The significance of research conducted on Arabidopsis thaliana cannot be overstated. This focus issue showcases how insights from Arabidopsis have opened new areas of biology and directly advanced our understanding of crops. Here, experts intimately involved in bridging between Arabidopsis and crops share their perspectives on the challenges and opportunities for translation. First, we examine the translatability of genetic modules from Arabidopsis into maize, emphasizing the need to publish well-executed translational experiments, regardless of outcome. Second, we highlight the landmark success of HB4, the first GM wheat cultivar on the market, whose abiotic tolerance is borne from direct translation and based on strategies first outlined in Arabidopsis. Third, we discuss the decades-long journey to engineer oilseed crops capable of producing omega-3 fish oils, with Arabidopsis serving as a critical intermediary. Fourth, we explore how direct translation of starch synthesizing proteins characterized in Arabidopsis helped uncover novel mechanisms and functions in crops, with potential valuable applications. Finally, we illustrate how shared molecular factors between Arabidopsis and barley exhibit distinct molecular wiring as exemplified in cuticular and stomatal development. Together, these vignettes underscore the pivotal role of Arabidopsis as a foundational model plant while highlighting the challenges of translating discoveries into field-ready, commercial cultivars with enhanced knowledge-based traits.

The cuticle covering aerial organs of land plants is well known to protect against desiccation. Cuticles also play diverse and specialized functions, including organ separation, depending on plant and tissue. Barley shows a distinctive cuticular wax bloom enriched in β-diketones on leaf sheaths, stem nodes and internodes and inflorescences. Barley also develops a sticky surface on the outer pericarp layer of its grain fruit leading to strongly adhered hulls, ‘covered grain’, important for embryo protection and seed dispersal. While the transcription factor-encoding gene HvNUDUM (HvNUD) appears essential for adherent hulls, little is understood about how the pericarp cuticle changes during adhesion or whether changes in pericarp cuticles contribute to another phenotype where hulls partially shed, called ‘skinning’. To that end, we screened barley lines for hull adhesion defects, focussing on the Eceriferum (= waxless, cer) mutants. Here, we show that the cer-xd allele causes defective wax blooms and compromised hull adhesion, and results from a mutation removing the last 10 amino acids of the GDS(L) [Gly, Asp, Ser, (Leu)]-motif esterase/lipase HvGDSL1. We used severe and moderate HvGDSL1 alleles to show that complete HvGDSL1 function is essential for leaf blade cuticular integrity, wax bloom deposition over inflorescences and leaf sheaths and pericarp cuticular ridge formation. Expression data suggest that HvGDSL1 may regulate hull adhesion independently of HvNUD. We found high conservation of HvGDSL1 among barley germplasm, so variation in HvGDSL1 unlikely leads to grain skinning in cultivated barley. Taken together, we reveal a single locus which controls adaptive cuticular properties across different organs in barley.

The cuticle is a chemically complex, lipid-rich seal covering aerial tissues in land plants. Secreted by the outer epidermis, cuticles protect against desiccation, organ fusion, pathogen attack and UV radiation exposure. Cuticles also mediate key interactions during reproduction and play other roles depending on the tissue, stage, and species. Reflecting these diverse functions, cuticle composition and structure can vary widely. We review the transcriptional control important for cuticle formation, with special attention to broadly conserved transcription factors where selective regulation of downstream targets drive cuticle specialization. We also discuss how transcriptional control of the cuticle intersects with networks controlling plasticity and cell fate, and explore their translational potential for crop improvement.

Summary: Cereal grain is a global food staple. Filial tissues in grain, such as the embryo and endosperm, grow embedded in the maternal ovule and surrounding ovary wall. Maternal tissues provide nutrition and protection but are progressively reduced and degraded as filial tissues expand, driving grain expansion between floret hulls. While we know some enzymatic players, regulation of this process remains less understood. However, our previous work in barley, Hordeum vulgare spp. vulgare suggested a role for the phytohormone jasmonate (JA). We edited the barley ALLENE OXIDASE CYCLASE ( HvAOC ) gene encoding a JA biosynthetic enzyme. We examined hvaoc mutants for developmental phenotypes, genetic interactions and effects on the transcriptome. Defective HvAOC alleles reduced JA levels, and caused enhanced vegetative growth, faster flowering and lengthened grain, all phenotypes rescued by MeJA treatment. HvAOC control of grain length appears parentally‐derived, consistent with enriched HvAOC expression in maternal compared to filial tissues. Grain dimension changes were uncoupled from hull size and correlated with larger endosperm cavities, more persistent maternal tissues and higher expression of genes linked to sucrose transport. This work provides the first genetic dissection of JA in barley and highlights roles in coordinating maternal tissues within grain.

