Humus
Humus
In classical[1] soil science, humus is the dark organic matter in soil that is formed by the decomposition of
plant and animal matter. It is a kind of soil organic matter. It is rich in nutrients and retains moisture in the
soil. Humus is the Latin word for "earth" or "ground".[2]
In agriculture, "humus" sometimes also is used to describe mature or natural compost extracted from a
woodland or other spontaneous source for use as a soil conditioner.[3] It is also used to describe a topsoil
horizon that contains organic matter (humus type,[4] humus form,[5] or humus profile[6]).
Humus has many nutrients that improve the health of soil, nitrogen being the most important. The ratio of
carbon to nitrogen (C:N) of humus commonly ranges between 8:1 and 15:1 with the median being about
12:1.[7] It also significantly improves (decreases) the bulk density of soil.[8] Humus is amorphous and lacks
the cellular structure characteristic of organisms.[9]
The solid residue of sewage sludge treatment, which is a secondary phase in the wastewater treatment
process, is also called humus.[10] When not judged contaminated by pathogens, toxic heavy metals, or
persistent organic pollutants according to standard tolerance levels, it is sometimes composted and used
as a soil amendment.[11]
Description
The primary materials needed for the process of humification are plant detritus and dead animals and
microbes, excreta of all soil-dwelling organisms, and also black carbon resulting from past fires.[12] The
composition of humus varies with that of primary (plant) materials and secondary microbial and animal
products. The decomposition rate of the different compounds will affect the composition of the humus.[13]
It is difficult to define humus precisely because it is a very complex substance which is still not fully
understood. Humus is different from decomposing soil organic matter. The latter looks rough and has
visible remains of the original plant or animal matter. Fully humified humus, on the contrary, has a uniformly
dark, spongy, and jelly-like appearance, and is amorphous; it may gradually decay over several years or
persist for millennia.[14] It has no determinate shape, structure, or quality. However, when examined under a
microscope, humus may reveal tiny plant, animal, or microbial remains that have been mechanically, but not
chemically, degraded.[15] This suggests an ambiguous boundary between humus and soil organic matter,
leading some authors to contest the use of the term humus and derived terms such as humic substances or
humification, proposing the Soil Continuum Model (SCM).[16] However, humus can be considered as having
distinct properties, mostly linked to its richness in functional groups, justifying its maintenance as a
specific term.[17]
Fully formed humus is essentially a collection of very large and complex molecules formed in part from
lignin and other polyphenolic molecules of the original plant material (foliage, wood, bark), in part from
similar molecules that have been produced by microbes.[18] During decomposition processes these
polyphenols are modified chemically so that they are able to join up with one another to form very large
molecules. Some parts of these molecules are modified in such a way that protein molecules, amino acids,
and amino sugars are able to attach themselves to the polyphenol "base" molecule. As protein contains
both nitrogen and sulfur, this attachment gives humus a moderate content of these two important plant
nutrients.[19]
Radiocarbon and other dating techniques have shown that the polyphenolic base of humus (mostly lignin
and black carbon) can be very old, but the protein and carbohydrate attachments much younger, while to
the light of modern concepts and methods the situation appears much more complex and unpredictable
than previously thought.[20] It seems that microbes are able to pull protein off humus molecules rather more
readily than they are able to break the polyphenolic base molecule itself. As protein is removed its place
may be taken by younger protein, or this younger protein may attach itself to another part of the humus
molecule.[21]
The most useful functions of humus are in improving soil structure, all the more when associated with
cations (e.g. calcium),[22] and in providing a very large surface area that can hold nutrient elements until
required by plants, an ion exchange function comparable to that of clay particles.[23]
Soil carbon sequestration is a major property of the soil, also considered as an ecosystem service.[24] Only
when it becomes stable and acquires its multi-century permanence, mostly via multiple interactions with
the soil matrix, molecular soil humus should be considered to be of significance in removing the
atmosphere's current carbon dioxide overload.[25]
There is little data available on the composition of humus because it is a complex mixture that is
challenging for researchers to analyze. Researchers in the 1940s and 1960s tried using chemical
separation to analyze plant and humic compounds in forest and agricultural soils, but this proved
impossible because extractants interacted with the analysed organic matter and created many artefacts.[26]
Further research has been done in more recent years, though it remains an active field of study.[27]
Humification
Microorganisms decompose a large portion of the soil organic matter into inorganic minerals that the roots
of plants can absorb as nutrients. This process is termed mineralization. In this process, nitrogen (nitrogen
