The four open reading frames (C1, C2, V1, V2) are shown. The complementary (C-sense) genes (encoding Rep and RepA) are required for replication, the virion (V-sense) genes (encoding movement protein, MP and coat protein, CP) for systemic infection. Transcription is bi-directional (transcripts shown in green). Transcription start sites are within the long intergenic region (LIR), termination is within the short intergenic region (SIR).