an intermediate level of freezing resistance during the spring [hence, these time-periods were referred as de-acclimation (DA)].Table 1 The estimates of LT50, calculated by fitting sigmoidal curve to electrolyte leakage values of temperature treatments, recorded for leaves collected during the different sampling time-points (from August 22, 2017 to September 18, 2018).Full size tableDuring the acclimation period (i.e., late in the growing season), plants acquired the highest resistance to freezing (Fig. 1). The low electrolyte leakage (= high freezing resistance) observed during this period might be due to changes in cell wall properties (such as increase in lignification and suberization of cell walls), which provide resistance to diffusion of electrolytes from cells of the leaves to the extracellular water47. Moreover, high freezing resistance may also be attributed to high leaf toughness and sclerophyllous habit of this evergreen species48. Further, it was found that freezing resistance was the lowest during mid-summer period. This pattern could be explained by a trade-of between plant growth rates and freezing resistance, where warmer temperatures favour plant allocation to growth49. These observations corroborated well with earlier reports that showed a rapid increase in ‘freezing resistance’ during the transition from summer to early winter and vice versa50.Figure 1LT50 [black point (with solid fill) on the curve] calculated by fitting sigmoidal curve to relative electrolyte leakage (REL %) values recorded during the three different acclimation phases. GOF indicates ‘goodness of fit’ test values for the fitted sigmoidal curves.Full size imagePhotosynthetic rates are higher during non-acclimation and de-acclimation periodIt was found that PN of R. anthopogon varied in the range from 8.336 to 17.64 μmol(CO2)m−2 s−1 and E from 2.281 to 4.912 mol(H2O)m−2 s−1, throughout its growing season. The Gs of leaves was estimated to be in the range from 0.110 to 0.265 mol (H2O) m−2 s−1. WUE, a ratio of PN and E, varied between 52.21 and 87.68 (Table 2). The gas exchange parameters of R. anthopogon varied significantly among the sampling time-points [referred to here as different acclimation phases of the growing period of evergreen shrub (Fig. 2, Table 3)]. In particular, PN was significantly lower on 18-September-2018 (referred as cold acclimation phase), whereas it was higher on 31-August-2018 and 15-June-2018 (referred as NA and DA phases, respectively). Similarly, Gs of leaves was significantly lower during cold acclimation in comparison to the rest of the acclimation phases (i.e., NA and DA). Further, WUE was significantly higher during cold acclimation, while it was lower during both NA and DA (p ≤ 0.05) (Fig. 2).Table 2 Variability in leaf gas exchange parameters of R. anthopogon during the different acclimation phases (NA = Non-acclimation, LA = Late cold acclimation and DA = De-acclimation).Full size tableFigure 2Variability in leaf gas exchange parameters of R. anthopogon during the three acclimation phases [i.e., Non-acclimation (31 August, 2018), Cold acclimation (18 September, 2018) and De-acclimation (15 June, 2018)]. Different alphabets (a, b, c) represent statistically significant values (p  More

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