Fig. 1.

Environmental inputs at different time points (Ancestral, Parental, Developmental and Current) have variable timescales of effect (Evolutionary time, Lifetime and Days-minutes-seconds). The ancestral environment (e.g., the ecological context and selection pressures faced by individuals, including resource abundance, competition and predation threat) is transmitted across generations genetically as sequence level variation, and thus is considered as part of a spectrum of environmental circumstances (i.e., abiotic and biotic factors) that impact behavioural expression⁶⁵. In the hypothetical example above, a behavioural phenotype is influenced by (left to right): (1) inherited gene sequence variation that evolved over time (purple vs. blue and red), (2) parental influences (which may include chromatin-based epigenetic effects, or other features under parental control, e.g., egg composition or oviposition site), (3) the environment experienced throughout development (including impacts on tissue structure or other mechanisms), and (4) the current environment experienced in real time. Solid black arrows indicate shifts among levels of a phenotype. We propose that timescales for behavioural effects are likely non-independent due to shared underlying regulatory mechanisms