Spermidine Food Sources and Dosing

The typical Western diet supplies somewhere between 7 and 15 mg of total polyamines per day — a mixture of putrescine, spermidine, and spermine — with spermidine itself usually contributing 1 to 3 mg in adults who eat little fermented or germinated food. The Bruneck cohort that produced the landmark 2018 BMJ mortality paper showed that participants in the highest tertile of dietary spermidine intake averaged closer to 12 mg/day, achieved primarily through a Mediterranean-leaning food pattern with substantial whole-grain, legume, fermented dairy, and mushroom content. The single richest practical food source is wheat germ, at roughly 24 mg of spermidine per 100 g — a heaping tablespoon stirred into yogurt or oatmeal delivers more spermidine than most people eat in a full day from all other sources combined. Natto (fermented soybean) is the highest-concentration food in the world at approximately 250 mg/kg, courtesy of Bacillus subtilis polyamine biosynthesis during fermentation. Aged cheeses — well-aged cheddar, parmigiano-reggiano, gruyère — concentrate spermidine through the polyamine output of the ripening bacterial cultures over months of aging. Mushrooms, particularly shiitake and oyster, contribute meaningfully on a serving basis. Legumes (peas, lentils, soybeans) and sprouted seeds round out the dietary picture. This deep-dive page walks through the food-source data quantitatively, traces the gut-microbiome contribution that roughly triples whatever the diet supplies, explains why aged and fermented foods are so much richer than their fresh counterparts, summarizes the human supplementation trials and their typical 1-5 mg/day dosing window, and gives practical guidance on building a daily intake plan from food first.

Table of Contents

  1. The Three Dietary Polyamines — What "Spermidine Intake" Actually Means
  2. Wheat Germ — The #1 Dietary Source
  3. Natto and the Fermentation Effect
  4. Aged Cheeses — Cheddar, Parmesan, Gruyère
  5. Mushrooms, Legumes, and Sprouted Seeds
  6. The Mediterranean Diet and the Bruneck Cohort
  7. The Gut Microbiome — Roughly Three Times the Dietary Input
  8. Age-Related Decline in Endogenous Polyamine Synthesis
  9. Cooking, Storage, and Bioavailability
  10. Wheat-Germ Extract, Synthetic, and Aged-Cheese Delivery Compared
  11. Human Supplementation Dose Range (1-5 mg/day SmartAge Window)
  12. Building a Daily Plan from Food First
  13. Key Research Papers
  14. Connections

The Three Dietary Polyamines — What "Spermidine Intake" Actually Means

Before discussing food sources, it is worth pinning down what gets measured. The dietary polyamines are three small aliphatic molecules: putrescine (a diamine, 4 carbons, 2 amine groups), spermidine (a triamine, 7 carbons, 3 amine groups), and spermine (a tetraamine, 10 carbons, 4 amine groups). The three are biosynthetically linked — putrescine is converted to spermidine by spermidine synthase, and spermidine is converted to spermine by spermine synthase, with decarboxylated S-adenosylmethionine donating the aminopropyl group at each step. The three interconvert biologically through a complementary recycling pathway as well, so a given cell or food sample contains all three in varying proportions.

Most food-composition studies measure all three and report them separately. When the literature refers to "spermidine intake," it almost always means spermidine specifically rather than total polyamines, but the broader concept of "polyamine intake" matters because the body can interconvert them to some extent. Foods that are rich in spermidine tend also to be rich in spermine and putrescine, and the Bruneck mortality paper used a polyamine composite as well as a spermidine-specific analysis, with similar results on both measures.

The food-composition database that underpins most modern work is the Atiya Ali / Strindberg et al. compilation from the early 2010s, supplemented by Munoz-Esparza and Loaec for European foods. Concentrations are usually reported in milligrams per kilogram (mg/kg) of food as eaten, which converts cleanly to milligrams per 100 g by dividing by 10. A practical daily target in the spermidine-longevity literature is somewhere between 6 and 12 mg/day of spermidine from food — the lower end of that range moves a typical Western diet up by 4-6 mg through deliberate food choices, and the upper end approximates the highest Bruneck tertile and the wheat-germ-extract supplementation arms of the clinical trials.

