In this work, the microstructure–property relationship of the heat-affected zone (HAZ) of a FH690 ultra-heavy marine steel plate was investigated based on insight of microstructure and crystallographic features. After multi-pass welding with a heat input of ~30 kJ/cm, an ~8 mm wide HAZ was obtained with a coarse grain HAZ (CGHAZ) of ~3.8 mm, fine grain HAZ (FGHAZ) of ~3.4 mm, and intercritical HAZ (ICHAZ) of ~1 mm. High impact toughness values of ~120 and 140 J at -60 °C were obtained for coarse grain HAZ and fine grain HAZ, respectively. The microstructure of the CGHAZ and FGHAZ was fine lath bainite. Although the average prior austenite grain size for the CGHAZ was ~75 μm, which was five times that of the FGHAZ (15 μm), a high density of high-angle grain boundaries (HAGBs) with misorientation higher than 45° was obtained in the CGHAZ. This is the underlying reason for the excellent low-temperature toughness of the HAZ. Thermo-dynamic calculations indicated that the high density of HAGBs in the CGHAZ was attributed to the decreased bainitic transformation temperature due to the reduced phase transformation driving force via the high nickel addition, leading to weak variant selection. In addition, the high nickel addition offered high hardenability for high hardness in the FGHAZ. The outcome of this study could provide an experimental and fundamental basis for designing high-strength ultra-heavy steel plates with excellent weldability.