INFORM October 2025

inform October 2025 Volume 36 (9)

OMEGA-6 to OMEGA-3 via GENE THERAPY

ALSO INSIDE: Designing tastier soy varieties

Cover crops engineered to make industrial lipids An olive oil ingredient that improves gut health

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One dose of Fat-1 gene therapy eases osteoarthritis in mice This article describes research showing a single dose of fat-1 gene therapy eased osteoarthritis symptoms in mice by converting omega-6 fatty acids into anti inflammatory omega-3s. The treatment reduced joint damage, inflammation, and signs of premature cellular aging tied to obesity. While promising, the therapy has only been tested in animals, and more work is needed before human trials can begin. Removing off-flavors from soy Scientists are developing new soybean varieties designed to eliminate the “beany” and “painty” off-flavors that turn consumers away from soy-based foods. By reducing polyunsaturated fatty acids and targeting key enzymes, they created high-oleic, flavor-null soybeans that retain nutrition without the unpleasant taste. Read about how these advances could make soy protein more appealing for use in snacks, beverages, and plant-based alternatives.

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Metabolic engineering enables lipid redesign Researchers have engineered cover crops like camelina and pennycress to produce acetyl-triacylglycerols (acetyl-TAGs), specialty plant oils with low viscosity and improved cold properties ideal for industrial uses. By optimizing enzymes, reducing competing pathways, and increasing precursor supply, they achieved nearly pure acetyl-TAG levels in seeds. The article describes how the cover crops were engineered as sustainable sources for biofuels, lubricants, and other biobased products. Scientists study a valuable ingredient in olive oil The compound 3,3-dimethyl-1-butanol (DMB) works in the human gut to inhibit some inflammatory diseases by preventing the formation of the trimethylamine enzyme. In animal models, this enzyme creates a compound in the liver that seems to facilitate a build up of fat and cholesterol in the arteries. An international research team developed a method that confirms DMB’s presence in extra virgin olive oil and reliably measures the amount. The next step is to better understand the contribution DMB makes to improving human health.

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CONTENTS

4 Index to Advertisers

5 Editor’s Letter 6 Division Update

26 Regulatory Review 28 Extracts & Distillates

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Julian Barnes Etienne Guillocheau Jerry King

Gary List Thais L. T. da Silva Raj Shah

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EDITOR’S LETTER

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An array of biotechnology applications

Innovation in biotechnology often starts with reimagining the familiar. This issue highlights three examples of how scientists are reshaping what we know about health, food, and sustain ability—by starting at the molecular level. In our cover story, researchers explore how a single dose of fat-1 gene therapy in mice transformed omega-6 fats into omega-3s leading to eased osteoarthritis symptoms and slowed signs of premature cellular aging. Another feature looks at soybeans—long a cornerstone of plant-based protein—and the effort to breed “flavor-null” varieties that eliminate the painty, beany notes that turn consumers away, opening the door to more appealing plant-based foods. Finally, we exam ine how engineered cover crops like camelina and pennycress are being redesigned to produce novel plant oils with industrial

potential, from lubricants to biofuels, offering a sustainable new platform for renewable materials. Together, these stories reveal how precise genetic and metabolic innovations are unlocking fresh possibilities across medicine, nutrition, and industry. They also remind us that the solutions to some of today’s most pressing challenges may lie in rethinking the building blocks of biology itself.

Yours in science,

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Biotechnology Division

2025 DIVISION ACHIEVEMENT AWARD WINNER Xavier Malcata is a professor at University of Porto, in Porto, Portugal and a senior researcher at LEPABE, a faculty of engi neering research unit operating in the fields of chemical, envi ronmental and biological engineering at the university. INFORM: Can you share your journey into the field of bio technology? What inspired you to pursue this path? Malcata: I was motivated from the very beginning by the pros pect of working in biotechnology since it offers an array of engineering opportunities. My journey started with a bach elor’s degree in chemical engineering from the University of Porto, followed by a chemical engineering doctorate with a minor in food science, biochemistry, and statistics from the University of Wisconsin, Madison. The research program for my dissertation covered reac tors with immobilized lipases aimed at tailoring edible fats. I eventually applied my background on modeling from first prin ciples to optimize biochemical processes in vitro . From then on, I have focused on various fields of biotechnology, from characterization and improvement of Portuguese traditional foods, through identification and study of adventitious probi otic cultures from non-dairy sources, to design and optimiza tion of photobioreactors for operations with microalgae. INFORM: Your work has had a substantial impact on the field of biotechnology. Can you discuss one or two key projects that you believe have made the most difference? Malcata: My studies of lipase immobilization on multiple sup ports helped pave the way for current industrial applications of immobilized lipases to engineer existing fats via inter- or trans-esterification with target fatty acids bearing a stronger nutritional role or a more pleasant sensory feature. Lipases were by that time seen as a solution in search of a problem. Their unique activation at the interface between aqueous media and hydrophobic materials (plastic polymers or insol uble lipids) had hampered deeper studies. The difficulties stemmed from simultaneously handling the interfacial chem ical reaction and the mass transfer between the biphasic sys tems that nature designed. Later, I delved into the possibility of using lipases for reactive distillation to produce volatile esters with sensory notes and combining separations with in situ enzyme-catalyzed reactions. INFORM: What are some of the biggest challenges you have faced in your research, and how did you overcome them? Malcata: The biggest challenges faced during my research were probably similar to those faced by most researchers. The need to find sustained sources of funding to cover the

