Brief Summary of Minutes of Annual Meeting

The 2025 NC-1203 meeting was held as a hybrid in-person and Zoom meeting on September 13-14, 2025, at Kansas State University in Manhattan, Kansas. Presentations of research progress and future directions were presented by the participants throughout the society meeting. Overall future research directions and writing of the renewal during Fall 2025 was discussed. Future meeting sites: In 2026 the meeting will be held at the University of Nevada, Reno (hosted by Dylan Kosma), and in 2027 the meeting will be held in conjunction with the 2027 Gordon Research Conference on Plant Lipids, in Pomona California, with more than half of the NC-1203 members planning to attend. Philip Bates will host the 2027 business meeting. The 2028 meeting will be held at the  University of Missouri and be hosted by Jay Thelen.

Research Activities and outputs:

Objective 1: Improve and extend methods for lipid characterization and measurement

Significant progress was made this year toward enhancing lipid and protein characterization techniques across diverse plant systems, with innovations that support both fundamental research and applied breeding. Researchers developed and refined analytical platforms that improve speed, increase the specificity of compound identification, enhance quantification and standardization, and extend approaches to measure additional plant tissue types.

Bates lab developed GC-FID based derivatization methods for rapid characterization of whole seed fatty acid composition and quantity for a variety of current and emerging oilseed crops including Camelina sativa, Thlaspi avernse (pennycress), Cuphea viscosissima, Brassica napus (var. Canola), Limnanthes alba (Meadowfoam) and Cannabis sativa L. (hemp) (Garneau, et al. 2025). This method will allow quicker screening of seed lipid content during research and breeding applications.

The Lee group developed a new mass spectrometry imaging (MSI) technique that can determine carbon-carbon double bond positions of PC lipids, called OzMALDI, by introducing ozone gas into MALDI source (Rensner, et al., 2025). Using this method, we could successfully visualize PC double-bond isomers in camelina and soybean seeds, engineered by Cahoon’s lab. Also demonstrated by the group is the use of unsupervised machine learning for the MSI data analysis with in vivo isotope labeling (Johnson, et al., 2025).

The Kosma Lab developed a GC-MS-based method for a complete characterization of external (pollenkitt) and internal fatty acids and specialized metabolites of pollen that requires a relatively small amount of pollen (~4-5 mg). This method has been used to investigate commercial and wild (Great Basin native) bee pollinated plant species. This will enable studies aimed at understanding pollen nutrition for pollinators in both ecological and agricultural contexts.

The Thelen lab has developed a multiplexed AQUA-MRM, LC-MS/MS assay for absolute quantitation of Arabidopsis acetyl-CoA carboxylase catalytic and effector proteins and demonstrated the utility of this approach to study the spatiotemporal regulation of this multienzyme complex that catalyzes the committed step of de novo fatty acid synthesis. 

The Welti lab is working with the Plant and Algal Lipid Interest Group of the International Lipidomics Society toward standardizing lipidomics results and creating reference mixtures. The group members have begun a multi-stage ring trial, in which they are analyzing the same samples.

Together, these advances position the project to deliver robust, scalable, and integrative analytical tools that support breeding, metabolic engineering, ecological research, and global data harmonization.

Objective 2: Identify lipid-related mechanisms to increase agricultural resilience

This year’s research advanced our understanding of how lipid metabolism contributes to plant resilience under environmental stress, nutrient limitation, and biotic pressure. Through multi-institutional collaborations, investigators explored genetic, enzymatic, structural, and signaling mechanisms that shape lipid function in plant defense, development, and stress adaptation. A major focus was uncovering metabolic bottlenecks and adaptations triggered by genetic modification.

