Mineral (nutrient)

In the context of nutrition, a mineral is a chemical element required as an essential nutrient by organisms to perform functions necessary for life.[1][2] Minerals originate in the earth and cannot be made by living organisms.[3] Plants get minerals from soil.[3] Most of the minerals in a human diet come from eating plants and animals or from drinking water.[3] As a group, minerals are one of the four groups of essential nutrients, the others of which are vitamins, essential fatty acids, and essential amino acids.[4]

The five major minerals in the human body are calcium, phosphorus, potassium, sodium, and magnesium.[1] All of the remaining elements in a human body are called "trace elements". The trace elements that have a specific biochemical function in the human body are iron, cobalt, copper, zinc, manganese, molybdenum, iodine, and selenium.[5]

Most chemical elements that are ingested by organisms are in the form of simple compounds. Plants absorb dissolved elements in soils, which are subsequently ingested by the herbivores that eat them, and the elements move up the food chain. Larger organisms may also consume soil (geophagia) or use mineral resources, such as salt licks, to obtain limited minerals unavailable through other dietary sources.

Bacteria and fungi play an essential role in the weathering of primary elements that results in the release of nutrients for their own nutrition and for the nutrition of other species in the ecological food chain. One element, cobalt, is available for use by animals only after having been processed into complex molecules (e.g., vitamin B12) by bacteria. Minerals are used by animals and microorganisms for the process of mineralizing structures, called "biomineralization", used to construct bones, seashells, eggshells, exoskeletons and mollusc shells.[6]

Essential chemical elements for humans

At least twenty chemical elements are known to be required to support human biochemical processes by serving structural and functional roles as well as electrolytes.[7] However, as many as twenty-nine elements in total (including hydrogen, carbon, nitrogen and oxygen) are suggested to be used by mammals, as inferred by biochemical and uptake studies.[8] Among the five major minerals, an adult female has 920-1000 g of calcium while an adult male has ~1.22 kg,[1] with 99% of it contained in bones and teeth, and the other 1% in extracellular fluids, intracellular structures and cell membranes.[1] Phosphorus makes up about 1% of a person's body weight.[9] The last four major minerals (potassium, sodium, chlorine and magnesium) make up only about 0.85% of the weight of the body. Together these eleven chemical elements make up 99.85% of the body.

Most of the known and suggested mineral nutrients are of relatively low atomic weight, and are reasonably common on land, or, at least, common in the ocean (iodine, sodium):

Nutritional elements in the periodic table
H  He
LiBe  B C N O F Ne
NaMg  AlSiP S ClAr
K CaSc TiVCrMnFeCoNiCuZnGaGeAsSeBrKr
RbSrY  ZrNbMoTcRuRhPdAgCdInSnSbTeI Xe
CsBaLa* HfTaW ReOsIrPtAuHgTlPbBiPoAtRn
FrRaAc** RfDbSgBhHsMtDsRgCnNhFlMcLvTsOg
 
  * CePrNdPmSmEuGdTbDyHoErTmYbLu
  **ThPaU NpPuAmCmBkCfEsFmMdNoLr
  The four organic basic elements
  Quantity elements
  Essential trace elements
  Suggested function from deprivation effects or active metabolic handling, but no clearly-identified biochemical function in humans
  Limited circumstantial evidence for trace benefits or biological action in mammals
  No evidence for biological action in mammals, but essential in some lower organisms.
(In the case of lanthanum, the definition of an essential nutrient as being indispensable and irreplaceable is not completely applicable due to the extreme similarity of the lanthanides. Thus Ce, Pr, and Nd may be substituted for La without ill effects for organisms using La, and the smaller Sm, Eu, and Gd may also be similarly substituted but cause slower growth.)

The following play important roles in biological processes:

DV = Daily Value (formerly Recommended Daily Allowance); UL = Tolerable Upper Intake Level (figures shown are for adults age 31-50, male or female neither pregnant nor lactating)