expenses incurred, as well as sophisticated analytical and pro cessing equipment; and to find graduate students available to pursue an academic career, motivated in advancing the state-of-the-art, autonomous in their performance, curious in their approach, thorough in their analysis, and resilient in their work. Research is a passionate game where one works at the edge of knowledge but never knows what will be found ahead. As Thomas Edison once said, “ I never failed in research; I just learned one thousand ways that do not work. ” INFORM: How has your research evolved over the years, and what emerging trends do you see in biotechnology that excite you? Malcata: I started from a classical background in chemical engineering supported by thermodynamics, chemical kinet ics, transport phenomena, and process control as structural pillars. As I became more proficient at handling and under standing viable microorganisms and their enzymes to bring about reactions with rates and selectivity at will, I realized that those same areas of knowledge applied at the microscale. But the phenomena take place simultaneously and are subjected to complex interactions that guarantee the best use of envi Xavier Malcata

YOUR AOCS COMMUNITY

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ronmental resources available in the medium. It was indeed a challenge to mechanistically model such microorganisms or their enzymes because the same laws of nature apply there that also apply to chemical processes at industrial scale. At present, I am considering specific yet integrated changes in the metabolic pathways of photosynthetic microorganisms to enhanced the production of commodities that may serve as cleaner fuels in the near future. INFORM: How do you believe this recognition will influ ence your future work in biotechnology? Malcata: I must confess that, whenever selected as a recipi ent of an award, especially if granted by a respected interna tional scientific or professional society, I feel an indescribable amount of pride to have my work recognized by peers world wide, and a unique sensation of accomplished duty. Over several decades, I have pursued a consistent and integrated research program to help advance the state-of-the-art in top ics motivated by my scientific curiosity, and the possibility of transferring that knowledge to undergraduate and graduate students. The track record I have built has also made a differ ence in my performance in the classroom. My lectures have plenty of “war stories,” coming from real life and actual expe rience—prone to emphasize practical applicability of advanced knowledge, and rational application of first principles to under stand and solve whatever situations and problems arise. INFORM: What role do you see AOCS playing in fostering collaboration among scientists and industry professionals in biotechnology? Malcata: AOCS addresses the interface between many dis tinct disciplines within the core field of fats and oils. It has established itself as an invaluable stakeholder, with comple mentary and unique expertise spread throughout its mem bership. Biotechnology is one of the best examples of how tools inspired by our surroundings (namely, biochemical pro cesses established in nature) can be used to our advantage to improve oils and fats, in both nutritional, functional, and envi

ronmental terms. In addition, both the fundamental issues of science and the applied issues of technology are included in the portfolio of activities AOCS offers—from topic-oriented workshops to the annual meeting with many thematic ses sions, from advanced books meant for professionals and the scientific community to a membership magazine covering daily issues of wide interest. Furthermore, its awards designed for public recognition of exemplary careers and outstanding per formance in academia, industry, or government contributes to the level of excellence it has consistently pursued. INFORM: What advice would you give to young sci entists and researchers aspiring to make an impact in biotechnology? Malcata: My advice to young scientists at the beginning of their careers is simple and straightforward, be tenacious with your approaches, but humble with your results. Research is a double-edged sword, in that it will eventually yield some use ful results—from either the understanding or the application points of view—but typically at the expense of many failures along the way. Therefore, a strong character is a must, be able to overcome misfortunes and poor outcomes while already looking for the next move and maintain an optimistic atti tude. On the other hand, research (especially in biotechnology) is rather expensive, and thus only possible at the expense of public money that might be used otherwise. This means that whatever successes are attained, they would have been hardly possible without the anonymous support of your fellow citi zens. They deserve consideration for their implicit input, well beyond the fame earned by the researcher himself. In fact, lasting impacts may come out of a fortunate breakthrough but are usually the outcome of long paths of consistent, resigned, incremental work, building on the results by peers and rely ing on multi-tasked teams holding complementary expertise. Finally, research cannot be blindly developed. It should ulti mately aim at improving the quality of life of mankind and the preservation of our common home called Earth—otherwise it will be meaningless and devoid of social value.