cycle) and the other nutrients (nutrient cycle) in the decomposed organic matter are recycled. Depending on
the conditions in which the decomposition occurs, a fraction of the organic matter does not mineralize and
instead is transformed by a process called humification. Prior to modern analytical methods, early evidence
led scientists to believe that humification resulted in concatenations of organic polymers resistant to the
action of microorganisms,[28] however recent research has demonstrated that microorganisms are capable
of digesting humus.[29]
Humification can occur naturally in soil or artificially in the production of compost. Organic matter is
humified by a combination of saprotrophic fungi, bacteria, microbes and animals such as earthworms,
nematodes, protozoa, and arthropods (see Soil biology). Plant remains, including those that animals
digested and excreted, contain organic compounds: sugars, starches, proteins, carbohydrates, lignins,
waxes, resins, and organic acids. Decay in the soil begins with the decomposition of sugars and starches
from carbohydrates, which decompose easily as detritivores initially invade the dead plant organs, while the
remaining cellulose and lignin decompose more slowly. Simple proteins, organic acids, starches, and
sugars decompose rapidly, while crude proteins, fats, waxes, and resins remain relatively unchanged for
longer periods of time.[30]
Lignin, which is quickly transformed by white-rot fungi,[31] is one of the primary precursors of humus,[32]
together with by-products of microbial[33] and animal[34] activity. The humus produced by humification is
thus a mixture of compounds and complex biological chemicals of plant, animal, and microbial origin that
has many functions and benefits in soil.[18] Some judge earthworm humus (vermicompost) to be the
optimal organic manure.[35]
Stability
Much of the humus in most soils has persisted for more than 100 years, rather than having been
decomposed into CO2, and can be regarded as stable; this organic matter has been protected from
decomposition by microbial or enzyme action because it is hidden (occluded) inside small aggregates of
soil particles, or tightly sorbed or complexed to clays.[36] Most humus that is not protected in this way is
decomposed within 10 years and can be regarded as less stable or more labile.[37] The mixing activity of
soil-consuming invertebrates (e.g. earthworms, termites, some millipedes) contribute to the stability of
humus by favouring the formation of organo-mineral complexes with clay at the inside of their guts,[38][39]
hence more carbon sequestration in humus forms such as mull and amphi, with well-developed mineral-
organic horizons, when compared with moder where most organic matter accumulates at the soil
surface.[40]
Stable humus contributes few plant-available nutrients in soil, but it helps maintain its physical structure.[41]
A very stable form of humus is formed from the slow oxidation (redox) of soil carbon after the
incorporation of finely powdered charcoal into the topsoil, suggested to result from the grinding and mixing
activity of a tropical earthworm.[42] This process is speculated to have been important in the formation of
the unusually fertile Amazonian terra preta do Indio.[43] However, some authors[16] suggest that complex soil
organic molecules may be much less stable than previously thought: "the available evidence does not
support the formation of large-molecular-size and persistent 'humic substances' in soils. Instead, soil
organic matter is a continuum of progressively decomposing organic compounds.″
Horizons
Humus has a characteristic black or dark brown color and is organic due to an accumulation of organic
carbon. Soil scientists use the capital letters O, A, B, C, and E to identify the master soil horizons, and
lowercase letters for distinctions of these horizons. Most soils have three major horizons: the surface
horizon (A), the subsoil (B), and the substratum (C). Some soils have an organic horizon (O) on the surface,
but this horizon can also be buried.[44] The master horizon (E) is used for subsurface horizons that have
significantly lost minerals (eluviation). Bedrock, which is not soil, uses the letter R. The richness of soil
horizons in humus determines their more or less dark color, generally decreasing from O to E, to the
exception of deep horizons of podzolic soils enriched with colloidal humic substances which have been
leached down the soil profile.[45]
The importance of chemically stable humus is thought by some to be the fertility it provides to soils in both
a physical and chemical sense,[46] though some agricultural experts put a greater focus on other features of
it, such as its ability to suppress disease.[47] It helps the soil retain moisture[48] by increasing
microporosity[49] and encourages the formation of good soil structure.[50][51] The incorporation of oxygen
into large organic molecular assemblages generates many active, negatively charged sites that bind to
positively charged ions (cations) of plant nutrients, making them more available to the plant by way of ion
exchange.[52] Humus allows soil organisms to feed and reproduce and is often described as the "life-force"
of the soil.[53][54]
The process that converts soil organic matter into humus feeds the population of microorganisms and
other creatures in the soil, and thus maintains high and healthy levels of soil life.[53][54]
The rate at which soil organic matter is converted into humus promotes (when fast, e.g. mull) or limits
