The roles of tetrapyrroles in plastid retrograde signaling and tolerance to environmental stresses.

MAIN CONCLUSION: This review provides new insights that tetrapyrrole signals play important roles in nuclear gene expression, chloroplast development and plant's resistance to environmental stresses. Higher plants contain many tetrapyrroles, including chlorophyll (Chl), heme, siroheme, phytochromobilin and some of their precursors, all of which have important biological functions. Genetic and physiological studies indicated that tetrapyrrole (mainly Mg-protoporphyrin IX) retrograde signals control photosynthesis-associated nuclear gene (PhANG) expression. Recent studies have shown that tetrapyrrole-derived signals may correlate with plant resistance to environmental stresses such as drought, high-light stress, water stress, osmotic stress, salinity and heavy metals. Signaling and physiological roles of Mg-protoIX-binding proteins (such as PAPP5, CRD and HSP90) and heme-binding proteins (such as HO and TSPO) and tetrapyrrole-signaling components (such as GUN1, ABI4 and CBFA) are summarized. Some of them positively regulate plant development and response to environmental stresses. The intermediate signaling components (such as PTM, HSP70-HSP90-HAP1 complex and PAPP5) between the nucleus and the plastid also positively regulate plant resistance to environmental stresses. This review provides new insights that genetically modified plants with enhanced tetrapyrrole levels have improved resistance to environmental stresses.