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Wheat Germ — The #1 Dietary Source

Wheat germ — the small, lipid-rich, vitamin-rich embryo of the wheat kernel that is removed during the production of white flour — is by a substantial margin the most concentrated routine food source of spermidine in the human diet. Reported concentrations cluster around 24 mg per 100 g, with some samples ranging from 18 to 30 mg/100 g depending on wheat cultivar, harvest conditions, and storage. A heaping tablespoon (about 7 g) of wheat germ thus contains roughly 1.7 mg of spermidine. A quarter cup (about 28 g) delivers approximately 6.7 mg — more than triple the typical full-day Western intake from all other sources combined.

The germ is the metabolically active embryonic tissue of the seed, and like other germinating or germinable plant tissues it has high polyamine concentrations because polyamines are essential for the rapid cell division that follows germination. Sprouted whole-grain breads, sprouted wheat berries, and sprouted-grain cereals retain meaningful spermidine content, though usually less per gram than isolated wheat germ. Wheat bran (the outer fibrous layer) has much less spermidine than the germ. White flour, with both bran and germ removed, has almost none — one of the many reasons that the shift from whole-grain to refined-flour diets in the twentieth century reduced background polyamine intake substantially.

Practically, wheat germ is sold in two forms: raw (refrigerated for stability because the lipid content oxidizes quickly at room temperature) and toasted (more shelf-stable but with slight loss of vitamin E and possibly some polyamine content during the toasting step, though the spermidine itself is heat-stable enough that the loss is modest). Either form works for dietary purposes. The lipid content makes wheat germ go rancid quickly once opened — store in the refrigerator or freezer in a tightly sealed container and use within 2-3 months for best results. The taste is mild and slightly nutty; it works well stirred into yogurt, oatmeal, smoothies, batters, and salads. A tablespoon-a-day habit is a reasonable foundation for anyone wanting to raise dietary spermidine intake without supplements.

Caveat for gluten-sensitive patients: wheat germ contains gluten and is not suitable for people with celiac disease or non-celiac gluten sensitivity. The polyamine concentration of other germinated cereal embryos (rye germ, barley germ) is similar in magnitude but they share the gluten problem. For gluten-free dietary spermidine, the main contributors become mushrooms, legumes, aged cheeses (some are essentially gluten-free), natto, and sprouted gluten-free grains and seeds.

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Natto and the Fermentation Effect

Natto is a traditional Japanese fermented soybean preparation in which whole cooked soybeans are inoculated with Bacillus subtilis var. natto and fermented for roughly 18-24 hours, producing the characteristic stringy, glutinous, pungent finished product. Natto has the highest spermidine concentration of any commonly consumed food on the global map — published values cluster around 250 mg/kg, or 25 mg per 100 g, which is roughly comparable to wheat germ on a per-weight basis. A standard 40-50 g serving of natto thus delivers approximately 10-12 mg of spermidine in a single meal.

The crucial observation is that the spermidine concentration in finished natto is roughly an order of magnitude higher than in the cooked soybeans used as the starting material. The polyamine content is built during fermentation by Bacillus subtilis itself, which has a robust polyamine biosynthesis pathway and produces both putrescine and spermidine as part of its normal metabolism. The fermentation effect is the most striking practical example of how microbial activity dominates dietary polyamine availability — a fermented version of almost any food has substantially more spermidine than its unfermented precursor.

Other fermented foods show the same effect to varying degrees. Tempeh (fermented soybean cake produced with Rhizopus molds) has elevated polyamines compared to plain soybeans, though less than natto. Miso (long-fermented soybean and grain paste) has meaningful concentrations and contributes broadly to Japanese-pattern dietary polyamine intake. Kimchi and sauerkraut (lactic-acid-fermented vegetables) contribute modest but non-trivial spermidine. Aged cheeses (discussed separately below) are the European equivalent of the fermentation amplification effect.

Natto remains an acquired taste for non-Japanese palates — the sulfurous aroma, the slimy mucilaginous texture, and the strong ammoniacal note from continued polyamine production during refrigerated storage are all challenging on first introduction. For dietary spermidine purposes the practical question is whether the patient can tolerate it as a regular item; many people find that natto stirred into rice with soy sauce, chopped green onion, and a raw egg yolk (the traditional preparation) is acceptable. Frozen natto stored well and thawed on the morning of consumption maintains its polyamine content. For patients who simply cannot tolerate the taste or texture, the alternative is to source polyamine intake from other foods and supplements.

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Aged Cheeses — Cheddar, Parmesan, Gruyère

Aged hard cheeses are the European counterpart to natto in the dietary polyamine economy. Fresh cheeses (mozzarella, ricotta, cottage cheese) have low polyamine concentrations because the bacterial cultures have not had time to produce them. Soft-ripened cheeses (brie, camembert) have moderate concentrations. Aged hard cheeses — varieties like cheddar, parmigiano-reggiano, grana padano, gruyère, manchego, aged gouda — have substantially higher concentrations because their bacterial and mold cultures continue to produce polyamines through months and years of ripening.