Dietary element DV [mg][10] UL [mg][11][12] Amount Category High nutrient density
dietary sources		 Insufficiency Excess
Potassium 3500 NE Quantity A systemic electrolyte and is essential in coregulating ATP with sodium Sweet potato, tomato, potato, beans, lentils, dairy products, seafood, banana, prune, carrot, orange[13] hypokalemia hyperkalemia
Chlorine 3400 3600 Quantity Needed for production of hydrochloric acid in the stomach and in cellular pump functions Table salt (sodium chloride) is the main dietary source. hypochloremia hyperchloremia
Sodium 2400 2300 Quantity A systemic electrolyte and is essential in coregulating ATP with potassium Table salt (sodium chloride, the main source), sea vegetables, milk, and spinach. hyponatremia hypernatremia
Calcium 1000 2500 Quantity Needed for muscle, heart and digestive system health, builds bone, supports synthesis and function of blood cells Dairy products, eggs, canned fish with bones (salmon, sardines), green leafy vegetables, nuts, seeds, tofu, thyme, oregano, dill, cinnamon.[14] hypocalcaemia hypercalcaemia
Phosphorus 1000 4000 Quantity A component of bones (see apatite), cells, in energy processing, in DNA and ATP (as phosphate) and many other functions Red meat, dairy foods, fish, poultry, bread, rice, oats.[15][16] In biological contexts, usually seen as phosphate[17] hypophosphatemia hyperphosphatemia
Magnesium 400 350 Quantity Required for processing ATP and for bones Spinach, legumes, nuts, seeds, whole grains, peanut butter, avocado[18] hypomagnesemia,
magnesium deficiency		 hypermagnesemia
Iron 18 45 Trace Required for many proteins and enzymes, notably hemoglobin to prevent anemia Meat, seafood, nuts, beans, dark chocolate[19] iron deficiency iron overload disorder
Zinc 15 40 Trace Pervasive and required for several enzymes such as carboxypeptidase, liver alcohol dehydrogenase, and carbonic anhydrase Oysters*, red meat, poultry, nuts, whole grains, dairy products[20] zinc deficiency zinc toxicity
Manganese 2 350 Trace A cofactor in enzyme functions Grains, legumes, seeds, nuts, leafy vegetables, tea, coffee[21] manganese deficiency manganism
Copper 2 11 Trace Required component of many redox enzymes, including cytochrome c oxidase Liver, seafood, oysters, nuts, seeds; some: whole grains, legumes[21] copper deficiency copper toxicity
Iodine 0.150 1.1 Trace Required for synthesis of thyroid hormones, thyroxine and triiodothyronine and to prevent goiter: Seaweed (kelp or kombu)*, grains, eggs, iodized salt[22] iodine deficiency iodism Hyperthyroidism[23]
Chromium 0.120 NE Trace Involved in glucose and lipid metabolism, although its mechanisms of action in the body and the amounts needed for optimal health are not well-defined[24][25] Broccoli, grape juice (especially red), meat, whole grain products[26] Chromium deficiency Chromium toxicity
Molybdenum 0.075 2 Trace The oxidases xanthine oxidase, aldehyde oxidase, and sulfite oxidase[27] Legumes, whole grains, nuts[21] molybdenum deficiency molybdenum toxicity[28]
Selenium 0.070 0.4 Trace Essential to activity of antioxidant enzymes like glutathione peroxidase Brazil nuts, seafoods, organ meats, meats, grains, dairy products, eggs[29] selenium deficiency selenosis
Cobalt none NE Trace Required in the synthesis of vitamin B12, but because bacteria are required to synthesize the vitamin, it is usually considered part of vitamin B12 Cobalt poisoning

* One serving exceeds UL (tolerable upper intake level).

Blood concentrations of minerals

Minerals are present in a healthy human being's blood at certain mass and molar concentrations. The figure below presents the concentrations of each of the chemical elements discussed in this article, from center-right to the right. Depending on the concentrations, some are in upper part of the picture, while others are in the lower part. The figure includes the relative values of other constituents of blood such as hormones. In the figure, minerals are color highlighted in purple.

Reference ranges for blood tests, sorted logarithmically by mass above the scale and by molarity below.

Dietary nutrition

Dietitians may recommend that minerals are best supplied by ingesting specific foods rich with the chemical element(s) of interest. The elements may be naturally present in the food (e.g., calcium in dairy milk) or added to the food (e.g., orange juice fortified with calcium; iodized salt, salt fortified with iodine). Dietary supplements can be formulated to contain several different chemical elements (as compounds), a combination of vitamins and/or other chemical compounds, or a single element (as a compound or mixture of compounds), such as calcium (as calcium carbonate, calcium citrate, etc.) or magnesium (as magnesium oxide, etc.), chromium (usually as chromium(III) picolinate), or iron (as iron bis-glycinate).

