Natural thermometers: chloroplasts as heat sensors.

Abstract

All organisms need to sense their surroundings and respond. This is especially true for plants, which are rooted to the ground and must contend with daily fluctuations of both above- and belowground environments. Of these, temperature is perhaps the most variable, with shifts throughout the day and microenvironments that affect one part of the plant differently from another (e.g., a leaf in the canopy vs. one in the shade). Heat stress is of particular concern and negatively affects basic cellular processes including protein folding and function, membrane fluidity, and cytoskeletal organization. The resulting dysfunctions impair growth through metabolic imbalances and the generation of reactive oxygen species (ROS) like hydrogen peroxide (H2O2). As such, plants have evolved numerous genetic and cellular mechanisms to sense heat and mitigate the adverse effects. Much progress has been made on understanding thermoprotective mechanisms, but how plants initially sense heat stress is still poorly understood. It is becoming clear, however, that multiple overlapping mechanisms are important (Hayes et al., 2021). For instance, photoreceptors, the circadian clock evening complex, RNA-based temperature switches, epigenetic mechanisms involving histone methylation, and the unfolded protein response can respond to heat and affect plant growth and development. More recent research has demonstrated that chloroplasts (specialized plastids that perform photosynthesis in plants and algae) also play important roles in sensing heat. These organelles are ideal temperature sensors for the cell as photosynthesis is particularly susceptible to heat. Under increased temperatures, it will rapidly produce ROS and other metabolites that can act as stress signaling molecules. In support of this model, two recent studies have revealed novel post-translational mechanisms within chloroplasts that allow cells to monitor heat stress and initiate a response. In the first, it was demonstrated that heat-induction of the toxic metabolite methylglyoxal (MG) affects thermotolerance in Arabidopsis thaliana by modifying the chloroplast protein import machinery, thereby reducing chloroplast function and photosynthesis (Ding et al., 2026).