A common regulatory switch controls a suite of C4 traits in multiple cell types.

The C4 photosynthetic pathway provided a major advantage to plants growing in hot, dry environments, including the ancestors of our most productive crops. Two traits were essential for the evolution of this pathway: increased vein density and the functionalization of bundle sheath cells for photosynthesis. Although GRAS transcriptional regulators, including SHORT ROOT (SHR), have been implicated in mediating leaf patterning in both C3 and C4 species, little is known about what controls the specialized features of the cells that mediate C4 metabolism and physiology. We show in the model monocot, Setaria viridis, that SHR regulates components of multiple cell identities, including chloroplast biogenesis and photosynthetic gene expression in bundle sheath cells, a central feature of C4 plants. Furthermore, we found that it also contributes to the two-cell compartmentalization of the characteristic four-carbon shuttle pathway. Disruption of SHR function clearly reduced photosynthetic capacity and seed yield in mutant plants under heat stress. Together, these results show how cell identities are remodeled by SHR to host the suite of traits characteristic of C4 regulation, which are a main engineering target in non-C4 crops to improve climate resilience.

Challenging the water stress index concept: Thermographic assessment of Arabidopsis transpiration.

Water stress may greatly limit plant functionality and growth. Stomatal closure and consequently reduced transpiration are considered as early and sensitive plant responses to drought and salinity stress. An important consequence of stomatal closure under water stress is the rise of leaf temperature (Tleaf ), yet Tleaf is not only fluctuating with stomatal closure. It is regulated by several plant parameters and environmental factors. Thermal imaging and different stress indices, incorporating actual leaf/crop temperature and reference temperatures, were developed in previous studies toward normalizing for effects unassociated to water stress on Tleaf , aiming at a more efficient water stress assessment. The concept of stress indices has not been extensively studied on the model plant Arabidopsis thaliana. Therefore, the aim of this study was to examine the different indices employed in previous studies in assessing rosette transpiration rate (E) in Arabidopsis plants grown under two different light environments and subjected to salinity. After salinity imposition, E was gravimetrically quantified, and thermal imaging was employed to quantify rosette (Trosette ) and artificial reference temperature (Twet, Tdry ). Trosette and several water stress indices were tested for their relation to E. Among the microclimatic growth conditions tested, RWSI1 ([Trosette - Twet ]/[Tdry - Twet ]) and RWSI2 ([Tdry - Trosette ]/[Tdry - Twet ]) were well linearly-related to E, irrespective of the light environment, while the sole use of either Twet or Tdry in different combinations with Trosette returned less accurate results. This study provides evidence that selected combinations of Trosette , Tdry , and Twet can be utilized to assess E under water stress irrespective of the light environment.

Phytochrome A Protects Tomato Plants From Injuries Induced by Continuous Light.

Plants perceive and transduce information about light quantity, quality, direction and photoperiod via several photoreceptors and use it to adjust their growth and development. A role for photoreceptors has been hypothesized in the injuries that tomato plants develop when exposed to continuous light as the light spectral distribution influences the injury severity. Up to now, however, only indirect clues suggested that phytochromes (PHY), red/far-red photoreceptors, are involved in the continuous-light-induced injuries in tomato. In this study, therefore, we exposed mutant and transgenic tomato plants lacking or over-expressing phytochromes to continuous light, with and without far-red light enrichment. The results show that PHYA over-expression confers complete tolerance to continuous light regardless the light spectrum. Under continuous light with low far-red content, PHYB1 and PHYB2 diminished and enhanced the injury, respectively, yet the effects were small. These results confirm that phytochrome signaling networks are involved in the induction of injury under continuous light. HIGHLIGHTS: - PHYA over-expression confers tolerance to continuous light regardless the light spectrum.- In the absence of far-red light, PHYB1 slightly diminishes the continuous light-induced injury.- Continuous light down-regulates photosynthesis genes in sensitive tomato lines.

Sucrose and Starch Content Negatively Correlates with PSII Maximum Quantum Efficiency in Tomato (Solanum lycopersicum) Exposed to Abnormal Light/Dark Cycles and Continuous Light.