Genetic changes due to breeding or bioengineering can lead to unexpected metabolic adaptations that may create bottlenecks to the desired metabolic accumulation. Understanding these metabolic adaptations is the first step in developing metabolism based rational engineering or breeding approaches. In 2025 three publications from multi-state collaborations investigated metabolic adaptations due to mutation or engineering of lipid metabolism. The Allen and Bates labs collaborated to investigate bottlenecks to lipid accumulation in vegetative tissues in tobacco. Key results indicate that oil-starch futile cycling can inhibit photosynthesis limiting total plant growth and oil accumulation, but stabilization of oil content limits the futile cycle (Johnson et. al., 2025). Additionally, heat stress can limit total oil accumulation (Murphy et. al., 2025). The Thelen, Allen, Koo, and Bates labs took a multi-omic approach to understand the effect of mutations of negative regulators of acetyl-CoA carboxylase the key regulatory point of fatty acid biosynthesis. Key results indicate that large changes to central carbon metabolism were induced that ultimately led to increases in both seed oil and protein content in the model plant Arabidopsis (Kataya et. al., 2025). Since activity of the biotin carboxylase subunit (BC) of acetyl-CoA carboxylase by effectors regulates fatty acid synthesis, Yokom determined the structure of the dimer of BC by cryo-EM. The Van Doren group quantified the accessibility of the BC surface using mass spectra collected by the Thelen group. Van Doren used crosslinks of BC identified by mass spec at PNNL to dock two of Yokom’s dimer of BC into a tetramer. Three of four potential binding sites appear to be available for effectors to bind, in principle, in the tetramer.

The Bates and Parchuri labs collaborated to characterize novel functionalities of key enzymes of oil biosynthesis DGAT1 and DGAT2 in a wide range of oil accumulating plants including Camelina sativa, pennycress, cotton, peanut, tung tree, castor. The results indicate never before recognized diversity in substrate selectivity for the sn-1,2 and sn-2,3 enantiomers of diacylglycerol the precursor to triacylglycerol. These results imply that different species may use a variety of mechanism in producing the final oil composition including triacylglycerol remodeling, and thus future bioengineering strategies to control oil composition will likely need to be crop specific. Additionally, the Bates lab discovered the key regulatory elements for DGAT1 expression in pollen, allowing for the first time complete replacement of endogenous oil biosynthetic enzymes with exogenous enzymes (McGuire et. al., 2025).

The Koo lab is investigating how GLUTAMATE RECEPTOR LIKE (GLR) proteins mediate the systemic wound response for jasmonate (JA) biosynthesis. Through a structure-function analysis, they identified a region that, when deleted, eliminates systemic JA biosynthesis. In collaboration with the Peck lab, they confirmed the correct targeting of the mutated GLR to the plasma membrane, validating the results of the GLR variants. In addition, the Koo and Peck labs collaborated to discover that a higher order mutant of a group of calcium signaling proteins may be involved in systemic wound response. Separately, the Koo lab has identified a novel chemical inhibitor that targets JA pathway and suppresses the wound response.  

The Wang Lab investigated the role of membrane phospholipid remodeling in rapeseed and camelina response to phosphate deficiency, The results show that the phosphate starvation induced phospholipase C, NPC4, promotes not only membrane lipid remodeling , but also P remobilization from old, senescent, source tissues to young, growing, sink tissues under P deficiency (Li et al., 2025). Manipulations of the pathway have potential to enhance crop plant growth and seed production, particularly under P-deficient conditions (Yang et al., 2025). Welti lab and Turkish collaborators investigated the effect of salicylic acid priming of wheat seeds on wheat tolerance for cadmium and showed that the positive effects of priming do not seem to be mediated through lipid alterations (Colak et al., 2025).

