Stress-induced dynamic adjustments of reproduction differentially affect fitness components of a semi-arid plant

1. Summer drought stress is considered the primary constraint to plant performance in Mediterranean ecosystems. However, little is known about the implications of summer stress for plant reproduction under real field conditions and, particularly, for the regulatory mechanisms of maternal investment in reproduction. 2. The relationship between plant physiological status at different reproductive stages over the course of the summer drought period and final reproductive output was modelled in the Mediterranean semi-arid specialist Helianthemum squamatum. 3. Plant physiological status, assessed by the chlorophyll fluorescence-based parameter Fv/Fm, and soil moisture content beneath each plant, were determined in the field at five key phenological moments in a total of 88 plants. We used Generalized Linear Mixed Models to evaluate the effect of plant physiological status at those different dates on several components of reproduction (number of flowers and seeds per plant, fruit-set and intra-fruit seed abortion). We included soil moisture as an additional predictor to statistically control its potential effect on reproduction. 4. Fv/Fm measured at midday was a significant predictor of reproductive output, but its significance varied over time and with the specific reproductive response variable. Fv/Fm measured at the onset of flowering was positively related to the number of flowers and seeds per plant, whereas Fv/Fm at the fruiting peak positively affected fruit-set. Soil moisture content was only significant when measured before flowering, being positively related to total flowers and seeds. The effect of stress on reproductive output acted either at an early stage of the reproductive season, by varying the number of flowers produced and seed primordia initiated, or at a later stage, by adjusting the number or ripe fruits. 5. Synthesis. Our results show a direct relationship between physiological status and reproduction, and highlight the importance of the timing of stress for reproductive success. They also show that small departures from the physiological optimum at specific reproductive stages may cause significant decreases in the reproductive output. We suggest that the dynamic adjustment of reproduction in response to stress is adaptive in fluctuating and unpredictable Mediterranean semi-arid environments, where an adequate temporal distribution of maternal resources determines the species' ability to withstand severe environmental conditions.