Abstract
The most conspicuous function of plastids is the oxygenic photosynthesis of chloroplasts, yet plastids are super-factories that produce a plethora of compounds that are indispensable for proper plant physiology and development. Given their origins as free-living prokaryotes, it is not surprising that plastids possess their own genomes whose expression is essential to plastid function. This semi-autonomous character of plastids requires the existence of sophisticated regulatory mechanisms that provide reliable communication between them and other cellular compartments. Such intracellular signaling is necessary for coordinating whole-cell responses to constantly varying environmental cues and cellular metabolic needs. This is achieved by plastids acting as receivers and transmitters of specific signals that coordinate expression of the nuclear and plastid genomes according to particular needs. In this review we will consider the so-called retrograde signaling occurring between plastids and nuclei, and between plastids and other organelles. Another important role of the plastid we will discuss is the involvement of plastid signaling in biotic and abiotic stress that, in addition to influencing retrograde signaling, has direct effects on several cellular compartments including the cell wall. We will also review recent evidence pointing to an intriguing function of chloroplasts in regulating intercellular symplasmic transport. Finally, we consider an intriguing yet less widely known aspect of plant biology, chloroplast signaling from the perspective of the entire plant. Thus, accumulating evidence highlights that chloroplasts, with their complex signaling pathways, provide a mechanism for exquisite regulation of plant development, metabolism and responses to the environment. As chloroplast processes are targeted for engineering for improved productivity the effect of such modifications on chloroplast signaling will have to be carefully considered in order to avoid unintended consequences on plant growth and development.
Highlights
According to the endosymbiotic theory plastids originated from free-living cyanobacteria that were engulfed by early eukaryotic cells
RNADEPENDENT RNA POLYMERASE1 (RDR1) does not seem to have a role in drought resistance (Hunter et al, 2013), other components of the RNAi components have been demonstrated to function in stress tolerance (Earley et al, 2010; Li et al, 2012; Westwood et al, 2013)
The first committed step in abscisic acid (ABA) synthesis is the cleavage of the carotenoid xanthopyhll by 9-cis-epoxycarotenoid dioxygenase (NCED) to produce the C15 compound xanthoxin, that is transported from the plastids into the cytosol by an unknown mechanism, where it will be converted into ABA (Finkelstein, 2013)
Summary
According to the endosymbiotic theory plastids originated from free-living cyanobacteria that were engulfed by early eukaryotic cells. An often over-looked aspect of cell biology is the physical interaction between organelles for coordination of signaling and metabolism (Figure 1B) This area is beginning to receive attention and we will examine these findings as they relate to chloroplasts and their roles in signaling. Given the dizzying array of signals that chloroplasts respond to and produce it is not surprising that there is considerable crosstalk between signaling pathways. This is evident during responses to biotic and abiotic stress (Nakashima et al, 2014; Trotta et al, 2014; Zhou et al, 2015). These consequences would not be limited to the chloroplasts, but could extend even to other plants cultivated in the vicinity of the engineered plants
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