Plastids communicate their developmental and physiological status to the nucleus via retrograde signaling, allowing nuclear gene expression to be adjusted appropriately. Signaling during plastid biogenesis and responses of mature chloroplasts to environmental changes are designated \"biogenic\" and \"operational\" controls, respectively. A prominent example of the investigation of biogenic signaling is the screen for gun (genomes uncoupled) mutants. Although the first five gun mutants were identified 30 years ago, the functions of GUN proteins in retrograde signaling remain controversial, and that of GUN1 is hotly disputed. Here, we provide background information and critically discuss recently proposed concepts that address GUN-related signaling and some novel gun mutants. Moreover, considering heme as a candidate in retrograde signaling, we revisit the spatial organization of heme biosynthesis and export from plastids. Although this review focuses on GUN pathways, we also highlight recent progress in the identification and elucidation of chloroplast-derived signals that regulate the acclimation response in green algae and plants. Here, stress-induced accumulation of unfolded/misassembled chloroplast proteins evokes a chloroplast-specific unfolded protein response, which leads to changes in the expression levels of nucleus-encoded chaperones and proteases to restore plastid protein homeostasis. We also address the importance of chloroplast-derived signals for activation of flavonoid biosynthesis leading to production of anthocyanins during stress acclimation through sucrose non-fermenting 1-related protein kinase 1. Finally, a framework for identification and quantification of intercompartmental signaling cascades at the proteomic and metabolomic levels is provided, and we discuss future directions of dissection of organelle-nucleus communication.

The chloroplastic thioredoxins (Trxs), a family of thiol-disulphide oxidoreductases, are reduced by either ferredoxin (Fd)-dependent Trx reductase (FTR) or reduced nicotinamide adenine dinucleotide phosphate (NADPH)-dependent Trx reductase (NTR). Two Trx systems are present in chloroplasts including Trxs, Trx-like proteins, and reductase FTR and NTRC. FTR is the main reductant for Trxs in chloroplasts, while the flavoprotein NTRC integrates NTR and Trx activity, and plays multiple roles in the Calvin cycle, the oxidative pentose phosphate pathway (OPPP), anti-peroxidation, tetrapyrrole metabolism, ATP and starch synthesis, and photoperiodic regulation. In addition, not only there exists a reduction potential transfer pathway across the thylakoid membrane, but also FTR and NTRC coordinate with each other to regulate chloroplast redox homeostasis. Herein, we summarise the physiological functions of these two Trx reduction systems, discuss how both regulate redox homeostasis in plant plastids, and emphasize the significance of chloroplast thioredoxin systems in maintaining photosynthetic efficiency in plants.

The arrangement of mitochondria and chloroplasts, together with the relative resistances of cell wall and chloroplast, determine the path of diffusion out of the leaf for (photo)respired CO2. Traditional photosynthesis models have assumed a tight arrangement of chloroplasts packed together against the cell wall with mitochondria located behind the chloroplasts, deep inside the cytosol. Accordingly, all (photo)respired CO2 must cross the chloroplast before diffusing out of the leaf. Different arrangements have recently been considered, where all or part of the (photo)respired CO2 diffuses through the cytosol without ever entering the chloroplast. Assumptions about the path for the (photo)respiratory flux are particularly relevant for the calculation of mesophyll conductance (gm). If (photo)respired CO2 can diffuse elsewhere besides the chloroplast, apparent gm is no longer a mere physical resistance but a flux-weighted variable sensitive to the ratio of (photo)respiration to net CO2 assimilation. We discuss existing photosynthesis models in conjunction with their treatment of the (photo)respiratory flux and present new equations applicable to the generalized case where (photo)respired CO2 can diffuse both into the chloroplast and through the cytosol. Additionally, we present a new generalized Δ13C model that incorporates this dual diffusion pathway. We assess how assumptions about the fate of (photo)respired CO2 affect the interpretation of photosynthetic data and the challenges it poses for the application of different models.

Retrograde signaling, defined as the signaling events leading from the plastids to the nucleus, coordinates the expression of plastid and nuclear genes and is crucial for metabolic as well as developmental processes of the plastids. In the recent past, the identification of various components that are involved in the generation and transmission of plastid-originated retrograde signals and the regulation of nuclear gene expression has only provided a glimpse of the plastid retrograde signaling network, which remains poorly understood. The basic assumptions underlying our current understanding of retrograde signaling stayed untouched for many years. Therefore, an attempt has been made in this review article to summarize established facts and recent advances regarding various retrograde signaling pathways derived from different sources, the identification of key elements mediating retrograde signal transduction and also to give an overview of possible signaling molecules that remain to be investigated.

Plants, as sessile organisms, have evolved an exquisitely tuned response network to survive environmental perturbations. Organelles-to-nucleus signaling, termed retrograde signaling, plays a key role in stress responses by communicating subcellular perturbations to the nucleus, thereby coordinating expression of stress specific nuclear genes essential for adaptive responses to hostile environment. Recently, several stress specific retrograde signals have been identified; most notable amongst them are reactive oxygen species, tetrapyrroles, 2-C-methyl-d-erythritol 2,4-cyclodiphosphate (MEcPP), unsaturated fatty acids, nitric oxide (NO), 3\'-phosphoadenosine 5\'-phosphate (PAP), and β-cyclocitral (β-CC). It is expected that this trend will continue to provide fundamental insight into the integrative network of sensory systems central to the adaptive responses of plants to the prevailing environment. This review focuses on the recent advancements in the field.