Dissecting the Opposing Roles of Thermal Intensity and Growing Degree Days in Regulating Spring Wheat Protein Content

Protein content (PC) stability is crucial for wheat quality. This study utilized partial least squares regression and structural equation modeling to distinguish the physiological effects of "thermal intensity" versus "thermal accumulation" on spring wheat PC across Inner Mongolia. Environmental factors were the dominant drivers of variation. Notably, the Erguna region achieved the highest PC (18.53%) despite recording the lowest total growing degree days. Structural equation modeling analysis revealed that thermal intensity during heading-to-anthesis exerted a strong positive effect on PC (path coefficient = 0.965), likely by enhancing nitrogen remobilization kinetics. Conversely, excessive thermal accumulation and sunshine duration during grain filling negatively impacted PC via a carbohydrate-driven "dilution effect". These findings suggest that superior PC formation requires a specific spatiotemporal coupling: high thermal intensity prior to anthesis to prime nitrogen transport, combined with low thermal accumulation post-anthesis to restrict carbon dilution. This study provides a physiological basis for optimizing wheat quality zoning by decoupling heat magnitude from duration under future climate scenarios.