Reported concentrations vary widely by variety, age, and producer:

The polyamines in aged cheese are produced by the secondary microbial flora — Lactobacillus, Lactococcus, Enterococcus, Brevibacterium, and others depending on the variety — that continue metabolic activity during the ripening period. Polyamine production scales roughly with aging time and with the diversity of the microbial community, which is why the most polyamine-rich cheeses tend to be the most pungently flavored ones — the same bacterial activity that produces the flavor compounds produces the polyamines.

A practical implication is that the spermidine content of cheese is not predictable from the label — it depends on age and bacterial community details that the producer may not disclose precisely. As a rough rule, the older and stinkier and more crystalline (those crunchy white crystals in aged parmesan are tyrosine, an aging marker), the more polyamines. Patients building a polyamine-rich diet can usefully prioritize an ounce or two of well-aged hard cheese daily as a flavor-positive contribution. The biogenic amine load of aged cheese is also a relevant consideration for patients on MAO inhibitors (a classic interaction with aged cheese is the tyramine reaction; spermidine and spermine themselves are not implicated in the MAOI reaction, but the aged-cheese category as a whole is restricted in that population).

For more on the cheese category in general nutrition terms, see Cheese. The connection to dietary polyamines is one of several reasons that the traditional Mediterranean diet pattern, which routinely includes 1-2 oz of aged cheese daily, appears to confer some of its longevity benefit through the polyamine route.

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Mushrooms, Legumes, and Sprouted Seeds

Mushrooms are a substantial and often-underappreciated source of dietary polyamines. Fungi as a kingdom have particularly active polyamine metabolism, and several common culinary mushrooms have spermidine concentrations in the 50-90 mg/kg range:

Legumes contribute meaningfully on a per-serving basis because they are eaten in larger portions than mushrooms:

Sprouted seeds in general are polyamine-rich, for the same germination biology reason that wheat germ is. Broccoli sprouts, alfalfa sprouts, sprouted sunflower seeds, sprouted pumpkin seeds, and sprouted radish seeds all contribute. Concentrations are not as systematically catalogued as for the major cereal germs, but the directional pattern is consistent — any rapidly germinating plant tissue has elevated polyamines as a matter of plant biology.

For more on the mushroom category, see Mushrooms and the related deep-dive on Medicinal Mushrooms. The legume category sits at Lentils and the broader Food hub.

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The Mediterranean Diet and the Bruneck Cohort

The Bruneck study is the single most influential observational cohort for the spermidine-mortality hypothesis. The cohort consists of approximately 800 adults living in the South Tyrolean alpine town of Bruneck, Italy, recruited in 1990 and followed prospectively for cardiovascular outcomes, cancer incidence, and all-cause mortality. The dietary assessment used food-frequency questionnaires translated into estimated polyamine intake through the Munoz-Esparza European food composition tables.

Kiechl, Pechlaner, Madeo, and colleagues published the polyamine-mortality analysis in the British Medical Journal in 2018. The key findings:

The honest interpretation of the Bruneck data is that observational cohort evidence cannot establish causation — the highest-spermidine tertile differs from the lowest in many ways beyond polyamine intake, and even rigorous statistical adjustment leaves residual confounding. The plausibility of the spermidine-specific mechanism is supported by the mechanistic work in autophagy and hypusinated eIF5A, by the parallel animal-model lifespan data, and by emerging short-term human trials of wheat-germ-extract supplementation, but a definitive randomized hard-outcome trial in humans has not yet been completed.

The practical lesson from Bruneck is twofold: first, that the magnitude of difference between the lowest and highest tertiles (6 vs 12 mg/day) is achievable through routine food choices without supplementation; and second, that the food pattern that produced the high-intake tertile is not an obscure prescriptive diet but a recognizable Mediterranean-leaning whole-food pattern that is independently associated with longevity benefit in dozens of cohorts. The polyamine angle is one of several plausible mechanisms by which the Mediterranean pattern delivers its benefit.

For more on the Mediterranean dietary pattern in general, see Mediterranean Diet. The cardiovascular-specific aspects of the Bruneck data are detailed in Cardiovascular Aging.

