Perceiving incoming environmental information is critical for optimizing plant growth and development. Multiple B-box proteins (BBXs) play essential roles in light-dependent developmental processes in plants. However, whether BBXs function as a signal integrator between light and temperature in tomato plants remains elusive. In this study, 31 genes were identified from the newly released tomato () genome sequences and were clustered into five subgroups. Gene structure and protein motif analyses showed relatively high conservation of closely clustered genes within each subgroup; however, genome mapping analysis indicated the uneven distribution of the genes on tomato chromosomes. Promoter -regulatory elements prediction and gene expression indicated that genes were highly responsive to light, hormones, and stress conditions. Reverse genetic approaches revealed that disruption of , and largely suppressed the cold tolerance of tomato plants. Furthermore, the impairment of , and suppressed the photosynthetic response immediately after cold stress. Due to the impairment of non-photochemical quenching (NPQ), the excess photon energy and electron flow excited by low temperature were not consumed in , and silenced plants, leading to the over reduction of electron carriers and damage of the photosystem. Our study emphasized the positive roles of light signaling transcription factors SlBBXs in cold tolerance in tomato plants, which may improve the current understanding of how plants integrate light and temperature signals to adapt to adverse environments.