The hydra and hormetic effects in a single discrete-time overcompensation model

The hydra effect and hormesis are commonly observed in pest management and cancer treatment. However, the interaction among intervention timing, intervention intensity, and density-dependent compensation remains unclear. In this paper, a discrete-time single-population model based on density-dependent overcompensation is developed to investigate hydra and hormetic effects induced by compensation strength and intervention intensity. Interventions are considered at three distinct timings of a generation: early-generation (before density-dependent growth), within-generation (during growth), and late-generation (after growth has occurred). The results show that early-generation interventions shrink the interval of parameter values for which both compensation strength and intervention intensity induce these paradoxical responses. For late-generation interventions, hydra and hormetic responses are driven by overcompensation strength rather than intervention intensity. Accordingly, late-generation timing modulates only the parameter region where overcompensation strength induces paradoxical responses. Interventions applied within-generation induce a dose-response to overcompensation that is monotonic (S-shaped) at low extrinsic mortality but becomes biphasic (inverted U-shaped) at higher mortality. These results quantify the coupling among density regulation, external intervention, and intrinsic growth. Fits to published hormesis datasets show that the model can reproduce recorded biphasic responses. The analysis demonstrates that intervention timing and intensity govern the emergence of parameter regions where multistability, hydra effects, and hormesis coexist by reshaping nonlinear density-dependent feedback. The proposed discrete-time overcompensation model provides criteria for identifying intervention thresholds and improving control strategies in ecological management and precision medicine.