ELONGATED HYPOCOTYL5 (HY5) and HY5 HOMOLOGUE (HYH) maintain shade avoidance suppression in UV-B.

Reductions in red to far-red ratio (R:FR) provide plants with an unambiguous signal of vegetational shade and are monitored by phytochrome photoreceptors. Plants integrate this information with other environmental cues to determine the proximity and density of encroaching vegetation. Shade-sensitive species respond to reductions in R:FR by initiating a suite of developmental adaptations termed shade avoidance. These include the elongation of stems to facilitate light foraging. Hypocotyl elongation is driven by increased auxin biosynthesis promoted by PHYTOCHROME INTERACTING FACTORs (PIF) 4, 5 and 7. UV-B perceived by the UV RESISTANCE LOCUS 8 (UVR8) photoreceptor rapidly inhibits shade avoidance, in part by suppressing PIF4/5 transcript accumulation and destabilising PIF4/5 protein. Here, we show that longer-term inhibition of shade avoidance is sustained by ELONGATED HYPOCOTYL 5 (HY5) and HY5 HOMOLOGUE (HYH), which regulate transcriptional reprogramming of genes involved in hormone signalling and cell wall modification. HY5 and HYH are elevated in UV-B and suppress the expression of XYLOGLUCAN ENDOTRANSGLUCOSYLASE/HYDROLASE (XTH) genes involved in cell wall loosening. They additionally increase expression GA2-OXIDASE1 (GA2ox1) and GA2ox2, encoding gibberellin catabolism enzymes that act redundantly to stabilise the PIF-inhibiting DELLA proteins. UVR8 therefore regulates temporally distinct signalling pathways to first rapidly inhibit and subsequently maintain suppression of shade avoidance following UV-B exposure.

Cell wall fucosylation in Arabidopsis influences control of leaf water loss and alters stomatal development and mechanical properties.

The Arabidopsis sensitive-to-freezing8 (sfr8) mutant exhibits reduced cell wall (CW) fucose levels and compromised freezing tolerance. To examine whether CW fucosylation also affects the response to desiccation, we tested the effect of leaf excision in sfr8 and the allelic mutant mur1-1. Leaf water loss was strikingly higher than in the wild type in these, but not other, fucosylation mutants. We hypothesized that reduced fucosylation in guard cell (GC) walls might limit stomatal closure through altering mechanical properties. Multifrequency atomic force microscopy (AFM) measurements revealed a reduced elastic modulus (E'), representing reduced stiffness, in sfr8 GC walls. Interestingly, however, we discovered a compensatory mechanism whereby a concomitant reduction in the storage modulus (E'') maintained a wild-type viscoelastic time response (tau) in sfr8. Stomata in intact leaf discs of sfr8 responded normally to a closure stimulus, abscisic acid, suggesting that the time response may relate more to closure properties than stiffness does. sfr8 stomatal pore complexes were larger than those of the wild type, and GCs lacked a fully developed cuticular ledge, both potential contributors to the greater leaf water loss in sfr8. We present data that indicate that fucosylation-dependent dimerization of the CW pectic domain rhamnogalacturonan-II may be essential for normal cuticular ledge development and leaf water retention.

Countering elevated CO2 induced Fe and Zn reduction in Arabidopsis seeds.

Growth at increased concentrations of CO2 induces a reduction in seed zinc (Zn) and iron (Fe). Using Arabidopsis thaliana, we investigated whether this could be mitigated by reducing the elevated CO2 -induced decrease in transpiration. We used an infrared imaging-based screen to isolate mutants in At1g08080 that encodes ALPHA CARBONIC ANHYDRASE 7 (ACA7). aca7 mutant alleles display wild-type (WT) responses to abscisic acid (ABA) and light but are compromised in their response to elevated CO2 . ACA7 is expressed in guard cells. When aca7 mutants are grown at 1000 ppm CO2 they exhibit higher transpiration and higher seed Fe and Zn content than WT grown under the same conditions. Our data show that by increasing transpiration it is possible to partially mitigate the reduction in seed Fe and Zn content when Arabidopsis is grown at elevated CO2 .

ABA signalling and metabolism are not essential for dark-induced stomatal closure but affect response speed.

Stomata are microscopic pores that open and close, acting to balance CO2 uptake with water loss. Stomata close in response to various signals including the drought hormone abscisic acid (ABA), microbe-associated-molecular-patterns, high CO2 levels, and darkness. The signalling pathways underlying ABA-induced stomatal closure are well known, however, the mechanism for dark-induced stomatal closure is less clear. ABA signalling has been suggested to play a role in dark-induced stomatal closure, but it is unclear how this occurs. Here we investigate the role of ABA in promoting dark-induced stomatal closure. Tracking stomatal movements on the surface of leaf discs we find, although steady state stomatal apertures are affected by mutations in ABA signalling and metabolism genes, all mutants investigated close in response to darkness. However, we observed a delayed response to darkness for certain ABA signalling and metabolism mutants. Investigating this further in the quadruple ABA receptor mutant (pyr1pyl1pyl2pyl4), compared with wild-type, we found reduced stomatal conductance kinetics. Although our results suggest a non-essential role for ABA in dark-induced stomatal closure, we show that ABA modulates the speed of the dark-induced closure response. These results highlight the role of ABA signalling and metabolic pathways as potential targets for enhancing stomatal movement kinetics.

Short- and Long-Term Effects of UVA on Arabidopsis Are Mediated by a Novel cGMP Phosphodiesterase.

Although UVA radiation (315-400 nm) represents 95% of the UV radiation reaching the earth's surface, surprisingly little is known about its effects on plants [1]. We show that in Arabidopsis, short-term exposure to UVA inhibits the opening of stomata, and this requires a reduction in the cytosolic level of cGMP. This process is independent of UVR8, the UVB receptor. A cGMP-activated phosphodiesterase (AtCN-PDE1) was responsible for the UVA-induced decrease in cGMP in Arabidopsis. AtCN-PDE1-like proteins form a clade within the large HD-domain/PDEase-like protein superfamily, but no eukaryotic members of this subfamily have been functionally characterized. These genes have been lost from the genomes of metazoans but are otherwise conserved as single-copy genes across the tree of life. In longer-term experiments, UVA radiation increased growth and decreased water-use efficiency. These experiments revealed that PDE1 is also a negative regulator of growth. As the PDE1 gene is ancient and not represented in animal lineages, it is likely that at least one element of cGMP signaling in plants has evolved differently to the system present in metazoans.

How Arabidopsis Talks to Itself about Its Water Supply.

Takahashi et al. (2018) report that the peptide CLE25 together with the BAM1, BAM3 LRR receptor-like kinases are involved in root-to-shoot communication during dehydration stress in Arabidopsis.