The Roston lab made progress in understanding freezing-activated galactolipid remodeler SFR2 as a reporter to uncover how plants gauge cold severity, employing yeast two hybrid analysis to identify yet another kinase potentially involved that was not previously associated with cold tolerance. Current data support a kinase/phosphatase pathway that modulates membrane repair in proportion to stress intensity. A future collaboration is planned with the Thelen lab to investigate the role of the kinases more broadly. In sorghum, time-bounded multi-omics found that while some cold responses to acclimation are conserved, many metabolite shifts are line-specific; photosynthetic readouts were reproducible only when measured within ~3 h of treatment, revealing strong time-of-day constraints, and lipid results were different between two tolerant lines. These findings motivated an updated model—daily temperature cycles synchronize lipid and soluble-metabolite programs essential for tolerance—which the Roston lab is now testing in collaboration with others at Nebraska and Frank Harmon of USDA.

The Louis Lab, in collaboration with the Cahoon and Welti Labs, recently identified a key role for triacylglycerols (TAGs), whose major constituents are fatty acids, in sorghum defense against the sugarcane aphid (SCA; Melanaphis sacchari). To investigate this, we employed electrospray ionization mass spectrometry (ESI-MS) to compare lipid profile alterations in SCA-resistant (SC265) and SCA-susceptible (SC1345) sorghum genotypes before and after aphid feeding. Our preliminary analyses revealed that sorghum lines differing in resistance also exhibit distinct lipid signatures. Notably, SCA feeding caused significant changes in TAG accumulation. While basal TAG levels were higher in the susceptible SC1345 line, seven days of SCA feeding led to a sharp reduction in several molecular species of TAGs in this genotype. In contrast, the resistant SC265 line maintained relatively stable TAG profiles under infestation. These findings suggest that TAG metabolism, and possibly TAG-derived metabolites, play a critical role in sorghum defense against SCA. Ongoing work, in collaboration with the Cahoon Lab at UNL, is focused on elucidating the specific mechanisms underlying this TAG-mediated defense. 

The Hoffmann-Benning lab has made progress in their Molecular Dynamics (MD) Modeling approach. They were able to show that PLAFP not only inserts into the membrane to detect and remove phosphatidic acid, it also dissociates from the membrane once the lipid is bound. This supports the hypothesis that protein and lipid jointly serve as a long-distance signal. To further investigate, they designed light-regulated gene expression for PLAFP and two control genes (FT, GRIP1) and were able to show movement of PLAFP throughout the plant. Gene-expression studies are currently in progress to support those findings.

The Schrick lab initiated MD simulations of lysophospholipid binding to the steroidogenic acute regulatory protein (StAR)-related lipid transfer (START) domain from homeodomain leucine-zipper (HD-Zip) IV transcription factor PDF2. These studies follow experimental data showing that PDF2 recruits lysophosphatidylcholine 18:1 in vivo and in vitro (Wojciechowska et al., 2024). The MD simulations address both lipid and membrane binding activities of the START domain and suggest that membrane contact occurs through key arginine residues in predicted loops on the periphery of the binding pocket. A separate study found that nuclear localization of HD-Zip IV proteins occurs through Importin alpha-mediated recognition of a classic nuclear localization sequence that overlaps with the DNA binding domain (Ahmad et al., 2024).

The Huang Lab is studying the role of the lipid droplets during plant-microbe interactions of important fungal diseases of soybean, tomato and strawberry. The group screened Plant-derived lipid biosynthesis inhibitors and identified a batch of inhibitors, including flavonoids, phthalides, fatty acids, and terpenes, that exhibited antimicrobial or toxin-suppressing effects. The tested fungal pathogens include Cercospora kikuchii, C. cf. flagellaris, and C. sojina, which cause Cercospora Leaf Blight and Frogeye Leaf Spot on soybeans. Additionally, Botrytis cinerea and Agroathelia rolfsii, which cause gray mold and southern blight diseases, respectively, have a broad host range; aggressive Neopestalotiopsis sp., which was reported as an emerging strawberry disease in recent years. A future collaboration is planned with Sanjaya and Welti to explore the functions of lipids in defense.