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One dose of

Despite affecting high numbers of American adults—more than 30 million according to the CDC— osteoarthritis (OA) has limited treatment options and no cure. OA is the most common form of arthritis, marked by discomfort, stiffness, and swelling in affected joints, frequently the knees, hips, hands, and spine. It is a painful, debilitating, progressive condition. If you are diagnosed with OA today, providers may recommend physical therapy, pain relieving medication, crutches, a cane, or joint replacement surgery, but they cannot offer a cure (https://tinyurl.com/492356). therapy eases osteoarthritis in mice Kelly Carroll Fat-1 gene

Farshid Guilak, professor of orthopedic surgery at Washington University in St. Louis, Missouri, is trying to change that. “It is one of the few diseases where we have absolutely nothing we can do to affect the disease process,” he said. Guilak and six colleagues recently published a study showing that gene therapy targeting a fatty acid desaturase successfully reduced the severity of OA in mice (https://doi.org/10.1073/pnas.2402954121). Gene therapy is a treatment that corrects a malfunctioning gene to prevent, or help the body resist, disease. Other research teams have developed sim ilar treatments for other diseases, including sickle cell, hemophilia A, and beta-thalassemia. The number of available therapies is growing rapidly. TRANSFORMING FAT Here researchers placed five-week-old mice on either a high-fat diet rich in omega-6 fatty acids or a regular diet. About four weeks later, the mice were given a single dose of fat-1 gene therapy, administered by an ade no-associated virus (AAV). Matlock Jeffries, director of the Oklahoma Medical Research Foundation’s Arthritis Research Center, said that this type of viral-me diated gene therapy does not change the host organism’s genome. It expresses the therapeutic gene over a long period, requiring infrequent administration. While AAV-administered gene therapies are not perma nent, they last a long time, Jeffries said, estimating that gene expression would gradually decrease after one to two years.

• Osteoarthritis is a common and painful condition of the joints with no available cure. • A single dose of fat-1 gene therapy systemically converts omega-6 fatty acids into omega-3 fatty acids and reduces the severity of injury- and age-related OA in experimental mouse models. • More experiments are needed to clarify how fat-1 gene therapy may translate to humans.

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The fat-1 gene therapy used in this study converts omega-6 fatty acids into omega-3 fatty acids based on serum analysis. The gene is not typically found in mammals like mice or humans, and scientists originally discovered it in small worms called Caenorhabditis elegans . PROFOUND EFFECTS Researchers induced OA in mice using an experimental surgical technique called destabilization of the medial meniscus (DMM). The procedure involves cutting a knee ligament which causes progressive cartilage breakdown, synovial membrane inflammation, subchondral bone

remodeling, and osteophyte formation in the weeks following surgery (https://doi.org/10.1038/s41598-018-21184-5). A single dose of fat-1 gene therapy reduced the amount of omega-6 fatty acids in the serum. It significantly reduced the mice’s weight and body fat at 28 weeks of age. Fat-1 gene therapy lowered fasting glucose levels and prostaglandin E2, an inflammatory lipid, in mice fed a high-fat diet. And the treatment reduced pro-inflammatory cytokines, including IL-6, IL-1α, and IL-1β in mice serum at 52 weeks of age. Fat-1 gene therapy also proved beneficial in two mouse models of OA: OA caused by DMM and OA caused by typi cal aging. Fat-1 gene therapy diminished the risk of cartilage