The dietary focus on chemical elements derives from an interest in supporting the biochemical reactions of metabolism with the required elemental components.[30] Appropriate intake levels of certain chemical elements have been demonstrated to be required to maintain optimal health. Diet can meet all the body's chemical element requirements, although supplements can be used when some requirements (e.g., calcium, which is found mainly in dairy products) are not adequately met by the diet, or when chronic or acute deficiencies arise from pathology, injury, etc. Research has supported that altering inorganic mineral compounds (carbonates, oxides, etc.) by reacting them with organic ligands (amino acids, organic acids, etc.) improves the bioavailability of the supplemented mineral.[31]

Elements considered possibly essential but not confirmed

Many ultratrace elements have been suggested as essential, but such claims have usually not been confirmed. Definitive evidence for efficacy comes from the characterization of a biomolecule containing the element with an identifiable and testable function.[5] One problem with identifying efficacy is that some elements are innocuous at low concentrations and are pervasive (examples: silicon and nickel in solid and dust), so proof of efficacy is lacking because deficiencies are difficult to reproduce.[30] Ultratrace elements of some minerals such as silicon and boron are known to have a role but the exact biochemical nature is unknown, and others such as arsenic and chromium are suspected to have a role in health, but with weaker evidence.[5] Roles for trace minerals include enzyme catalysis, attracting substrate molecules, redox reactions, and structural or regulatory effects on protein binding.[5]

Element Description Excess
Bromine Possibly important to basement membrane architecture and tissue development, as a needed catalyst to make collagen IV.[32] bromism
Arsenic Essential in rat, hamster, goat and chicken models, but no biochemical mechanism known in humans.[33] arsenic poisoning
Nickel Nickel is an essential component of several enzymes, including urease and hydrogenase.[34] Although not required by humans, some are thought to be required by gut bacteria, such as urease required by some varieties of Bifidobacterium.[35] In humans, nickel may be a cofactor or structural component of certain metalloenzymes involved in hydrolysis, redox rections, and gene expression. Nickel deficiency depressed growth in goats, pigs, and sheep, and diminished circulating thyroid hormone concentration in rats.[36] Nickel toxicity
Fluorine Fluorine (as fluoride) is not considered an essential element because humans do not require it for growth or to sustain life. Research indicates that the primary dental benefit from fluoride occurs at the surface from topical exposure.[37][38] Fluoride poisoning
Boron Boron is an essential plant nutrient, required primarily for maintaining the integrity of cell walls.[39][40][41] Boron has been shown to be essential to complete the life cycle in representatives of all phylogenetic kingdoms, including the model species danio rerio (zebrafish) and Xenopus laevis (African clawed frog).[34][42] In animals, supplemental boron has been shown to reduce calcium excretion and activate vitamin D.[43] Nontoxic
Lithium It is not known whether lithium has a physiological role in any species,[44] but nutritional studies in mammals have indicated its importance to health, leading to a suggestion that it be classed as an essential trace element. Lithium toxicity
Strontium Strontium has been found to be involved in the utilization of calcium in the body. It has promoting action on calcium uptake into bone at moderate dietary strontium levels, but a rachitogenic (rickets-producing) action at higher dietary levels.[45] Rachitogenic (causing Rickets)
Other Silicon and vanadium have established, albeit specialized, biochemical roles as structural or functional cofactors in other organisms, and are possibly, even probably, used by mammals (including humans). By contrast, tungsten, lanthanum, and cadmium have specialized biochemical uses in certain lower organisms, but these elements appear not to be utilized by humans.[8] Other elements considered to be possibly essential include aluminium, germanium, lead, rubidium, and tin.[34][46][47] Multiple

Mineral ecology

Recent studies have shown a tight linkage between living organisms and chemical elements on this planet. This led to the redefinition of minerals as "an element or compound, amorphous or crystalline, formed through 'biogeochemical' processes. The addition of 'bio' reflects a greater appreciation, although an incomplete understanding, of the processes of mineral formation by living forms."[48]:621 Biologists and geologists have only recently started to appreciate the magnitude of mineral biogeoengineering. Bacteria have contributed to the formation of minerals for billions of years and critically define the biogeochemical mineral cycles on this planet. Microorganisms can precipitate metals from solution contributing to the formation of ore deposits in addition to their ability to catalyze mineral dissolution, to respire, precipitate, and form minerals.[49][50][51]