(when slow, e.g. mor) the coexistence of plants, animals, and microorganisms in the soil.[55]
"Effective humus" and "stable humus" are additional sources of nutrients for microbes: the former
provides a readily available supply, and the latter acts as a long-term storage reservoir.[56]
Decomposition of dead plant material causes complex organic compounds to be slowly oxidized (lignin-
like humus) or to decompose into simpler forms (sugars and amino sugars, and aliphatic and phenolic
organic acids), which are further transformed into microbial biomass (microbial humus) or reorganized
and further oxidized into humic assemblages (fulvic acids and humic acids), which bind to clay minerals
and metal hydroxides.[57] The ability of plants to absorb humic substances with their roots and
metabolize them has been long debated.[58] There is now a consensus that humus functions hormonally
rather than simply nutritionally in plant physiology,[59][60] and that organic sunstances exuded by roots
and transformed in humus by soil organisms are an evolved strategy by which plants "talk" to the soil.[61]
Humus is a negatively charged colloidal substance which increases the cation-exchange capacity of soil,
hence its ability to store nutrients by chelation.[62] While these nutrient cations are available to plants,
they are held in the soil and prevented from being leached by rain or irrigation.[52]
Humus can hold the equivalent of 80–90% of its weight in moisture and therefore increases the soil's
capacity to withstand drought.[63]
The biochemical structure of humus enables it to moderate, i.e. buffer, excessive acidic or alkaline soil
conditions.[64]
During humification, microbes secrete sticky, gum-like mucilages; these contribute to the crumby
structure (tilth) of the soil by adhering particles together and allowing greater aeration of the soil.[65]
Toxic substances such as heavy metals and excess nutrients can be chelated, i.e., bound to the organic
molecules of humus, and so prevented from leaching away.[66]
The dark, usually brown or black, color of humus helps to warm cold soils in spring.[67]
Humus can contribute to climate change mitigation through its carbon sequestration potential.[68]
Artificial humic acid and artificial fulvic acid synthesized from agricultural litter can increase the content
of dissolved organic matter and total organic carbon in soil.[69]
See also
Biochar
Biomass
Biotic material
Detritus
Glomalin
Humic acid
Organic matter
Plant litter
Soil horizon
Soil science
Terra preta
References
1. Popkin, Gabriel (27 July 2021), A soil-science revolution upends plans to fight climate change (https://w
ww.quantamagazine.org/a-soil-science-revolution-upends-plans-to-fight-climate-change-20210727/) ,
Quanta Magazine, retrieved 9 June 2024, " "The latest edition of The Nature and Properties of Soils,
published in 2016, cites Lehmann's 2015 paper and acknowledges that "our understanding of the
nature and genesis of soil humus has advanced greatly since the turn of the century, requiring that
some long-accepted concepts be revised or abandoned"."
4. Chertov, Oleg G.; Komarov, Alexander S.; Crocker, Graham; Grace, Peter; Klir, Jan; Körschens, Martin;
Poulton, Paul R.; Richter, Daniel (1997). "Simulating trends of soil organic carbon in seven long-term
experiments using the SOMM model of the humus types" (https://fr.articles.sk/book/17536355/c3552
3) . Geoderma. 81 (1–2): 121–135. Bibcode:1997Geode..81..121C (https://ui.adsabs.harvard.edu/ab
s/1997Geode..81..121C) . doi:10.1016/S0016-7061(97)00085-2 (https://doi.org/10.1016%2FS0016-7
061%2897%2900085-2) . Retrieved 9 June 2024.
5. Brêthes, Alain; Brun, Jean-Jacques; Jabiol, Bernard; Ponge, Jean-François; Toutain, François (1995).
"Classification of forest humus forms: a French proposal" (https://www.researchgate.net/publication/
45341270) . Annales des Sciences Forestières. 52 (6): 535–46. doi:10.1051/forest:19950602 (https://
doi.org/10.1051%2Fforest%3A19950602) . Retrieved 16 June 2024.
6. Bernier, Nicolas (1998). "Earthworm feeding activity and development of the humus profile" (https://w
ww.academia.edu/34816078) . Biology and Fertility of Soils. 26 (3): 215–23.
Bibcode:1998BioFS..26..215B (https://ui.adsabs.harvard.edu/abs/1998BioFS..26..215B) .
doi:10.1007/s003740050370 (https://doi.org/10.1007%2Fs003740050370) . Retrieved 16 June
2024.
8. Bauer, Armand (1974). "Influence of soil organic matter on bulk density and available water capacity of
soils" (https://library.ndsu.edu/ir/bitstream/handle/10365/24299/ndfr_19740501_v31_iss05_044.pd
f) (PDF). Farm Research. 31 (5): 44–52. Retrieved 23 June 2024.
9. Whitehead, D. C.; Tinsley, J. (1963). "The biochemistry of humus formation" (https://fr.articles.sk/boo
k/1689524/861585) . Journal of the Science of Food and Agriculture. 14 (12): 849–57.
Bibcode:1963JSFA...14..849W (https://ui.adsabs.harvard.edu/abs/1963JSFA...14..849W) .
doi:10.1002/jsfa.2740141201 (https://doi.org/10.1002%2Fjsfa.2740141201) . Retrieved 23 June
2024.
11. Brinton, William F. (2020). "Compost quality standards and guidelines, final report" (https://compost.cs
s.cornell.edu/Brinton.pdf) (PDF). Ithaca, New York: Cornell University. Retrieved 7 July 2024.
12. Guggenberger, Georg (2005). "Humification and mineralization in soils". In Buscot, François; Varma,
Ajit (eds.). Microorganisms in soils: roles in genesis and Functions (http://ndl.ethernet.edu.et/bitstrea
m/123456789/75774/1/Franc%C2%B8ois%20Buscot.pdf#page=102) (PDF). Soil biology. Vol. 3.