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ically, and that really improves overall metabolic health and joint health.” Philip Calder, professor of nutritional immunology at the University of Southampton, United Kingdom, said in an email that the study’s findings are consistent with previous research on the connections between dietary fatty acids and inflamma tion-related diseases. In 2015, Thiago Belchoir published research demonstrat ing that omega-3 fatty acids protect against diet-induced obesity and inflammation through the transcription fac tor PPARγ (https://doi.org/10.1002/mnfr.201400914). And in 2020, Guilak and colleagues showed that fat-1 transgenic mice exhibited reduced OA severity (https://doi.org/10.1186/ s13075-020-02170-7). FATTY ACIDS AND CELLULAR AGING Guilak’s current study also indicates that obesity causes pre mature joint aging. The research shows that high-fat diet-in duced obesity increases cellular senescence, a cellular state associated with aging where cells are alive but no longer divid ing. Higher amounts of senescence markers—β-galactosidase, p21, and p16—were measured in the fat tissues and joints of obese mice with DMM-induced OA. These findings suggest that high-fat diets may lead to premature cellular aging and Some omega-6 fatty acids are common elements of the human diet. Linoleic acid (LA), found in grapeseed, cotton seed, sunflower, corn, and canola oils, as well as nuts, seeds, mixed grains, and meats, is the most prevalent omega-6 fatty acid in the diet. When metabolized, LA breaks down into other fatty acids, including arachidonic acid (ARA). ARA is also found in meat, eggs, and other foods and is a critical component of cell membranes. It also contributes to brain development, and wound healing (https://doi.org/10.1016/j.jare.2017.11.004). If there is too little LA in the diet, the body’s skin and epi thelial barriers can become compromised. There is also evi dence that higher consumption of LA improves heart and metabolic health. Research has shown that higher levels of LA in the blood are tied to lower risks of coronary heart disease, stroke, and type 2 diabetes. It remains unclear how LA is medi ating these effects. In a 2024 review paper, Kristina H. Jackson and colleagues wrote: “Taken together, these epidemiological findings strongly suggest that higher LA (and in some settings, ARA) levels are linked with improved outcomes” (https://doi. org/10.1186/s12944-024-02246-2). The fatty acid controversy Both omega-6 and omega-3 fatty acids are essential to health. Despite the excess of omega-6 in the typical Western diet and their association with obesity and systemic inflammation, no single element of the human diet, is a villain. Furthermore, not all omega-6 fatty acids behave the same way inside the body.

degeneration and synovitis, inflammation of the synovial mem brane that lines the joint. The researchers analyzed the ther apeutic effects on the trabeculae, spongy bone structures in joints that distribute load and absorb shock. Mice with DMM induced OA had fewer but thicker trabeculae with greater sep aration. Fat-1 gene therapy counteracted these effects in bone microarchitecture. CONSISTENT OUTCOMES Through this series of experiments, Guilak and his colleagues were able to use gene therapy to alter the systemic fatty acid composition and inflammatory profile in obese mice, which reduced the severity of OA. “They found that injecting into the whole mouse this virus that contains this gene ... did result in improvements in OA outcomes,” said Jefferies. “The histology was better, and it does appear, probably, that the inflammation around the joint was better. And also, interestingly enough, it looks like the mice weighed less even when they were fed a high-fat diet.” Guilak emphasized that the study indicates fat-1 con verts omega-6 fatty acids into even more beneficial ome ga-3s, thereby improving metabolism and joint health. He said through the expression of fat-1 , you can “take your hamburger and fries and convert it to a salmon oil in your body automat

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contribute to poor joint health. “One of our problems is, when we have really bad diets, we age our joints and other parts of our body faster,” Guilak said. Fat-1 gene therapy successfully mitigated cellular aging. Mice fed a high-fat diet and given fat-1 gene therapy showed reduced levels of cellular senescence markers. Converting omega-6 fatty acids to omega-3 fatty acids in mice via gene therapy, slowed down that accelerated aging process, Guilak said. Obesity contributes to OA progression and is a signifi cant, preventable risk factor. However, for many years people believed that the connection between obesity and OA was due to weight-related increased loading and eventual breakdown of the joints. Guilak said that people dismissed OA develop ment in obese people as wear and tear on overloaded joints. Researchers have revealed a more complex relationship, how ever. Systemic, persistent inflammation has a significant role in OA development. “It turns out a major component is the inflammatory and metabolic environment that occurs with obesity,” said Guilak. “They have this low-grade, chronic, systemic inflammation.” Circulating pro-inflammatory mediators, such as cytokines IL-1β, TNF-α, and IL-6, impact OA (https://tinyurl.com/923781). Guilak said fat produces many of these pro-inflammatory many researchers to conclude that the omega-6/omega-3 ratio does not convey the nuanced role of individual fatty acids. The ratio may also be unhelpful when determin ing the optimal diet of fatty acids. Some researchers pre fer to use an omega-3 index, which is the sum of two omega-3 fatty acids: EPA and DHA. Evidence suggests that the omega-3 index is a helpful tool in predicting a person’s risk of coronary heart disease and sudden cardiac death (https://doi.org/10.1016/j.atherosclerosis.2017.05.007). Despite advances in understanding fatty acid metab olism and the role of dietary components in inflammation and disease, many aspects remain unknown. Future studies will continue to parse out the roles of individual fatty acids in the body and determine the optimal levels for health. The effects of ARA are complex. A systematic review in 2019, found that higher levels of ARA were not associ ated with better or worse health outcomes. People who consumed more than the typical amount of ARA (100-200 mg per day) up to between 1,000 and 1,500 mg per day did not experience adverse effects on their blood lipids, immune system function, or inflammation. Interestingly, a small study of elderly Japanese men indicated that ARA may counteract age-related cognitive decline (https://doi. org/10.1017/S0007114519000692). The complex effects of omega-6 fatty acids has led