Summary: Land plants secrete a protective outer cuticular layer with diverse functions. Barley ( Hordeum vulgare L.) develops two cuticular specialisations: the β‐diketone rich wax bloom on vegetative tissues and an adherent grain surface which sticks to the hulls, leading to barley's distinctive ‘covered’ grain phenotype. Two barley SHINE transcription factors, HvWIN1 and NUD, promote the wax bloom and covered grain phenotypes, respectively, yet we know little about other genes involved. We investigated a barley mutant showing defects in both cuticular specialisations to better understand the networks and processes underlying these traits. We identified the barley BODYGUARD1 ( HvBDG1 ) gene encoding an α/β hydrolase as crucial for leaf cuticular integrity and wax‐bloom deposition. Mining tetraploid wheat ( Triticum turgidum ssp. durum ) mutant populations demonstrated that BDG1 and WIN1 orthologues also control the wax bloom in wheat. We further reveal that NUD and HvWIN1 retain functional, independent overlap in leaf cuticle integrity and hull adhesion in barley, functions which involve upregulation of HvBDG1 and other direct or indirect targets, shared and distinct, depending on the stage and tissue. Our work greatly expands our knowledge of genetic and developmental mechanisms underlying shared and distinctive cuticular features in plants.

The outer epidermis of land plants secretes a cuticular layer, a hydrophobic diffusion barrier which minimises water loss into the atmosphere and protects from pests, ultraviolet light and organ fusion. Cuticles typically comprise a polyester cutin matrix embedded and overlaid with cuticular waxes, but their exact chemical make-up, structure and functions can vary widely depending on the tissue and species. Barley shows two such cuticular specialisations: (1) deposition of a thick β-diketone-rich wax bloom on multiple organs at reproductive stage, common in other Poeceae species and linked to yield; and, (2) secretion of a sticky layer on the grain fruit (caryopsis) pericarp cuticle which adheres to inner floral hulls, leading to barley’s distinctive ‘covered’ grain used in animal feed and malting. Two SHINE/WAX-INDUCER transcription factors in barley, HvWIN1 and NUD, promote the wax bloom and hull to caryopsis adhesion, respectively, yet little is understood about other genes involved. Leveraging near-isogenic lines of wax-deficient mutants, we identify the barley BODYGUARD1 ( HvBDG1 ) gene encoding an α/β-hydrolase essential for leaf cuticular integrity and wax bloom deposition. Modelling of functional and defective alleles suggests that HvBDG1 N-terminal region control of protein flexibility is important for HvBDG1 function. In addition to their role in controlling barley epicuticular wax deposition, we show that both HvBDG1 and HvWIN1 are essential for strong hull to caryopsis adhesion. Along with NUD, these gene products differentially contribute to ultrastructural changes on the pericarp associated with a cuticular building programme driven by NUD and HvWIN1 regulation of cuticle metabolism and transport and cell wall-related genes, and correlate with shifts in pericarp surface chemistry. We also show that the previously asserted ‘grain-specific’ role of NUD should be revised, as our findings reveal that it is essential for maintaining leaf cuticle integrity. Our analyses in barley suggest that NUD and HvWIN1 control cuticular specialisations and cuticle integrity in part via promotion of HvBDG1 expression, while HvWIN1 and NUD likely act independently from each other. Lastly, mining tetraploid wheat mutant populations followed by crossing to combine mutated homoeologues demonstrated that BDG1 and WIN1 orthologues also control wax bloom in wheat. Taken together, our work greatly expands the genetic networks and molecular activities important for cuticle development in cereals and the underlying mechanisms for both shared and species-specific cuticular specialisations.