Reactive oxygen species triggering systemic programmed cell death process via elevation of nuclear calcium ion level in tomatoes resisting tobacco mosaic virus

Programmed cell death (PCD) plays a positive role in the systemic response of plants to pathogen resistance. It has been confirmed that local tobacco mosaic virus (TMV) infecting tomato leaves can induce systemic PCD process in root-tip tissues. But up to now the underlying physiological mechanisms are poorly understood. This study focused on the detailed investigation of the physiological responses of root-tip cells during the initiation of systemic PCD. Physiological, biochemical examination and cytological observation showed that 1 day post-inoculation (dpi) of TMV inoculation there was an increase in calcium fluorescence intensity in root tip tissue cells. Then at 2 dpi, 4 dpi, 8 dpi and 15 dpi, the fluorescence intensity of calcium ion continued to increase. However, at 5 dpi, the reactive oxygen species (ROS) began to accumulate in the root-tip cells. And finally at 20 dpi, the obvious PCD reaction was detected. In addition, the experimental results also showed that the above process involved the elevation of two types of intracellular Ca2+, including cytoplasmic calcium ([Ca2+]cyt) and nuclear calcium ([Ca2+]nuc). The [Ca2+]cyt, as a pilot signal could lead to the subsequent elevation of intracellular ROS concentration. Then, the high levels of ROS stimulated an increase of [Ca2+]nuc and eventually caused PCD reactions in the root-tip tissues. In particular, the high level of nuclear calcium is an essential mediator in systemic PCD of plants.