cytokines, which circulate throughout the body, including the joints, and contribute to chronic conditions like OA over time. “It could take 20 years to develop osteoarthritis,” he said. Jeffries emphasized that both aging and obesity are gener ally associated with increased systemic inflammation. “As we age, our immune system tends to become more inflammatory all over the body, and obesity induces increased inflammation all over the body, and both of those things result in increased rates of OA.” Obesity and systemic inflammation play a role in the development of many diseases beyond OA. People who are obese have higher rates of hypertension, type 2 dia betes, heart disease, and some cancers (https://tinyurl. com/5085085). “It is hard to find a disease that is not impacted by metabolic function and obesity, and as we dig down further, dietary composition,” Guilak said. THE FUTURE OF FAT-1 GENE THERAPY Obesity remains a significant challenge in America. In 2023, more than 20 percent of adults in the US were obese, with the highest rates in the Midwest and South. In 23 states, more than one in three adults was obese (https://tinyurl. com/9154189). Available treatments for obesity include diet modifi cations, exercise, bariatric surgery, and GLP-1 agonists like Semaglutide. Each treatment option comes with challenges. Lifestyle modifications can be difficult to maintain over the long term, while medications can have side effects, and sur gery is invasive and costly. There may eventually be a gene therapy available to treat obesity. Researchers have only studied fat-1 gene therapy in mice and pigs, but Guilak sees a future where this type of gene ther apy may be available to treat people. Gene therapies come with risks and often significant monetary costs. “We would think about it only in really extreme cases where diet and exercise are not appropriate or would not work,” he said. An example population would include patients who are obese, immobile, and at extreme health risk. Guilak also sees poten tial use for fat-1 gene therapy in obese pets with obesity-re lated health challenges. Future studies will detemine how fat-1 gene therapy may translate from mouse models to humans. Guilak’s team hopes to uncover how to regulate fat-1 gene therapy bet ter and address potential safety concerns. Notably, the mice that received fat-1 gene therapy in Guilak’s studies showed no adverse side effects. “In everything we measured, we only saw beneficial effects,” he said. “We are a long way away from get ting into human studies.” Kelly Carroll is an assistant professor at Bellarmine University and freelance writer and editor. She can be contacted at kcarroll@bellarmine.edu.

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Processing soybeans to make soy-based products can set off lipid oxidation reactions, giving the foods off-flavors. Source: iStock

Removing off-flavors from soy Katie Cottingham

It is not every day that researchers want to develop a fla vorless, bland food, but that has been the goal of Kristin Bilyeu and her colleagues for the past several years. “We wanted to develop a bland protein source using our knowledge of genetics and biochemistry to reduce off-flavors for soy in food,” says Bilyeu. She is a research molecular biologist at the US Department of Agriculture Agricultural Research Service (USDA-ARS) and adjunct professor at the University of Missouri in Columbia. Soy is high in protein, an important component of the body including mus cle, skin, and bone. But during the storage of products like soy milk and soy burgers PUFA in residual oil in the protein meal can oxidize, leading to a bad taste and smell. “Painty” and “beany” are just a couple of the negative terms that trained tasters on sensory panels use to describe these unpleasant char acteristics. These objectionable flavors deter many consumers from trying soy based products. So, Bilyeu teamed up with Bongkosh Vardhanabhuti, an associate pro fessor of food science, and Andrew Scaboo, an assistant professor of plant science and technology, both at the University of Missouri. They set out to develop brand-new varieties of soybeans that provide the health benefits without the funk.

• Lipid oxidation reactions produce off-flavors, making soy products taste “painty” or “beany.” • Researchers are developing new soybean varieties that have higher oleic acid and lower levels of polyunsaturated fatty acid that are more easily oxidized. • The work led to high oleic acid soybeans that could become snack food ingredients.