Light is most important to plants as it fuels photosynthesis and provides clues about the environment. If provided in unnatural long photoperiods, however, it can be harmful and even lethal. Tomato (Solanum lycopersicum), for example, develops mottled chlorosis and necrosis when exposed to continuous light. Understanding the mechanism of these injuries is valuable, as important pathways regulating photosynthesis, such as circadian, retrograde and light signaling pathways are probably involved. Here, we use non-targeted metabolomics and transcriptomics analysis as well as hypothesis-driven experiments with continuous light-tolerant and -sensitive tomato lines to explore the long-standing proposed role of carbohydrate accumulation in this disorder. Analysis of metabolomics and transcriptomics data reveals a clear effect of continuous light on sugar metabolism and photosynthesis. A strong negative correlation between sucrose and starch content with the severity of continuous light-induced damage quantified as the maximum quantum efficiency of PSII (Fv/Fm) was found across several abnormal light/dark cycles, supporting the hypothesis that carbohydrates play an important role in the continuous light-induced injury. We postulate that the continuous light-induced injury in tomato is caused by down-regulation of photosynthesis, showing characteristics of both cytokinin-regulated senescence and light-modulated retrograde signaling. Molecular mechanisms linking carbohydrate accumulation with down-regulation of carbon-fixing enzymes are discussed.

On the induction of injury in tomato under continuous light: circadian asynchrony as the main triggering factor.

Unlike other species, when tomato plants (Solanum lycopersicum L.) are deprived of at least 8h of darkness per day, they develop a potentially lethal injury. In an effort to understand why continuous light (CL) is injurious to tomato, we tested five factors, which potentially could be responsible for triggering the injury in CL-grown tomato: (i) differences in the light spectral distribution between sunlight and artificial light, (ii) continuous light signalling, (iii) continuous supply of light for photosynthesis, (iv) continuous photo-oxidative pressure and (v) circadian asynchrony - a mismatch between the internal circadian clock frequency and the external light/dark cycles. Our results strongly suggest that continuous-light-induced injury does not result from the unnatural spectral distribution of artificial light nor from the continuity of light per se. Instead, circadian asynchrony seems to be the main factor inducing the CL-induced injury, but the mechanism is not by the earlier hypothesised circadian pattern in sensitivity for photoinhibition. Here, however, we show for the first time diurnal fluctuations in sensitivity to photoinhibition during normal photoperiods. Similarly, we also report for the first time diurnal and circadian rhythms in the maximum quantum efficiency of PSII (Fv/Fm) and the parameter F0.

Continuous-light tolerance in tomato is graft-transferable.

Continuous light induces a potentially lethal injury in domesticated tomato (Solanum lycopersicum) plants. Recently, continuous-light tolerance was reported in several wild tomato species, yet the molecular mechanisms underpinning tolerance/sensitivity are still elusive. Here, we investigated from which part of the plant continuous-light tolerance originates and whether this trait acts systemically within the plant. By exposing grafted plants bearing both tolerant and sensitive shoots, the trait was functionally located in the shoot rather than the roots. Additionally, an increase in continuous-light tolerance was observed in sensitive plants when a continuous-light-tolerant shoot was grafted on it. Cultivation of greenhouse tomatoes under continuous light promises high yield increases. Our results show that to pursuit this, the trait should be bred into scion rather than rootstock lines. In addition, identifying the nature of the signal/molecule(s) and/or the mechanism of graft-induced, continuous-light tolerance can potentially result in a better understanding of important physiological processes like long-distance signaling.

A single locus confers tolerance to continuous light and allows substantial yield increase in tomato.

An important constraint for plant biomass production is the natural day length. Artificial light allows for longer photoperiods, but tomato plants develop a detrimental leaf injury when grown under continuous light--a still poorly understood phenomenon discovered in the 1920s. Here, we report a dominant locus on chromosome 7 of wild tomato species that confers continuous light tolerance. Genetic evidence, RNAseq data, silencing experiments and sequence analysis all point to the type III light harvesting chlorophyll a/b binding protein 13 (CAB-13) gene as a major factor responsible for the tolerance. In Arabidopsis thaliana, this protein is thought to have a regulatory role balancing light harvesting by photosystems I and II. Introgressing the tolerance into modern tomato hybrid lines, results in up to 20% yield increase, showing that limitations for crop productivity, caused by the adaptation of plants to the terrestrial 24-h day/night cycle, can be overcome.

Plants under continuous light.

Continuous light is an essential tool for understanding the plant circadian clock. Additionally, continuous light might increase greenhouse food production. However, using continuous light in research and practice has its challenges. For instance, most of the circadian clock-oriented experiments were performed under continuous light; consequently, interactions between the circadian clock and the light signaling pathway were overlooked. Furthermore, in some plant species continuous light induces severe injury, which is only poorly understood so far. In this review paper, we aim to combine the current knowledge with a modern conceptual framework. Modern genomic tools and rediscovered continuous light-tolerant tomato species (Solanum spp.) could boost the understanding of the physiology of plants under continuous light.