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The Gut Microbiome — Roughly Three Times the Dietary Input

One of the most important and least-appreciated facts about spermidine biology is that the gut microbiome produces substantially more polyamines than the diet supplies. The estimate from tracer studies and from germ-free vs conventional animal comparisons is that intestinal microbial polyamine biosynthesis contributes roughly three times the daily polyamine load reaching the systemic circulation, dwarfing dietary input. The dominant microbial producers are Bacteroides species, Fusobacterium, Clostridium, Escherichia coli, and various lactic-acid bacteria — all common commensals in the healthy adult colon.

The implication is that the relevant variable for systemic polyamine availability is not just dietary intake but also the composition and activity of the colonic microbiome. A patient with a robust, diverse, polyamine-producing microbiome may reach adequate systemic polyamine levels even on a relatively low dietary intake. Conversely, a patient with antibiotic-disrupted microbiome, inflammatory bowel disease, or other dysbiosis may have substantially reduced microbial polyamine production despite adequate dietary intake.

The microbiome-polyamine link is bidirectional. Polyamine intake (both dietary and microbial) shapes the microbiome itself — polyamines are growth substrates for some bacterial species and selectively favor certain community structures. Fermentable fibers (the prebiotics) feed the bacteria that produce polyamines, so a fiber-rich diet supports both microbial polyamine production directly (more bacterial biomass) and indirectly (community composition favoring producer species). Resistant starch, beta-glucans (from oats and mushrooms), pectins (from fruit), and inulin-type fructans (from onions, garlic, leeks, jerusalem artichokes) all support the colonic environment that produces polyamines.

The clinical implication is that any intervention that disrupts the microbiome — especially broad-spectrum antibiotic courses — can be expected to transiently reduce systemic polyamine availability for weeks to months until the microbiome recovers. Patients planning a spermidine-rich dietary or supplemental intervention may want to time it to begin after, not during, an unavoidable antibiotic course, and to combine the polyamine intervention with prebiotic fiber support and (potentially) a multi-strain probiotic.

For more on the microbiome side of this picture, see Gut Microbiome and the related Prebiotics and Probiotics pages.

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The cellular machinery for endogenous polyamine biosynthesis declines progressively with chronological age. The rate-limiting enzyme, ornithine decarboxylase (ODC1), has lower steady-state activity in tissues from older animals and older humans. Spermidine synthase and S-adenosylmethionine decarboxylase (which produces the aminopropyl donor for the synthase step) similarly decline. The net result is that an older adult, given the same dietary polyamine input as a younger adult, ends up with lower tissue polyamine concentrations.

Hofer and colleagues documented the age-related decline in cardiac tissue spermidine in mice and showed that dietary supplementation could restore the levels to those of younger animals. Pucciarelli and Madeo's laboratory has shown analogous patterns in human peripheral blood mononuclear cells — older donors have lower baseline spermidine and lower hypusinated eIF5A than younger donors, and exogenous spermidine raises both. The decline appears to be roughly 30-50% between young adulthood and the eighth decade of life, with substantial inter-individual variation.

This age-related decline is part of the rationale for why dietary or supplemental spermidine repletion might be particularly relevant in older adults, even those whose dietary intake appears adequate on paper. The aging body simply does not retain or produce polyamines as efficiently as the younger body, and an intake that was sufficient at age 30 may be borderline at age 70. The Bruneck cohort data showed that the highest dietary tertile's mortality benefit was particularly pronounced in the older participants, consistent with the age-related-decline framework — the older you are, the more replacement value the dietary input provides.

The same age-related decline pattern is documented for other endogenously synthesized molecules central to the longevity story — NAD+, glutathione, coenzyme Q10, and others. Spermidine fits comfortably in that category of "endogenous molecule whose levels fall with age and whose replacement may have anti-aging value." The parallels to NAD+ and NMN, Glutathione, and Coenzyme Q10 repletion strategies are deliberate and conceptually unified.

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Cooking, Storage, and Bioavailability

Spermidine itself is a small, chemically stable molecule. It survives ordinary cooking temperatures essentially intact — boiling, baking, sauteing, grilling, and pressure cooking do not significantly degrade it. The cooked weight of a portion is therefore a reasonable basis for estimating polyamine intake, and standard food-composition databases generally report values for foods as eaten rather than as raw weight.

That said, there are some practical considerations:

The practical bottom line is that almost any way of cooking polyamine-rich foods preserves most of the polyamine content. The only meaningful loss is from discarding boiling liquid in the classical drain-the-water vegetable preparations. The fermentation and germination amplifications are large — reaching for the fermented or sprouted version of a food is one of the single highest-yield dietary moves.