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PUFAS: GOOD FOR HEALTH, BAD FOR SOY PROCESSING

of enzymes that catalyze PUFA oxidation. Heat treatment can reduce lipoxygenase activity and produce compounds that can help mask beany or painty compounds. Bathing soybeans in an alkaline solution can also lessen the impact of these enzymes. Bilyeu and her team, however, went right to the source— the genes encoding lipoxygenases, as well as genes encoding fatty acid desaturases that can convert other fatty acids into PUFAs. “My work has been to find non-functional versions of each of those enzymes to try to stop compounds from going through the oxidation pathway,” says Bilyeu. Together with Scaboo, she developed new soybean varieties that have differ ent defects in these two classes of enzymes. “We have been so fortunate to have the USDA-ARS Germplasm Resources Information Network (GRIN, https:// www.ars-grin.gov/), which is a seed bank that provides researchers with over 20,000 different accessions of soybean from all over the world,” says Bilyeu. “It is a treasure.” The team screened GRIN’s naturally diverse collection by pheno type and genotype to discover the best varieties for their work. They also used chemical mutagens to make changes to soy bean plants that they screened for appropriate phenotypes. Because they are not directly changing the genes with molec ular biology techniques, the varieties are not considered to be genetically modified organisms, or GMOs. The first good candidate soybean variety (referred to as HOLL) had high levels of oleic acid and low levels of linolenic acid (the PUFA linked to off-flavors in soybeans). They patented and trademarked this variety as Soyleic® (https://soyleic.com/). Its oil is about 80 percent oleic acid and about 2.5 percent lin olenic acid. The team also developed HOLL varieties with other beneficial attributes, such less raffinose, a carbohydrate with out nutritional value, to make the soybean protein meal more

Soy is an unusual seed oil crop. Unlike canola or sunflower seeds that contain up to 50 percent oil, soybean seeds are only about 18-22 percent oil but about 40 percent protein ( https:// doi.org/10.1186/s12870-019-2199-7, https://doi.org/10.1007/ s00122-022-04222-9). The protein in soybeans is complete, meaning it contains all of the essential amino acids. This is important for consumers who are vegetarians or vegans and avoid eating meat. To unlock the oil and protein, soybean seeds are broken open by mechanical grinding. But that exposes the oil to lipid oxidation reactions since its composition is more than 60 per cent polyunsaturated fatty acids (PUFAs) and 25 percent oleic acid, which is monounsaturated. Consuming PUFAs can reduce the risk of heart disease and is recommended for a healthy diet. However, PUFAs are more susceptible to lipid oxidation. Oxidation reduces shelf life and produces unpleas ant off-flavors and smells from compounds such as alde hydes, ketones, and alcohols (https://doi.org/10.3390/ foods12050923). “The flavor of the oil will change; however, lipid oxidation will also affect the flavor perception of the pro teins,” says Vardhanabhuti. That is because a little bit of the oil carries through some of the processing steps along with the protein as it is modified from flakes to soy protein concentrate and soy protein isolate, making them taste bad as well, she explains. FLAVOR-NULL, NOT FLAVORFUL Oxygen in the air and enzymes in the plant can oxidize PUFAs, resulting in undesirable compounds. Industry has a few tools at its disposal to dampen the effects of lipoxygenases, a family

Soybeans called Super L are high in oleic acid and low in linolenic acid, the fatty acid most likely to be oxidized in a lipid oxidation reaction. Source: Kristin Bilyeu

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Developing flavor-null soybeans requires meticulous genotyping analyses, and this screen shows data from plants that are homozygous mutant (red), homozygous wild-type (blue), or heterozygous (green) for a seed composition trait. Source: Kristin Bilyeu

attractive as an animal feed. Additional experiments eventu ally led the team to the flavor-null variety called Super that combines the HOLL/Soyleic® trait and defective lipoxygenase enzymes with the carbohydrate trait. “The idea was to see what I can put all in one package to bring more value through out the value chain,” says Bilyeu. On the food science side, Vardhanabhuti’s team con ducted sensory panels that tasted samples of the new soybean varieties. To tie the sensory data to the chemis try, Vardhanabhuti collaborated with another group at the University of Missouri that analyzed the new varieties with gas chromatography-mass spectrometry. This method pro vides information on volatile compounds that usually have a smell associated with them. The researchers analyzed raw soy slurries and soy milk they made in the lab. “Overall, the Super soybean had the lowest concentration of vola tiles that have been shown to correlate with off-flavor,” says Vardhanabhuti. Bilyeu wondered if she could go even lower and reduce linolenic acid level to less than 1 percent of the oil. Scaboo already had developed a soybean variety with a suite of dys functional enzyme genes that they then bred with Super, resulting in a variety with 10 defective enzyme genes. This Super-L variety has ultra-low linolenic acid—less than 1 per cent. “The little bit of linolenic acid is due to a leak-through of the biochemical pathway, so even though I have done my best to destroy all parts of it, there is still a little bit of the biochem istry capability left,” says Bilyeu. Vardhanabhuti’s team will test Super-L soon for its sensory and chemical properties and should have results by summer 2026. GETTING FARMERS AND INDUSTRY ONBOARD The soybean industry is a large-scale production that is difficult to change, so the researchers are working to fit their needs, as well as those of farmers. “Everybody in the value chain has to perceive their role as being a winner,” says Bilyeu.