The Yandeau-Nelson lab, in partial collaboration with previous NC-1203 member Basil Nikolau, have assessed the role of the genetic redundancy in cuticular wax biosynthesis in maize, focusing on the 28 genes encoding ketoacyl-CoA synthetases. Taking a synthetic biology approach, they have tested different iterations of the fatty acid elongase pathway from maize (i.e. ZmFAE) in yeast and have demonstrated differences in very long chain fatty acid profiles among the different ZmFAE combinations. The Yandeau-Nelson team has also functionally characterized maize glossy2 and glossy2-like within the ZmFAE systems, and also in single and double mutants in maize.  Importantly, cuticular wax structure changes in gl2; gl2-like double mutants and is accompanied by a change in the cuticle’s function as a water barrier. A future collaboration is planned with the Louis lab to assess whether these mutants experience changed aphid-plant interactions.

Together, these discoveries highlight the central role of lipid pathways in plant resilience—from membrane integrity and nutrient remobilization to stress signaling and defense. The integration of structural biology, multi-omics, and synthetic biology continues to drive innovation in crop improvement, positioning lipid metabolism as a powerful lever for agricultural sustainability.

Objective 3: Develop crops with improved yield and/or functionality

This year’s collaborative research advanced the frontier of crop improvement by harnessing lipid metabolism to enhance yield, oil content, stress resilience, and value-added traits across a range of species. Through strategic metabolic engineering, gene discovery, and translational field studies, the project demonstrated how lipid pathways can be leveraged to meet the growing demands for food, fuel, and climate-resilient agriculture. A central theme was the enhancement of oil accumulation and composition in both seeds and vegetative tissues.

The Bates lab demonstrated that recently discovered triacylglycerol remodeling enzymes from Physaria fendleri could be utilized to enhance both total oil accumulation, and accumulation of valuable hydroxylated fatty acids in Arabidopsis thaliana and Camelina sativa.

The Mukherjee, Durrett, and Allen labs described the changes in soybean seed size as a consequence of SDP1 suppression, which have now been verified in multiple generations. Importantly, observed increases in seed size do not negatively impact seed number; investigations as to the underlying mechanism are ongoing. Ongoing collaborative work will generate and characterize stable mutant lines that investigate carbon partitioning and lipid turnover in developing soybean seeds.

The Allen lab in collaboration with Koo and Thelen reported the beta oxidation of fatty acids including some that were lipid-derived in a recent report (Koley et al., 2025). The study indicated that in multiple seed and a high-oil leaf system, that fat fatty acid oxidation occurs across development and appears to be, in collaboration with Koo and Thelen, reported the beta oxidation of fatty acids, including some that were lipid-derived, a regular part of metabolism.

The Koo lab collaborated with the Welti and Allen labs to boost plant biomass oil production using pathway engineering. The inducible expression of a plastid-localized phospholipase A1 (PLA1) called DEFECTIVE AND ANTHER DEHISCENCE 1 (DAD1) has led to the accumulation of TAGs in leaves, forming lipid droplets (LDs) (Kimberlin et al., 2025). Lipidomics and FAME analyses revealed that the accumulated TAGs were primarily composed of 18:3 fatty acids coming from MGDG and DGDG. Similar results were observed when DAD1 or its Nicotiana benthamiana homolog, NbDAD1, were transiently expressed in N. benthamiana. A stable transgenic soybean line harboring the inducible PLA1 also successfully increased leaf TAGs. Building on this work, the Koo lab is developing genetic tools and new lines with greater oil content. 

In collaboration with the Allen and Lee labs, the Durrett lab metabolically engineered camelina and pennycress lines to accumulate almost pure levels of acetyl-TAG, an unusually structured lipid with useful properties (Alkotami et al., 2024). The trace amounts of endogenous TAG remaining were localized to the embryonic axis of the seed. Future work will investigate the source and role of this residual TAG.

The Kosma lab recently developed a variety of potato that has reduced sprouting without compromising overall tuber yield (Vulavala et al., 2024). This unexpected discovery came about through CRISPR-Cas9 gene editing targeting transcription factors that regulate specialized lipid metabolism (wound suberin) during tuber wound healing.