Dordrecht, The Netherlands: Springer. pp. 85–106. doi:10.1007/3-540-26609-7_4 (https://doi.org/10.1
007%2F3-540-26609-7_4) . ISBN 978-3-540-26609-9. Archived (https://web.archive.org/web/202407
07083204/http://ndl.ethernet.edu.et/bitstream/123456789/75774/1/Franc%C2%B8ois%20Buscot.pdf
#page=102) (PDF) from the original on 7 July 2024. Retrieved 8 September 2024.
13. Kögel-Knabner, Ingrid; Zech, Wolfgang; Hatcher, Patrick G. (1988). "Chemical composition of the
organic matter in forest soils: the humus layer" (https://fr.articles.sk/book/34689934/0bcef3) .
Journal of Plant Nutrition and Soil Science. 151 (5): 331–40. doi:10.1002/jpln.19881510512 (https://do
i.org/10.1002%2Fjpln.19881510512) . Retrieved 14 July 2024.
14. Waksman, Selman A. (1936). Humus: origin, chemical composition and importance in nature (https://cit
eseerx.ist.psu.edu/document?repid=rep1&type=pdf&doi=45020e04c07d0fa28dca0093772951b65197
eb2e) . Baltimore, Maryland: Williams & Wilkins. ISBN 9780598966629. Retrieved 14 July 2024.
15. Bernier, Nicolas; Ponge, Jean-François (1994). "Humus form dynamics during the sylvogenetic cycle in
a mountain spruce forest" (https://www.researchgate.net/publication/46312511) . Soil Biology and
Biochemistry. 26 (2): 183–220. Bibcode:1994SBiBi..26..183B (https://ui.adsabs.harvard.edu/abs/1994
SBiBi..26..183B) . doi:10.1016/0038-0717(94)90161-9 (https://doi.org/10.1016%2F0038-0717%289
4%2990161-9) . Retrieved 14 July 2024.
16. Lehmann, Johannes; Kleber, Markus (2015). "The contentious nature of soil organic matter" (https://th
emarea.org/wp-content/uploads/2018/08/Lehmann-and-Kebbler-2015.pdf) (PDF). Nature. 528
(7580): 60–68. Bibcode:2015Natur.528...60L (https://ui.adsabs.harvard.edu/abs/2015Natur.528...60
L) . doi:10.1038/nature16069 (https://doi.org/10.1038%2Fnature16069) . PMID 26595271 (https://p
ubmed.ncbi.nlm.nih.gov/26595271) . Retrieved 14 July 2024.
17. Ponge, Jean-François (2022). "Humus: dark side of life or intractable "aether"?" (https://www.research
gate.net/publication/360175852) . Pedosphere. 32 (4): 660–64. Bibcode:2022Pedos..32..660P (http
s://ui.adsabs.harvard.edu/abs/2022Pedos..32..660P) . doi:10.1016/S1002-0160(21)60013-9 (https://
doi.org/10.1016%2FS1002-0160%2821%2960013-9) . Retrieved 14 July 2024.
18. Dou, Sen; Shan, Jun; Song, Xiangyun; Cao, Rui; Wu, Meng; Li, Chenglin; Guan, Song (April 2020). "Are
humic substances soil microbial residues or unique synthesized compounds? A perspective on their
distinctiveness" (https://www.researchgate.net/publication/338991840) . Pedosphere. 30 (2): 159–
67. Bibcode:2020Pedos..30..159D (https://ui.adsabs.harvard.edu/abs/2020Pedos..30..159D) .
doi:10.1016/S1002-0160(20)60001-7 (https://doi.org/10.1016%2FS1002-0160%2820%2960001-7) .
Retrieved 21 July 2024.
19. Das, Subhasich; Bhattacharya, Satya Sundar (2017). "Significance of soil organic matter in relation to
plants and their products". In Siddiqui, Mohammed Wasim; Bansal, Vasudha (eds.). Plant secondary
metabolites. Volume 3. Their roles in stress ecophysiology (https://www.academia.edu/82083954) .
Palm Bay, Florida: Apple Academic Press. pp. 39–61. ISBN 978-1-77188-356-6. Retrieved 26 August
2024.
20. Piccolo, Alessandro (December 2002). "The supramolecular structure of humic substances: a novel
understanding of humus chemistry and implications in soil science" (https://www.researchgate.net/pu
blication/222526145) . Advances in Agronomy. 75: 57–134. doi:10.1016/S0065-2113(02)75003-7 (htt
ps://doi.org/10.1016%2FS0065-2113%2802%2975003-7) . ISBN 978-0-12-000793-6. Retrieved
4 August 2024.
21. Paul, Eldor A. (2016). "The nature and dynamics of soil organic matter: plant inputs, microbial
transformations, and organic matter stabilization" (https://www.nrel.colostate.edu/assets/nrel_files/la
bs/paul-lab/docs/Paul_SBBreview2016.pdf) (PDF). Soil Biology and Biochemistry. 98: 109–26.