“On the research end, we need to make sure that these soybean seeds behave like any other soybean seeds that a farmer would grow so that they are incentivized to grow them and know that they will be as profitable as their other seeds,” says Bilyeu. To help farmers feel they are not los ing anything by growing the new soybeans, researchers are conducting tests to make sure good yields are possible for different geographical locations and environments. Another aspect is helping farmers overcome negative perceptions of how growing alternative protein sources like soybeans for potential use in non-animal meat products could impact their communities. Companies need to see that the varieties can be grown on a large scale and that the protein would be useful for many types of products. “They need a source of nutritious protein that has the functional properties that work in their products, and the less offending the flavor is, the better,” says Bilyeu. For example, Vardhanabhuti says that if the protein will be used in ice cream, it should be able to foam, but foaming is not desirable in many types of beverages. The team’s efforts combining the HOLL trait with other properties, such as low raffinose content, can also make the soybeans more valuable for industry. Currently, the team is in discussions with companies to see how they can work together. A food company has expressed interest in including the new soybean varieties into snacks. Bilyeu notes that alter native protein ingredient companies and seed companies could also be interested in collaborating on these soy varieties. “I am really excited to be able to talk to these different entities and let them know what kind of research we have here and the fact that the soybean is available under different agreements,” she says. “We want to be good partners at ARS.” Katie Cottingham is a freelance science writer and editor whose work has appeared in publications, such as Science , Scientific American , and Smithsonian Magazine . She can be contacted at katie.cottingham@yahoo.com.

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16 • inform October 2025, Vol. 36 (9)

In our research group, when we think about plant oils, we think beyond the conventional. Across the plant kingdom, hundreds of unique triacylglycerol (TAG) structures, the main components of oils, are produced by different plant species. Some of these TAGs contain unusual fatty acids (FA) with atypical chain lengths, functional groups, or double bonds in unconventional positions. The difference in structures gives plant oils distinct physical and chemical properties which make them useful in many industries. However, a lot of species that naturally produce them are not suitable for large-scale production. This raises a key question for our work: How can we produce these valuable, structurally diverse oils in crops with favorable agronomic traits? Metabolic engineering enables lipid redesign Linah Alkotami and Timothy Durrett

Genetically engineered oilseed crops could serve as alternative plat forms to make these oils more accessible. Yet, researchers report having difficulty achieving native-like oil levels in engineered crops. Recent advances in systems biology and molecular techniques give us a better understanding of the complex pathways involved in the biosynthesis of diverse oils. Our lab recently demonstrated a prom ising result in engineering specialized oils ( https://doi.org/10.1073/ pnas.2412542121 ). Through pathway optimization, we enabled the pro O O sn-1 sn-1 Euonymus alatus

• Acetyl-TAGs have low viscosity and better cold properties, making them suitable for industrial applications. • Native plants that produce acetyl TAGs lack traits needed for large-scale cultivation. • Selective enzymes reduced competition from endogenous pathways and increased substrate availability resulting in almost pure levels of acetyl-TAG in transgenic camelina and pennycress seeds.

O

sn-2

O

sn-2

sn-3

sn-3

O

O O

O

O

O

O

O

Euonymus fortunei

Celastrus scandens

TAG Acetyl-TAG An image of seeds from species such as Euonymus alatus and E. fortunei , which naturally accumulate acetyl-TAGs at high levels. Structures of TAG and acetyl-TAG with the acetyl group indicated in red. Unlike conventional TAGs, which contain three long-chain fatty acids, acetyl-TAGs feature an acetate group at the sn-3 position, lowering viscosity and improving cold flow properties. Source: Durrett Lab.

Acetyl-CoA pool

Acetyl-TAG

BIOENGINEERING

inform October 2025, Vol. 36 (9) • 17

Euonymus alatus

sn-1

sn-1

O

O

O

sn-2

O

sn-2

sn-3

sn-3

O

O O

O

O

O

O

O

Euonymus fortunei

Celastrus scandens

duction of acetyl-triacylglycerol (acetyl-TAG), a unique TAG structure, in oilseed cover crops at levels comparable to those in native species. ACETYL-TAGS: WHY THEY MATTER Acetyl-TAGs differ from conventional TAGs in that the sn-3 position on the glycerol backbone contains an acetate group rather than a third long-chain fatty acid. As a result, ace tyl-TAGs have lower viscosity and melting points, making them suitable for use in lubricants, plasticizers, and drop-in biofuels. Certain native species, such as members of the Euonymus or Celastrus genera, predominantly produce acetyl-TAG in seeds, although trace amounts have been observed in the fruit and arils. The abundance of acetyl-TAG in seeds varies among native species, with some levels reaching 98 percent. Although promising, these species suffer from agronomic challenges such as a long growth cycle, non-uniform maturation of seeds, and the need for manual harvesting due to incompatibility with harvesting machinery. To engineer high levels of acetyl-TAGs, we needed crop species that could combine agronomic value, transformation ease, and scalability. COVER CROPS AS BIOTECHNOLOGY PLATFORMS Our lab focuses on two oilseeds, camelina ( Camelina sativa ) and pennycress ( Thlaspi arvense ). Both belong to the mus tard family (Brassicaceae), similar to rapeseed, canola, and the model plant Arabidopsis. Their close evolutionary relation ship to Arabidopsis allows us to draw on the extensive knowl

