The causes and impacts for heat stress in spring maize during grain filling in the North China Plain - A review
High-temperature stress (HTS) at the grain-filling stage in spring maize (Zea mays L.) is the main obstacle to increasing productivity in the North China Plain (NCP). To solve this problem, the physiological mechanisms of HTS, and its causes and impacts, must be understood. The HTS threshold of the duration and rate in grain filling, photosynthetic characteristics (e.g., the thermal stability of thylakoid membrane, chlorophyll and electron transfer, photosynthetic carbon assimilation), water status (e.g., leaf water potential, turgor and leaf relative water content) and signal transduction in maize are reviewed. The HTS threshold for spring maize is highly desirable to be appraised to prevent damages by unfavorable temperatures during grain filling in this region. HTS has negative impacts on maize photosynthesis by damaging the stability of the thylakoid membrane structure and degrading chlorophyll, which reduces light energy absorption, transfer and photosynthetic carbon assimilation. In addition, photosynthesis can be deleteriously affected due to inhibited root growth under HTS in which plants decrease their water-absorbing capacity, leaf water potential, turgor, leaf relative water content, and stomatal conductance. Inhibited photosynthesis decrease the supply of photosynthates to the grain, leading to falling of kernel weight and even grain yield. However, maize does not respond passively to HTS. The plant transduces the abscisic acid (ABA) signal to express heat shock proteins (HSPs), which are molecular chaperones that participate in protein refolding and degradation caused by HTS. HSPs stabilize target protein configurations and indirectly improve thylakoid membrane structure stability, light energy absorption and passing, electron transport, and fixed carbon assimilation, leading to improved photosynthesis. ABA also induces stomatal closure to maintain a good water status for photosynthesis. Based on understanding of such mechanisms, strategies for alleviating HTS at the grain-filling stage in spring maize are summarized. Eight strategies have the potential to improve the ability of spring maize to avoid or tolerate HTS in this study, e.g., adjusting sowing date to avoid HTS, breeding heat-tolerance varieties, and tillage methods, optimizing irrigation, heat acclimation, regulating chemicals, nutritional management, and planting geometric design to tolerate HTS. Based on the single technology breakthrough, a comprehensive integrated technical system is needed to improve heat tolerance and increase the spring maize yield in the NCP.