Singlet oxygen-triggered chloroplast-to-nucleus retrograde signalling pathways: An emerging perspective

Singlet oxygen (¹ O₂ ) is a prime cause of photo-damage of the photosynthetic apparatus. The chlorophyll molecules in the photosystem II reaction center and in the light-harvesting antenna complex are major sources of ¹ O₂ generation. It has been thought that the generation of ¹ O₂ mainly takes place in the appressed regions of the thylakoid membranes, namely, the grana core, where most of the active photosystem II complexes are localized. Apart from being a toxic molecule, new evidence suggests that ¹ O₂ significantly contributes to chloroplast-to-nucleus retrograde signalling that primes acclimation and cell death responses. Interestingly, recent studies reveal that chloroplasts operate two distinct ¹ O₂ -triggered retrograde signalling pathways in which β-carotene and a nuclear-encoded chloroplast protein EXECUTER1 play essential roles as signalling mediators. The coexistence of these mediators raises several questions: their crosstalk, source(s) of ¹ O₂ , downstream signalling components, and the perception and reaction mechanism of these mediators towards ¹ O₂ . In this review, we mainly discuss the molecular genetic basis of the mode of action of these two putative ¹ O₂ sensors and their corresponding retrograde signalling pathways. In addition, we also propose the possible existence of an alternative source of ¹ O₂ , which is spatially and functionally separated from the grana core.