Bibcode:2016SBiBi..98..109P (https://ui.adsabs.harvard.edu/abs/2016SBiBi..98..109P) .
doi:10.1016/j.soilbio.2016.04.001 (https://doi.org/10.1016%2Fj.soilbio.2016.04.001) . Retrieved
11 August 2024.
22. Huang, Xue Ru; Li, H.; Li, Song; Xiong, Hailing; Jiang, Xianjun (May 2016). "Role of cationic polarization
in humus-increased soil aggregate stability" (https://www.researchgate.net/publication/303509978) .
European Journal of Soil Science. 67 (3): 341–50. Bibcode:2016EuJSS..67..341H (https://ui.adsabs.har
vard.edu/abs/2016EuJSS..67..341H) . doi:10.1111/ejss.12342 (https://doi.org/10.1111%2Fejss.1234
2) . Retrieved 11 August 2024.
23. Shoba, V. N.; Chudnenko, K. V. (August 2014). "Ion exchange properties of humus acids" (https://www.r
esearchgate.net/publication/269385340) . Eurasian Soil Science. 47 (8): 761–71.
Bibcode:2014EurSS..47..761S (https://ui.adsabs.harvard.edu/abs/2014EurSS..47..761S) .
doi:10.1134/S1064229314080110 (https://doi.org/10.1134%2FS1064229314080110) . Retrieved
11 August 2024.
24. Lal, Rattan; Negassa, Wakene; Lorenz, Klaus (August 2015). "Carbon sequestration in soil" (https://ww
w.researchgate.net/publication/283457192) . Current Opinion in Environmental Sustainability. 15: 79–
86. Bibcode:2015COES...15...79L (https://ui.adsabs.harvard.edu/abs/2015COES...15...79L) .
doi:10.1016/j.cosust.2015.09.002 (https://doi.org/10.1016%2Fj.cosust.2015.09.002) . Retrieved
18 August 2024.
25. Dynarski, Katherine A.; Bossio, Deborah A.; Scow, Kate M. (13 November 2020). "Dynamic stability of
soil carbon: reassessing the "permanence" of soil carbon sequestration" (https://doi.org/10.3389%2Ff
envs.2020.514701) . Frontiers in Environmental Science. 8 (714701). doi:10.3389/fenvs.2020.514701
(https://doi.org/10.3389%2Ffenvs.2020.514701) .
26. Kleber, Markus; Lehmann, Johannes (8 March 2019). "Humic substances extracted by alkali are invalid
proxies for the dynamics and functions of organic matter in terrestrial and aquatic ecosystems" (http
s://acsess.onlinelibrary.wiley.com/doi/pdfdirect/10.2134/jeq2019.01.0036) . Journal of
Environmental Quality. 48 (2): 207–16. Bibcode:2019JEnvQ..48..207K (https://ui.adsabs.harvard.edu/a
bs/2019JEnvQ..48..207K) . doi:10.2134/jeq2019.01.0036 (https://doi.org/10.2134%2Fjeq2019.01.00
36) . PMID 30951127 (https://pubmed.ncbi.nlm.nih.gov/30951127) . Retrieved 25 August 2024.
27. Baveye, Philippe C.; Wander, Michelle (6 March 2019). "The (bio)chemistry of soil humus and humic
substances: why is the "new view" still considered novel after more than 80 years?" (https://doi.org/10.
3389%2Ffenvs.2019.00027) . Frontiers in Environmental Science. 7 (27).
doi:10.3389/fenvs.2019.00027 (https://doi.org/10.3389%2Ffenvs.2019.00027) .
28. Brady, Nyle C. (1984). The nature and properties of soils (https://www.academia.edu/23641831)
(9th ed.). New York, New York: Macmillan Publishing Company. p. 265. ISBN 978-0029460306.
Retrieved 1 September 2024.
29. Popkin, Gabriel (2021). "A soil-science revolution upends plans to fight climate change" (https://www.q
uantamagazine.org/a-soil-science-revolution-upends-plans-to-fight-climate-change-20210727/) .
Quanta Magazine. Retrieved 1 September 2024. "Soil researchers have concluded that even the largest,
most complex molecules can be quickly devoured by soil's abundant and voracious microbes"
30. Krishna, M. P.; Mohan, Mahesh (July 2017). "Litter decomposition in forest ecosystems: a review" (http
s://www.academia.edu/119720860) . Energy, Ecology and Environment. 2 (3): 236–49.
doi:10.1007/s40974-017-0064-9 (https://doi.org/10.1007%2Fs40974-017-0064-9) . Retrieved
8 September 2024.
31. Levin, Laura; Forchiassin, Flavia (9 May 2001). "Ligninolytic enzymes of the white rot basidiomycete
Trametes trogii" (https://www.academia.edu/120239930) . Acta Biotechnologica. 21 (2): 179–86.