The Cahoon lab successfully engineered oilseed camelina to produce the high-value ketocarotenoid astaxanthin, a red pigment with exceptional antioxidant properties. Utilizing genes from the plant Adonis aestivalis, their strategy led to the nearly complete conversion of β-carotene to ketocarotenoids, primarily astaxanthin. Field trials demonstrated a maximal accumulation of approximately 135 μg/g seed weight, with astaxanthin making up about 47 μg/g. The research also showed that the extracted oil from these seeds had enhanced oxidative stability, making it useful for food applications like oleogels. This breakthrough provides a new, sustainable plant-based source for a commercially valuable pigment used in aquaculture and the food industry, reducing reliance on less efficient methods (Kim et al., 2025).

The Cahoon and Clemente labs successfully engineered high-yielding sorghum to accumulate energy-dense vegetable oil in its leaves and stems, a novel approach for producing renewable diesel and sustainable aviation fuel. Using a "push-pull-protect" genetic strategy, the engineered sorghum accumulated oil in its leaves and stems to amounts of up to 5.5% DW and 3.5% DW respectively under field conditions. These results represented a 78-fold increase in leaves and a 58-fold increase in stems compared to wild-type plants. Their work highlights the effectiveness of a lab-to-field pipeline for developing high-vegetative-oil biomass crops. This achievement demonstrates a powerful new approach to creating feedstocks for sustainable fuels, offering a promising solution to strengthen American energy independence (Park et al., 2025).

The Dhankher Lab cloned and characterized MGAT1 genes encoding monoacylglycerol acyltransferase from Camelina sativa. Overexpression of CsMGAT1 in Camelina under the control of seed-specific BcNA1 (napin) promoter from Brassica napus, resulted in a significant increase in seed yield and oil contents. Camelina transgenics exhibited a 33-57% increase in seed yield, 8–10% seed oil content, >20% oil yields per plant, and altered polyunsaturated fatty acid (PUFA) content, compared to their parental wild-type (WT) plants. Results from [¹⁴C]acetate labeling of Camelina developing embryos expressing CsMGAT in culture indicated increased rates of radiolabeled fatty acid incorporation into glycerolipids compared to WT embryos. These findings suggest that overexpression of MGAT positively impacts fatty acid flux into lipids, contributing to increased oil accumulation and potentially enhancing seed yield in Camelina.

Narayanan Lab investigated the expression of the genes regulating lipid metabolic changes contributing to heat tolerance in peanut genotypes. They conducted a comprehensive lipidome analysis of 52 peanut recombinant inbred lines (RILs) of F6 population derived from a cross between the heat-tolerant genotype ICGS76 and the heat-susceptible TamrunOL02, under optimum (29/20°C) and heat stress conditions (38/28°C). They found that the sequestration of unsaturated acyl chains from membrane lipids to triacylglycerols (TG) and sterol esters (SE) helps to reduce the unsaturation levels in membrane lipids which in turn helps to maintain optimal membrane fluidity and integrity under heat stress conditions. They further investigated the expression patterns of key genes involved in the identified lipid remodeling. The tested genes were diacylglycerol acyltransferases (DGAT1-2, DGAT3-3), fatty acid desaturase (FAD3-2), phospholipid:diacylglycerol acyltransferase (PDAT), acyl-coA:sterol acyltransferase (ASAT), phospholipid:sterol acyl transferase (PSAT) and heat inducible lipase (HIL1). Gene expression analysis revealed an upregulation of ASAT, PSAT, DGAT3-3 and PDAT, that uniquely regulate the acylation of sterols and TGs. This result confirmed the role of TGs and SEs in heat stress tolerance through acyl sequestration. FAD3-2 which converts 18:2 fatty acids to 18:3, showed decreased expression, potentially contributing to reduced fatty acid unsaturation levels in membrane lipids by lowering the 18:3 fatty-acid amount under heat stress. The identified genes (ASAT, PSAT, DGAT3-3, PDAT, and FAD3-2) and associated lipid-related mechanisms of heat-stress tolerance will help develop heat-tolerant peanut varieties. Further, the genes will help develop molecular markers associated with heat tolerance that will speed up the peanut breeding programs for heat tolerance.