Most minerals are inorganic in nature. Mineral nutrients refers to the smaller class of minerals that are metabolized for growth, development, and vitality of living organisms.[48][52][53] Mineral nutrients are recycled by bacteria that are freely suspended in the vast water columns of the worlds oceans. They absorb dissolved organic matter containing mineral nutrients as they scavenge through the dying individuals that fall out of large phytoplankton blooms. Flagellates are effective bacteriovores and are also commonly found in the marine water column. The flagellates are preyed upon by zooplankton while the phytoplankton concentrates on the larger particulate matter that is suspended in the water column as they are consumed by larger zooplankton, with fish as the top predator. Mineral nutrients cycle through this marine food chain, from bacteria and phytoplankton to flagellates and zooplankton which are then eaten by fish. The bacteria are important in this chain because only they have the physiological ability to absorb the dissolved mineral nutrients from the sea. These recycling principals from marine environments apply to many soil and freshwater ecosystems as well.[54][55] In terrestrial ecosystems, fungi play similar roles as bacteria: they mobilize nutritional elements composing matter that is inaccessible for other organisms and transport acquired nutrients to nutritionally scarce patches of ecosystem.[56]

See also

References

  1. 1 2 3 4 Berdanier, Carolyn D.; Dwyer, Johanna T.; Heber, David (2013). Handbook of Nutrition and Food (3rd ed.). CRC Press. p. 199. ISBN 978-1-4665-0572-8. Retrieved 3 July 2016.
  2. ↑ "Minerals". MedlinePlus, National Library of Medicine, US National Institutes of Health. 22 December 2016. Retrieved 24 December 2016.
  3. 1 2 3 "Minerals". Micronutrient Information Center, Linus Pauling Institute, Oregon State University, Corvallis. 2016. Retrieved 19 December 2016. Minerals are elements that originate in the Earth and cannot be made by living organisms. Plants obtain minerals from the soil, and most of the minerals in our diets come directly from plants or indirectly from animal sources. Minerals may also be present in the water we drink, but this varies with geographic locale. Minerals from plant sources may also vary from place to place, because soil mineral content varies geographically.
  4. ↑ "Vitamin and mineral supplement fact sheets". Office of Dietary Supplements, US National Institutes of Health, Bethesda, MD. 2016. Retrieved 19 December 2016.
  5. 1 2 3 4 Berdanier, Carolyn D.; Dwyer, Johanna T.; Heber, David (19 April 2016). Handbook of Nutrition and Food, Third Edition. CRC Press. pp. 211–224. ISBN 978-1-4665-0572-8. Retrieved 3 July 2016.
  6. ↑ Harris, Ph.D., Edward D. (1 January 2014). Minerals in Food Nutrition, Metabolism, Bioactivity (chapter 3.4) (1st ed.). Lancaster, PA: DEStech Publications, Inc. p. 378. ISBN 978-1-932078-97-8. Retrieved 27 December 2016.
  7. ↑ Nelson, David L.; Michael M. Cox (2000-02-15). Lehninger Principles of Biochemistry, Third Edition (3 Har/Com ed.). W. H. Freeman. p. 1200. ISBN 1-57259-931-6.
  8. 1 2 Ultratrace minerals. Authors: Nielsen, Forrest H. USDA, ARS Source: Modern nutrition in health and disease / editors, Maurice E. Shils ... et al.. Baltimore : Williams & Wilkins, c1999., p. 283-303. Issue Date: 1999 URI:
  9. ↑ "Phosphorus in diet". MedlinePlus, National Library of Medicine, US National Institutes of Health. 2 December 2016. Retrieved 24 December 2016.
  10. ↑ U.S. Food and Drug Administration 14. Appendix F(mg)
  11. ↑ Dietary Reference Intakes (DRIs): Elements Food and Nutrition Board, Institute of Medicine, National Academies (2011)(mg)