doi:10.1002/1521-3846(200105)21:2<179::AID-ABIO179>3.0.CO;2-2 (https://doi.org/10.1002%2F1521-
3846%28200105%2921%3A2%3C179%3A%3AAID-ABIO179%3E3.0.CO%3B2-2) . Retrieved
15 September 2024.
32. González-Pérez, Martha; Vidal Torrado, Pablo; Colnago, Luiz A.; Martin-Neto, Ladislau; Otero, Xosé L.;
Milori, Débora M. B. P.; Haenel Gomes, Felipe (31 August 2008). "13C NMR and FTIR spectroscopy
characterization of humic acids in spodosols under tropical rain forest in southeastern Brazil" (https://
www.academia.edu/14026276) . Geoderma. 146 (3–4): 425–33. Bibcode:2008Geode.146..425G (htt
ps://ui.adsabs.harvard.edu/abs/2008Geode.146..425G) . doi:10.1016/j.geoderma.2008.06.018 (http
s://doi.org/10.1016%2Fj.geoderma.2008.06.018) . Retrieved 15 September 2024.
33. Knicker, Heike; Almendros, Gonzalo; González-Vila, Francisco Javier; Lüdemann, Hans-Dietrich; Martín,
Fracisco (November–December 1995). "13C and 15N NMR analysis of some fungal melanins in
comparison with soil organic matter" (https://www.academia.edu/78009567) . Organic Geochemistry.
23 (11–12): 1023–28. Bibcode:1995OrGeo..23.1023K (https://ui.adsabs.harvard.edu/abs/1995OrGe
o..23.1023K) . doi:10.1016/0146-6380(95)00094-1 (https://doi.org/10.1016%2F0146-6380%2895%29
00094-1) . Retrieved 15 September 2024.
34. Muscolo, Adele; Bovalo, Francesco; Gionfriddo, Francesco; Nardi, Serenella (August 1999). "Earthworm
humic matter produces auxin-like effects on Daucus carota cell growth and nitrate metabolism" (http
s://www.academia.edu/78825632) . Soil Biology and Biochemistry. 31 (9): 1303–11.
Bibcode:1999SBiBi..31.1303M (https://ui.adsabs.harvard.edu/abs/1999SBiBi..31.1303M) .
doi:10.1016/S0038-0717(99)00049-8 (https://doi.org/10.1016%2FS0038-0717%2899%2900049-8) .
Retrieved 15 September 2024.
35. Oyege, Ivan; Sridhar, B. B. Maruthi (10 November 2023). "Effects of vermicompost on soil and plant
health and promoting sustainable agriculture" (https://doi.org/10.3390%2Fsoilsystems7040101) .
Soil Systems. 7 (4): 101. doi:10.3390/soilsystems7040101 (https://doi.org/10.3390%2Fsoilsystems70
40101) .
36. Dungait, J. A.; Hopkins, D. W.; Gregory, A. S.; Whitmore, A. P. (14 February 2012). "Soil organic matter
turnover is governed by accessibility not recalcitrance" (https://www.desmog.com/wp-content/upload
s/files/Dungait%20SOM%20article.pdf) (PDF). Global Change Biology. 18 (6): 1781–96.
doi:10.1111/j.1365-2486.2012.02665.x (https://doi.org/10.1111%2Fj.1365-2486.2012.02665.x) .
Retrieved 22 September 2024.
37. Baldock, Jeffrey A.; Skjemstad, Jan Otto (July 2000). "Role of the soil matrix and minerals in protecting
natural organic materials against biological attack" (https://www.academia.edu/78009563) . Organic
Geochemistry. 31 (7): 697–710. doi:10.1016/S0146-6380(00)00049-8 (https://doi.org/10.1016%2FS01
46-6380%2800%2900049-8) . Retrieved 22 September 2024.
38. Angst, Šárka; Mueller, Carsten W.; Cajthaml, Tomáš; Angst, Gerrit; Lhotáková, Zuzana; Bartuška, Martin;
Špaldoňová, Alexandra; Frouz, Jan (1 March 2017). "Stabilization of soil organic matter by earthworms
is connected with physical protection rather than with chemical changes of organic matter" (https://w
ww.academia.edu/80259832) . Geoderma. 289: 29–35. doi:10.1016/j.geoderma.2016.11.017 (http
s://doi.org/10.1016%2Fj.geoderma.2016.11.017) . Retrieved 6 October 2024.
39. Brauman, Alain (July 2000). "Effect of gut transit and mound deposit on soil organic matter
transformations in the soil feeding termite: a review" (https://fr.articles.sk/book/18221151/100de8) .
European Journal of Soil Biology. 36 (3–4): 117–25. doi:10.1016/S1164-5563(00)01058-X (https://doi.
org/10.1016%2FS1164-5563%2800%2901058-X) . Retrieved 6 October 2024.