These integrated efforts underscore the potential of lipid-based strategies to improve crop performance, resilience, and market value across diverse agricultural systems.

Milestones:

• Developed method for rapid characterization of whole seed fatty acid composition and quantity for a variety of current and emerging oilseed crops (Garneau et al. 2025).
• Developed a new MSI technique to visualize carbon-carbon double bonds of lipids (Rensner et al., 2025).
• Demonstrated the use of unsupervised machine learning for the data analysis of MSI with in vivo isotope labeling (Johnson et al., 2025).
• Identified DIACYLGLYCEROL ACYLTRANSFERASE 1 regulatory elements essential for expression in pollen, and key to ultimate replacement of endogenous acyltransferase with those leading to higher yields or altered fatty acid compositions (McGuire et al. 2025).
• Engineered a new plant-based platform using the oilseed camelina to produce the high-value pigment astaxanthin for use in food and aquaculture (Kim et al., 2025).
• Identified metabolic adaptations to mutations and genetic engineering that lipid total oil accumulation in seeds and leaves (Kataya et al., 2025; Murphy et. al., 2025; Johnson et. al., 2025; Kimberlin et al., 2025). This knowledge will be the basis for new hypothesis driven approaches to enhance lipid accumulation in different plant tissues.
• Demonstrated fatty acid catabolism is active throughout development in filling seed and transgenic high-oil leaves (Koley et al., 2025).
• Developed a new pathway to create high-yielding sorghum capable of producing large amounts of vegetable oil in its leaves and stems for renewable diesel and sustainable aviation fuel (Park et al., 2025).
• Identified a lysophospholipid ligand for a transcriptional regulator that plays a role in epidermal development and response to phosphate limitation (Wojciechowska et al., 2024).
• Characterized the nuclear localization mechanism of a lipid-binding transcription factor implicated in regulation of seed oil levels (Ahmad et al., 2024)
• Developed a new variety of potato with reduced sprouting during cold storage that stands to significantly reduce postharvest potato tuber yield losses (Vulavala et al., 2024).
• Identified that membrane fluidity changes in response to temperature gated rice colonization by Magnaporthe oryzae (Richter et al. 2024).
• Identified lipid traits key to improving maize cold tolerance (Ojeda-Rivera et al. 2025).
• Narayanan spent three months of her sabbatical (March-May 2025) in the Allen Lab and established a new collaborative program that focuses on carbon metabolism for enhancing the quality of oilseed crops under optimal and heat-stress conditions.