  12. ↑ Dietary Reference Intakes : Electrolytes and Water The National Academies (2004)
  13. ↑ "Dietary Guidelines for Americans 2005: Appendix B-1. Food Sources of Potassium". United States Department of Agriculture. 2005.
  14. ↑ Adam Drewnowski (2010). "The Nutrient Rich Foods Index helps to identify healthy, affordable foods" (PDF). The American Journal of Clinical Nutrition. 91(suppl): 1095S–1101S.
  15. ↑ "NHS Choices:Vitamins and minerals – Others". Retrieved November 8, 2011.
  16. ↑ Corbridge, D. E. C. (1995-02-01). Phosphorus: An Outline of Its Chemistry, Biochemistry, and Technology (5th ed.). Amsterdam: Elsevier Science Pub Co. p. 1220. ISBN 0-444-89307-5.
  17. ↑ "Linus Pauling Institute at Oregon State University". Retrieved 2008-11-29.
  18. ↑ "Magnesium—Fact Sheet for Health Professionals". National Institutes of Health. 2016.
  19. ↑ "Iron—Dietary Supplement Fact Sheet". National Institutes of Health. 2016.
  20. ↑ "Zinc—Fact Sheet for Health Professionals". National Institutes of Health. 2016.
  21. 1 2 3 Schlenker, Eleanor; Gilbert, Joyce Ann (28 August 2014). Williams' Essentials of Nutrition and Diet Therapy. Elsevier Health Sciences. pp. 162–3. ISBN 978-0-323-29401-0. Retrieved 15 July 2016.
  22. ↑ "Iodine—Fact Sheet for Health Professionals". National Institutes of Health. 2016.
  23. ↑ Jameson, J. Larry; De Groot, Leslie J. (25 February 2015). Endocrinology: Adult and Pediatric. Elsevier Health Sciences. p. 1510. ISBN 978-0-323-32195-2. Retrieved 14 July 2016.
  24. ↑ Kim, Myoung Jin; Anderson, John; Mallory, Caroline (1 February 2014). Human Nutrition. Jones & Bartlett Publishers. p. 241. ISBN 978-1-4496-4742-1. Retrieved 10 July 2016.
  25. ↑ Gropper, Sareen S.; Smith, Jack L. (1 June 2012). Advanced Nutrition and Human Metabolism. Cengage Learning. pp. 527–8. ISBN 1-133-10405-3. Retrieved 10 July 2016.
  26. ↑ "Chromium". Office of Dietary Supplements, US National Institutes of Health. 2016. Retrieved 10 July 2016.
  27. ↑ Sardesai VM (December 1993). "Molybdenum: an essential trace element". Nutr Clin Pract. 8 (6): 277–81. PMID 8302261. doi:10.1177/0115426593008006277.
  28. ↑ Momcilović, B. (September 1999). "A case report of acute human molybdenum toxicity from a dietary molybdenum supplement—a new member of the "Lucor metallicum" family.". Archives of Industrial Hygiene and Toxicology. De Gruyter. 50 (3): 289–97. PMID 10649845.
  29. ↑ "Selenium—Fact Sheet for Health Professionals". National Institutes of Health. 2016.
  30. 1 2 Lippard, Stephen J.; Jeremy M. Berg (1994). Principles of Bioinorganic Chemistry. Mill Valley, CA: University Science Books. p. 411. ISBN 0-935702-72-5.
  31. ↑ Ashmead, H. DeWayne (1993). The Roles of Amino Acid Chelates in Animal Nutrition. Westwood: Noyes Publications.
  32. ↑ A. Scott McCall; Christopher F. Cummings; Gautam Bhave; Roberto Vanacore; Andrea Page-McCaw; Billy G. Hudson (5 June 2014). "Bromine Is an Essential Trace Element for Assembly of Collagen IV Scaffolds in Tissue Development and Architecture". Cell. 157 (6): 1380–1392. PMC 4144415 Freely accessible. PMID 24906154. doi:10.1016/j.cell.2014.05.009.
  33. ↑ Anke M. Arsenic. In: Mertz W. ed., Trace elements in human and Animal Nutrition, 5th ed. Orlando, FL: Academic Press, 1986, 347–372; Uthus E.O., Evidency for arsenical essentiality, Environ. Geochem. Health, 1992, 14:54–56; Uthus E.O., Arsenic essentiality and factors affecting its importance. In: Chappell W.R, Abernathy C.O, Cothern C.R. eds., Arsenic Exposure and Health. Northwood, UK: Science and Technology Letters, 1994, 199–208.
  34. 1 2 3 Berdanier, Carolyn D.; Dwyer, Johanna T.; Heber, David (19 April 2016). Handbook of Nutrition and Food, Third Edition. CRC Press. pp. 211–26. ISBN 978-1-4665-0572-8. Retrieved 3 July 2016.