40. Andreetta, Anna; Ciampalini, Rossano; Moretti, Pierpaolo; Vingiani, Simona; Poggio, Giorgio; Matteucci,
Giorgio; Tescari, Francesca; Carnicelli, Stefano (2011). "Forest humus forms as potential indicators of
soil carbon storage in Mediterranean environments" (https://www.researchgate.net/publication/22641
7489) . Biology and Fertility of Soils. 47: 31–40. doi:10.1007/s00374-010-0499-z (https://doi.org/10.1
007%2Fs00374-010-0499-z) . Retrieved 6 October 2024.
41. Oades, J. Malcolm (February 1984). "Soil organic matter and structural stability: mechanisms and
implications for management" (https://edisciplinas.usp.br/pluginfile.php/5168021/mod_resource/con
tent/1/Grupo%206_Oades%2C%201984.%20Soil%20organic%20matter%20and%20structural%20stabil
ity%20mechanisms%20and%20implications%20for%20management.pdf) (PDF). Plant and Soil. 76
(1–3): 319–337. Bibcode:1984PlSoi..76..319O (https://ui.adsabs.harvard.edu/abs/1984PlSoi..76..319
O) . doi:10.1007/BF02205590 (https://doi.org/10.1007%2FBF02205590) . S2CID 7195036 (https://a
pi.semanticscholar.org/CorpusID:7195036) . Retrieved 13 October 2024.
42. Ponge, Jean-François; Topoliantz, Stéphanie; Ballof, Sylvain; Rossi, Jean-Pierre; Lavelle, Patrick;
Betsch, Jean-Marie; Gaucher, Philippe (July 2006). "Ingestion of charcoal by the Amazonian
earthworm Pontoscolex corethrurus: a potential for tropical soil fertility" (https://www.researchgate.ne
t/publication/44735820) . Soil Biology and Biochemistry. 38 (7): 2008–9.
doi:10.1016/j.soilbio.2005.12.024 (https://doi.org/10.1016%2Fj.soilbio.2005.12.024) . Retrieved
13 October 2024.
43. Arroyo-Kalin, Manuel (July 2017). "Amazonian Dark Earths". In Nicosia, Cristiano; Stoops, Georges
(eds.). Archaeological soil and sediment micromorphology (https://www.researchgate.net/publication/3
19444794) . Hoboken, New Jersey: Wiley. pp. 345–57. doi:10.1002/9781118941065.ch33 (https://do
i.org/10.1002%2F9781118941065.ch33) . ISBN 9781118941065. Retrieved 13 October 2024.
44. Gerlach, Renate; Fischer, Peter; Eckmeier, Eileen; Hilgers, Alexandra. "Buried dark soil horizons and
archaeological features in the Neolithic settlement region of the Lower Rhine area, NW Germany:
formation, geochemistry and chronostratigraphy" (https://www.academia.edu/77065464) .
Quaternary International. 265: 191–204. doi:10.1016/j.quaint.2011.10.007 (https://doi.org/10.1016%2F
j.quaint.2011.10.007) . Retrieved 20 October 2024.
45. Sanborn, Paul; Lamontagne, Luc; Hendershot, William. "Podzolic soils of Canada: genesis, distribution,
and classification" (https://doi.org/10.4141%2Fcjss10024) . Canadian Journal of Soil Science. 91 (5):
843–80. doi:10.4141/cjss10024 (https://doi.org/10.4141%2Fcjss10024) .
46. Hargitai, László (December 1993). "The role of organic matter content and humus quality in the
maintenance of soil fertility and in environmental protection" (https://fr.articles.sk/book/19958903/07
e3a6) . Landscape and Urban Planning. 27 (2–4): 161–67. Bibcode:1993LUrbP..27..161H (https://ui.ad
sabs.harvard.edu/abs/1993LUrbP..27..161H) . doi:10.1016/0169-2046(93)90044-E (https://doi.org/1
0.1016%2F0169-2046%2893%2990044-E) . Retrieved 27 October 2024.
47. Hoitink, Harry A. J.; Fahy, Peter C. (September 1986). "Basis for the control of soilborne plant
pathogens with composts" (https://fr.articles.sk/book/18744629/4076fa) . Annual Review of
Phytopathology. 24: 93–114. doi:10.1146/annurev.py.24.090186.000521 (https://doi.org/10.1146%2Fa
nnurev.py.24.090186.000521) . Retrieved 27 October 2024.
48. Lal, Rattan (September 2020). "Soil organic matter and water retention" (https://www.researchgate.ne
t/publication/341213360) . Agronomy Journal. 116 (5): 3265–77. doi:10.1002/agj2.20282 (https://do
i.org/10.1002%2Fagj2.20282) . Retrieved 27 October 2024.
49. de Macedo, José Ronaldo; do Amaral Meneguelli, Neli; Ottoni Filho, Theophilo Benedicto; Lima, Jorge
Araújo de Sousa (February 2007). "Estimation of field capacity and moisture retention based on
regression analysis involving chemical and physical properties in Alfisols and Ultisols of the state of
Rio de Janeiro" (https://fr.articles.sk/book/32902177/18be8b) . Communications in Soil Science and
Plant Analysis. 33 (13–14): 2037–55. Bibcode:2002CSSPA..33.2037D (https://ui.adsabs.harvard.edu/a
bs/2002CSSPA..33.2037D) . doi:10.1081/CSS-120005747 (https://doi.org/10.1081%2FCSS-1200057
47) . S2CID 98466747 (https://api.semanticscholar.org/CorpusID:98466747) . Retrieved 27 October
2024.