TAG edge of lipid metabolism and oil biosynthesis available for this well-studied model species. Because the genomes of both camelina and pennycress are sequenced and publicly available, we can directly apply insights from Arabidopsis to identify and manipulate key genes involved in oil production. In addition, both plants are prime candidates for biotechnology because Acetyl-TAG

Acetyl-CoA pool

Acetyl-TAG

1

EfDAcT

FAE1 18:1-CoA FAE1 20:1-CoA 22:1-CoA FA elongation 3

DAG

Acyl-CoA pool

DGAT1

2

RNAi

TAG

WT Simplified pathway of the last step of TAG and acetyl-TAG biosynthesis. Numbers represent the metabolic engineering strategies used in this study to optimize acetyl-TAG production: (1) Expression of high-activity EfDAcT enzyme; (2) suppression of endogenous DGAT1 to reduce competing TAG synthesis; (3) elimination of fatty acid elongation to increase acetyl-CoA supply. Source: Durrett Lab EfDAcT 1 EfDAcT DGAT1-RNAi 2 EfDAcT DGAT1-RNAi fae1 background 3

Pennycress

0%

68%

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98%

18 • inform October 2025, Vol. 36 (9)

Genes of intrest

Selectable marker

Harvested seeds screened for selectable markers

T-DNA

Dipped plants covered overnight to maintain high humidity

Flowers dipped in Agrobacterium suspension

Transgenic seeds (DsRed+) under green light and red lense filter

Vir genes

Agrobacterium Tumefaciens

Wildtype seeds under green light and red lense filter

Vacuum applied to enhance infiltration of Agrobacterium

Plants transferred to growth chambers for recovery and seed maturation

Camelina at early flowering stage

Agrobacterium suspension

Agrobacterium -mediated transformation of Camelina and Pennycress Most plant species are difficult to genetically transform, often requiring laborious tissue culture to regenerate plants from transformed cells. An important exception is the model plant Arabidopsis thaliana , which can be effi ciently transformed through Agrobacterium -mediated flo ral dip, bypassing tissue culture entirely. This approach has been extended to two emerging oilseed crops, Camelina sativa and Thlaspi arvense (field pennycress), making them useful platforms for plant biotechnology. Agrobacterium tumefaciens is a soil bacterium that developing flowers in an Agrobacterium suspension con taining a surfactant to reduce surface tension. A key part of the protocol involves using vacuum infiltration to draw the bacterial suspension deeper into floral tissues, increasing access to target cells. Following infiltration, plants are maintained under high humidity to support infection. Seeds are harvested from treated plants, and transgenic progeny are identified via reporter genes such as DsRed or selectable markers such as herbicide resistance.

they can easily be genetically modified using Agrobacterium mediated floral dip transformation with vacuum infiltration (see sidebar). Agronomically, both crops fit well into existing agricultural systems. They grow in the off-season between major food crops, tolerate diverse environmental conditions, and require minimal water and fertilizer. Compared to other oilseed crops such as soybean, they have short growing cycles and a relatively high oil yield. As cover crops, they naturally transfers a segment of its plasmid DNA (T-DNA) into plant cells during infection. This DNA integrates into the plant genome, enabling stable expression of intro duced genes. Researchers have taken advantage of this ability, replacing the bacterium’s tumor-inducing genes with desired transgenes, thus harnessing Agrobacterium ’s natural DNA delivery machinery for crop improvement. For camelina and pennycress, the method involves growing plants until early flowering, then immersing

suppress weeds, reduce erosion, and retain nutrients, contributing to soil health. BUILDING THE PATHWAY Our work involved multiple metabolic engineering strategies, with the goal of matching the acetyl-TAG levels found in some Euonymus seeds—over 90 mol percent. To reach these levels, we first needed to build the acetyl-TAG biosynthetic pathway in camelina and pennycress. Using this method, transformation efficiencies reach about one percent in camelina and half a percent in pen nycress. These protocols allow simple, tissue-culture-free introduction of diverse traits, including altered fatty acid composition and increased oil content. They can also be applied to implement genome editing in camelina and pennycress, enabling precise mutation of genes of inter est. Together, these advances accelerate the agronomic improvement of these oilseeds.