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Wheat-Germ Extract, Synthetic, and Aged-Cheese Delivery Compared

Three different supplemental and dietary delivery formats for spermidine are used in the modern marketplace and the academic literature:

  1. Wheat-germ-extract supplements — concentrated polyamine-rich extracts of wheat germ, typically standardized to deliver a specified dose of spermidine (commonly 1-3 mg per capsule, with 1.2 mg being the most common standardized dose). These were the formulation used in the Wirth, Schwarz, and Schroeder human trials, including the SmartAge cognitive trial. The advantages: well-characterized clinical safety record, food-matrix delivery with the co-present nutrients (vitamin E, B vitamins, healthy fats) intact, mechanism of action confirmed in the same product format used in trials. The disadvantages: standardized dose is still relatively low compared to the high-intake Bruneck tertile, gluten content (relevant for celiac and non-celiac gluten-sensitive patients), product-to-product variation in standardization rigor across brands.
  2. Synthetic spermidine — chemically synthesized spermidine trihydrochloride or spermidine free base, sold as powder or in capsules. The advantages: precise dose control, no gluten, can deliver higher doses per capsule, generally lower cost per milligram of spermidine. The disadvantages: no food matrix, less mature human clinical safety record than wheat-germ extract (though no specific safety signal has emerged), regulatory ambiguity in some jurisdictions, sometimes a slightly metallic or bitter taste in powder form.
  3. Aged-cheese delivery — getting the polyamine dose through routine consumption of aged cheese as part of meals. The advantages: pleasant to consume, integrates seamlessly with Mediterranean-pattern eating, provides additional nutrients (calcium, vitamin K2, protein, beneficial fatty acids), no separate supplement-taking ritual. The disadvantages: variable polyamine content by variety and age, calorically dense (high saturated fat and sodium content), tyramine and other biogenic amines may matter for MAOI patients and migraine-sensitive individuals, lactose-intolerant patients may need to focus on aged hard cheeses (which are essentially lactose-free) rather than fresh varieties.

For most patients, the best practical approach is a combination — build a daily dietary baseline through wheat germ, mushrooms, legumes, and aged cheese, and add a wheat-germ-extract supplement only if the dietary baseline does not reach the target range or if the patient prefers supplemental insurance. The wheat-germ-extract supplements used in the clinical trials are widely available; common brands include Longevity Labs/spermidineLIFE, Primeadine, and Oxford Healthspan, all standardized to specified spermidine content per serving and using wheat-germ-extract as the source material.

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Human Supplementation Dose Range (1-5 mg/day SmartAge Window)

The dose range that has been studied in formal human trials is narrow: most studies have used between 1 and 5 mg of spermidine per day in supplemental form, with 1.2 mg being the most common single-capsule dose and 3-5 mg being the upper end of trial-tested daily intake. The major human trials and their dosing:

The upper end of safe supplemental intake in humans is not formally established because the trials have not pushed the dose higher than necessary. Animal studies have used very high doses (10-50 mg/kg, which would scale to hundreds of mg/day human-equivalent) without obvious acute toxicity. The conservative practical position is that the 1-5 mg/day supplemental window has good safety data, doses in the 5-15 mg/day range (achievable through diet and supplements combined) have substantial observational safety data from the Bruneck cohort and similar populations who naturally consume those amounts, and substantially higher pharmacologic doses are not established for general use.

Most patients translating the literature into a personal protocol target somewhere in the 1-3 mg/day supplemental range on top of a polyamine-rich dietary baseline, for a total intake roughly approximating the Bruneck high-tertile of 10-12 mg/day. This is also approximately the dose range used in the active SmartAge and follow-on trials and the range with the most direct human evidence.

The cancer caveat noted in the Autophagy Induction page applies to dosing decisions as well — polyamines including spermidine are required for cell proliferation, and isolated polyamine biology in oncology has been studied as a potential growth-promotion concern. The clinical evidence on dietary spermidine and cancer outcomes is mixed but not pointing in a strongly worrying direction; the Bruneck cohort showed no excess cancer mortality in the highest-intake tertile. Patients with active cancer or recent cancer history should discuss the dietary and supplemental intake with their oncology team before launching a deliberate polyamine-elevation strategy.

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Building a Daily Plan from Food First

A workable food-first daily plan that approximates the Bruneck high-intake tertile looks something like this. The numbers below are intended as a rough sketch rather than a prescriptive recipe.