  35. ↑ Sigel, Astrid; Sigel, Helmut; Sigel, Roland K. O. (27 January 2014). Interrelations between Essential Metal Ions and Human Diseases. Springer Science & Business Media. p. 349. ISBN 978-94-007-7500-8. Retrieved 4 July 2016.
  36. ↑ Institute of Medicine (29 September 2006). Dietary Reference Intakes: The Essential Guide to Nutrient Requirements. National Academies Press. pp. 313–19, 415–22. ISBN 978-0-309-15742-1. Retrieved 21 June 2016.
  37. ↑ Mitsuo Kakei, Toshiro Sakae and Masayoshi Yoshikawa (2012). "Aspects Regarding Fluoride Treatment for Reinforcement and Remineralization of Apatite Crystals". Journal of Hard Tissue Biology. 21 (3): 475–6. Retrieved 2017-06-01.
  38. ↑ Peter Loskill, Christian Zeitz, Samuel Grandthyll, Nicolas Thewes, Frank Müller, Markus Bischoff, Mathias, Herrmann, Karin Jacobs (2013). "Reduced Adhesion of Oral Bacteria on Hydroxyapatite by Fluoride Treatment". Langmuir. doi:10.1021/la4008558. Retrieved 2017-06-01.
  39. ↑ Mahler, R. L. "Essential Plant Micronutrients. Boron in Idaho" (PDF). University of Idaho. Archived from the original (PDF) on 1 October 2009. Retrieved 2009-05-05.
  40. ↑ "Functions of Boron in Plant Nutrition" (PDF). U.S. Borax Inc. Archived from the original (PDF) on 20 March 2009.
  41. ↑ Blevins, Dale G.; Lukaszewski, KM (1998). "Functions of Boron in Plant Nutrition". Annual Review of Plant Physiology and Plant Molecular Biology. 49 (1): 481–500. PMID 15012243. doi:10.1146/annurev.arplant.49.1.481.
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  43. ↑ Nielsen, Forrest H. (1997). "Boron in human and animal nutrition". Plant and Soil. 193 (2): 199–208. ISSN 0032-079X. doi:10.1023/A:1004276311956.
  44. ↑ "Some Facts about Lithium". ENC Labs. Retrieved 2010-10-15.
  45. ↑ "The biological role of strontium". Retrieved 2010-10-06.
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  48. 1 2 Skinner, H. C. W. (2005). "Biominerals". Mineralogical Magazine. 69 (5): 621–641. doi:10.1180/0026461056950275.
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  51. ↑ González-Muñoz, M. T.; Rodriguez-Navarro, C.; Martinez-Ruiz, F.; Arias, J. M.; Merroun, M. L.; Rodriguez-Gallego, M. (2010). "Bacterial biomineralization: new insights from Myxococcus-induced mineral precipitation". Geological Society, London, Special Publications. 336 (1): 31–50. doi:10.1144/SP336.3.
  52. ↑ Kirkby, H.; Kirkby, E. A.; Cakmak, I. (1996). "Effect of mineral nutritional status on shoot-root partitioning of photoassimilates and cycling of mineral nutrients" (PDF). Journal of Experimental Biology. 47 (S1255): 1255. PMID 21245257. doi:10.1093/jxb/47.Special_Issue.1255.
  53. ↑ Adame, L. (2002). "Leaf absorption of mineral nutrients in carnivorous plants stimulates root nutrient uptake" (PDF). New Phytologist. 155: 89–100. doi:10.1046/j.1469-8137.2002.00441.x.
  54. ↑ Azam, F.; Fenchel, T.; Field, J. G.; Gray, J. S.; Meyer-Reil, L. A.; Thingstad, F. (1983). "The ecological role of water-column microbes in the sea" (PDF). Mar. Ecol. Prog. Ser. 10: 257–263. doi:10.3354/meps010257.
  55. ↑ Uroz, S.; Calvaruso, C.; Turpault, M.; Frey-Klett, Pascale (2009). "Mineral weathering by bacteria: ecology, actors and mechanisms" (PDF). Trends in Microbiology. 17 (8): 378–87. PMID 19660952. doi:10.1016/j.tim.2009.05.004.
  56. ↑ J. Dighton (2007). "Nutrient Cycling by Saprotrophic Fungi in Terrestrial Habitats". In Kubicek, Christian P.; Druzhinina, Irina S. Environmental and microbial relationships (2nd ed.). Berlin: Springer. pp. 287–300. ISBN 978-3-540-71840-6.

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