Summary
Only a limited number of dominant resistance genes acting against plant viruses have been cloned, and further functional studies of these have been almost entirely limited to the resistance genes
Rx
against
Potato virus X
(PVX) and
N
against
Tobacco mosaic virus
(TMV). Recently, the cell‐to‐cell movement protein (NS
M
) of
Tomato spotted wilt virus
(TSWV) has been identified as the avirulence determinant (Avr) of Sw‐5b‐mediated resistance, a dominant resistance gene which belongs to the class of SD‐CC‐NB‐LRR (Solanaceae domain‐coiled coil‐nucleotide‐binding‐leucine‐rich repeat, SD‐CNL) resistance genes. On transient expression of the NS
M
protein in tomato and transgenic
Nicotiana benthamiana
harbouring the
Sw‐5b
gene, a hypersensitive cell death response (HR) is triggered. Here, it is shown that high accumulation of the Sw‐5b protein in
N. benthamiana
leaves, achieved by co‐expression of the Sw‐5b protein with RNA silencing suppressors (RSSs), leads to auto‐activity in the absence of NS
M
. In a similar approach, Sw‐5a, the highest conserved paralogue of Sw‐5b from
Solanum peruvianum
, also triggered HR by auto‐activation, whereas the highest conserved orthologue from susceptible
S. lycopersicum
, named Sw‐5a
S
, did not. However, neither of the last two homologues was able to trigger an NS
M
‐dependent HR. Truncated and mutated versions of these Sw‐5 proteins revealed that the NB‐ARC [nucleotide‐binding adaptor shared by Apaf‐1 (from humans), R proteins and CED‐4 (from nematodes)] domain is sufficient for the triggering of HR and seems to be suppressed by the SD‐CC domain. Furthermore, a single mutation was sufficient to restore auto‐activity within the NB‐ARC domain of Sw‐5a
S
. When the latter domain was fused to the Sw‐5b LRR domain, NS
M
‐dependent HR triggering was regained, but not in the presence of its own Sw‐5a
S
LRR domain. Expression analysis
in planta
revealed a nucleocytoplasmic localization pattern of Sw‐5b, in which the SD‐CC domain seems to be required for nuclear translocation. Although the Sw‐5 N‐terminal CC domain, in contrast with
Rx
, contains an additional SD, most findings from this study support a conserved role of domains within NB‐LRR (NLR) proteins against plant viruses.