1. Ahmad, B., Lerma-Reyes, R., Mukherjee, T., Nguyen, H.V., Weber, A.L., Cummings, E.E., Schulze, W.X., Comer, J.R., Schrick, K. 2024. Nuclear localization of HD-Zip transcription factor GLABRA2 is driven by Importin α. J Exp Bot. 75(20):6441-6461. doi:10.1093/jxb/erae326.
2. Garneau MG, Parchuri P, Zander N, Bates PD (2025) Rapid quantification of whole seed fatty acid amount, composition, and shape phenotypes from diverse oilseed species with large differences in seed size. Plant Methods 21: 67
3. McGuire ST, Shockey J, Bates PD (2025) The first intron and promoter of Arabidopsis DIACYLGLYCEROL ACYLTRANSFERASE 1 exert synergistic effects on pollen and embryo lipid accumulation. New Phytologist 245: 263-281
4. Kataya A, Nascimento JRS, Xu C, Garneau MG, Koley S, Kimberlin A, Mukherjee T, Mooney BP, Xu D, Bates PD, Allen DK, Koo AJ, Thelen JJ (2025) Comparative Omics Reveals Unanticipated Metabolic Rearrangements in a High-Oil Mutant of Plastid Acetyl-CoA Carboxylase. Journal of Proteome Research 24: 2675-2688
5. Kim, H.,Liu, L.,Han, L.,Park, K.,Kim, H. J.,Nguyen, T.,Nazarenus, T. J.,Cahoon, R. E.,Haslam, R. P.,Ciftci, O.,Napier, J. A. & Cahoon, E. B. (2025) Oilseed-based metabolic engineering of astaxanthin and related ketocarotenoids using a plant-derived pathway: Lab-to-field-to-application, Plant Biotechnology Journal. 23, 3451-3464 [https://doi.org/10.1111/pbi.70148].
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11. Dong, J., Croslow, S. W., Lane, S. T., Castro, D. C.,Blanford, J., Zhou, S.,Park, K., Burgess, S., Root, M., Cahoon, E. B., Shanklin, J., Sweedler, J. V., Zhao, H. & Hudson, M. E. (2025) Enhancing lipid production in plant cells through automated high-throughput genome engineering and phenotyping, The Plant Cell. 37 [10.1093/plcell/koaf026].
12. Murphy KM, Johnson BS, Harmon C, Gutierrez J, Sheng H, Kenney S, Gutierrez-Ortega K, Wickramanayake J, Fischer A, Brown A, Czymmek KJ, Bates PD, Allen DK, Gehan MA (2025) Excessive leaf oil modulates the plant abiotic stress response via reduced stomatal aperture in tobacco (Nicotiana tabacum). The Plant Journal 121: e70067
13. Gautam, B., Kim, H., Wang, C., Park, K., Cahoon, E. B. & Sedbrook, J. C. (2025) Meeting Liquid Biofuel and Bioproduct Goals: Biotechnological Design of the Intermediate Oilseeds Pennycress and Camelina, and Beyond, Journal of Experimental Botany [10.1093/jxb/eraf415].
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16. Johnson BS, Allen DK, Bates PD (2025) Triacylglycerol stability limits futile cycles and inhibition of carbon capture in oil-accumulating leaves. Plant Physiology 197: kiae121. doi: 10.1093/plphys/kiae121 
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22. Yang B, Fan R, Yao S, Lou H, Li J, Guo L, Wang X. Non-specific phospholipase Cs and their potential for crop improvement. J Exp Bot. 2025; eraf334, https://doi.org/10.1093/jxb/eraf334

23. Kosma DK, Graça J, Molina I (2024) Update on the structure and regulated biosynthesis of the apoplastic polymers cutin and suberin. Plant Physiology 197: kiae653 https://doi.org/10.1093/plphys/kiae653
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25. Ojeda-Rivera JO, Barnes AC, Ainsworth EA, Angelovici R, Basso B, Brindisi LJ, Brooks MD, Busch W, Buttelmann GL, Castellano MJ, Chen J, Costich DE, de Leon N, Emmett BD, Ertl D, Fitzsimmons SL, Flint-Garcia SA, Gore MA, Guan K, Hale CO, Herr S, Hirsch CN, Holding DH, Holland JB, Hsu S-K, Hua J, Hufford MB, Kaeppler SM, Leary EN, Liu Z-Y, Mahama AA, McCubbin TJ, Messina CD, Michael TP, Miller SJ, Murray SC, Okumoto S, Oren E, Park AN, Piñeros MA, Pugh NA, Raboy V, Rellán-Álvarez R, Romay MC, Rooney T, Roston RL, Sawers RJH, Schnable JC, Schulz AJ, Scott MP, Springer NM, Washburn JD, Zambrano MA, Zhai J, Zou J, Buckler ES. 2025. Designing a nitrogen-efficient cold-tolerant maize for modern agricultural systems, The Plant Cell, Volume 37, Issue 7, koaf139, https://doi.org/10.1093/plcell/koaf139
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