Patients who cannot or will not eat one of the major categories (no wheat for celiac, no cheese for vegan, no soy for allergy) can still reach the target by emphasizing the other categories more heavily. A gluten-free patient might rely more heavily on mushrooms, legumes, aged cheese, and sprouted gluten-free seeds. A vegan patient might rely more heavily on natto/tempeh, wheat germ, mushrooms, sprouted legumes, and sprouted seeds. A patient with cheese restriction might rely more heavily on wheat germ, natto, mushrooms, and legumes. The category diversity in the polyamine sources is one of their underappreciated practical strengths — almost any reasonable dietary pattern can be adjusted to reach the target intake.

For patients who cannot or do not want to engineer the diet to this degree, a 1.2-3 mg/day wheat-germ-extract supplement on top of an ordinary varied diet is a reasonable alternative. The supplement does not replicate the broader Mediterranean-pattern benefit of the food-first approach, but it does deliver the spermidine itself in a form that has been tested in human trials.

For the broader rationale behind dietary intervention for cognitive and cardiovascular aging, see Cognitive Function and Cardiovascular Aging. For the autophagy mechanism that underlies most of the benefit, see Autophagy Induction.

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Key Research Papers

  1. Kiechl S, Pechlaner R, Willeit P, Notdurfter M, Paulweber B, Willeit K, et al. (2018). Higher spermidine intake is linked to lower mortality: a prospective population-based study. American Journal of Clinical Nutrition 108(2):371-380. — PubMed
  2. Madeo F, Eisenberg T, Pietrocola F, Kroemer G (2018). Spermidine in health and disease. Science 359(6374):eaan2788. — PubMed
  3. Schwarz C, Stekovic S, Wirth M, Benson G, Royer P, Sigrist SJ, et al. (2018). Safety and tolerability of spermidine supplementation in mice and older adults with subjective cognitive decline. Aging 10(1):19-33. — PubMed
  4. Wirth M, Benson G, Schwarz C, Kobe T, Grittner U, Schmitz D, et al. (2018). The effect of spermidine on memory performance in older adults at risk for dementia: a randomized controlled trial. Cortex 109:181-188. — PubMed
  5. Schroeder S, Hofer SJ, Zimmermann A, Pechlaner R, Dammbrueck C, Pendl T, et al. (2021). Dietary spermidine improves cognitive function. Cell Reports 35(2):108985. — PubMed
  6. Atiya Ali M, Poortvliet E, Stromberg R, Yngve A (2011). Polyamines in foods: development of a food database. Food & Nutrition Research 55. — PubMed
  7. Munoz-Esparza NC, Latorre-Moratalla ML, Comas-Baste O, Toro-Funes N, Veciana-Nogues MT, Vidal-Carou MC (2019). Polyamines in food. Frontiers in Nutrition 6:108. — PubMed
  8. Pucciarelli S, Moreschini B, Micozzi D, De Fronzo GS, Carpi FM, Polzonetti V, et al. (2012). Spermidine and spermine are enriched in whole blood of nona/centenarians. Rejuvenation Research 15(6):590-595. — PubMed
  9. Soda K, Kano Y, Sakuragi M, Takao K, Lefor A, Konishi F (2009). Long-term oral polyamine intake increases blood polyamine concentrations. Journal of Nutritional Science and Vitaminology 55(4):361-366. — PubMed
  10. Eisenberg T, Abdellatif M, Schroeder S, Primessnig U, Stekovic S, Pendl T, et al. (2016). Cardioprotection and lifespan extension by the natural polyamine spermidine. Nature Medicine 22(12):1428-1438. — PubMed
  11. Matsumoto M, Kibe R, Ooga T, Aiba Y, Kurihara S, Sawaki E, et al. (2012). Impact of intestinal microbiota on intestinal luminal metabolome. Scientific Reports 2:233. — PubMed
  12. Hofer SJ, Liang Y, Zimmermann A, Schroeder S, Dengjel J, Kroemer G, et al. (2021). Spermidine-induced hypusination preserves mitochondrial and cognitive function during aging. Aging Cell 20(4):e13328. — PubMed
  13. Loaec G, Niquet-Leridon C, Henry N, Jacolot P, Volpoet G, Goudemand E, et al. (2014). Effects of variety, agronomic factors, and drying on the amount of free asparagine and crude protein in French wheat. Mapping of the potential acrylamide production. Journal of Cereal Science 60(1):153-159. — PubMed

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