50. Hempfling, Reinhold; Schulten, Hans-Rolf; Horn, Rainer (June 1990). "Relevance of humus composition
to the physical/mechanical stability of agricultural soils: a study by direct pyrolysis-mass
spectrometry" (https://fr.articles.sk/book/8543777/01704b) . Journal of Analytical and Applied
Pyrolysis. 17 (3): 275–81. doi:10.1016/0165-2370(90)85016-G (https://doi.org/10.1016%2F0165-237
0%2890%2985016-G) . Retrieved 3 November 2024.
51. Piccolo, Alessandro (1996). "Humus and soil conservation". In Piccolo, Alessandro (ed.). Humic
substances in terrestrial ecosystems (https://www.researchgate.net/publication/281451183) .
Amsterdam, The Netherlands: Elsevier. pp. 225–64. doi:10.1016/B978-044481516-3/50006-2 (https://
doi.org/10.1016%2FB978-044481516-3%2F50006-2) . ISBN 978-0-444-81516-3. Retrieved
3 November 2024.
52. Szalay, Alex (October–November 1964). "Cation exchange properties of humic acids and their
importance in the geochemical enrichment of UO2++ and other cations" (https://fr.articles.sk/book/19
639940/3bac09) . Geochimica et Cosmochimica Acta. 28 (10–11): 1605–14.
Bibcode:1964GeCoA..28.1605S (https://ui.adsabs.harvard.edu/abs/1964GeCoA..28.1605S) .
doi:10.1016/0016-7037(64)90009-2 (https://doi.org/10.1016%2F0016-7037%2864%2990009-2) .
Retrieved 3 November 2024.
53. Elo, Seija; Maunuksela, Liisa; Salkinoja-Salonen, Mirja; Smolander, Aino; Haahtela, Kielo (February
2000). "Humus bacteria of Norway spruce stands: plant growth promoting properties and birch, red
fescue and alder colonizing capacity" (https://doi.org/10.1111%2Fj.1574-6941.2000.tb00679.x) .
FEMS Microbiology Ecology. 31 (2): 143–52. doi:10.1111/j.1574-6941.2000.tb00679.x (https://doi.org/
10.1111%2Fj.1574-6941.2000.tb00679.x) . PMID 10640667 (https://pubmed.ncbi.nlm.nih.gov/10640
667) .
54. Vreeken-Buijs, Madelein J.; Hassink, Jan; Brussaard, Lijbert (1998). "Relationships of soil
microarthropod biomass with organic matter and pore size distribution in soils under different land
use" (https://www.academia.edu/65368490) . Soil Biology and Biochemistry. 30 (1): 97–106.
Bibcode:1998SBiBi..30...97V (https://ui.adsabs.harvard.edu/abs/1998SBiBi..30...97V) .
doi:10.1016/S0038-0717(97)00064-3 (https://doi.org/10.1016%2FS0038-0717%2897%2900064-3) .
Retrieved 3 November 2024.
55. Ponge, Jean-François (July 2003). "Humus forms in terrestrial ecosystems: a framework to
biodiversity" (https://www.academia.edu/20508983) . Soil Biology and Biochemistry. 35 (7): 935–45.
doi:10.1016/S0038-0717(03)00149-4 (https://doi.org/10.1016%2FS0038-0717%2803%2900149-4) .
Retrieved 10 November 2024.
56. Hodges, R. D. (1991). "Soil organic matter: its central position in organic farming". In Wilson, W. S.
(ed.). Advances in soil organic matter research: the impact on agriculture and the environment (https://fr.
articles.sk/book/81383430/4b29a3) . Sawston, United Kingdom: Woodhead Publishing. pp. 355–64.
doi:10.1016/b978-1-85573-813-3.50040-8 (https://doi.org/10.1016%2Fb978-1-85573-813-3.50040-
8) . ISBN 978-1-85573-813-3. Retrieved 10 November 2024.
57. Gunina, Anna; Kuzyakov, Yakov (April 2022). "From energy to (soil organic) matter" (https://doi.org/10.
1111%2Fgcb.16071) . Global Change Biology. 28 (7): 2169–82. doi:10.1111/gcb.16071 (https://doi.or
g/10.1111%2Fgcb.16071) .
58. Senn, T. L.; Kingman, Alta R.; Godley, W. C. (1973). "A review of humus and humic acids" (https://www.h
umintech.com/fileadmin/content_images/agriculture/information/articles_pdf/A-Review-of-Humus-an
d-Humic-Acids_T.L.Senn__A.R.Kingsmann.pdf) (PDF). Research Series, South Carolina Agricultural
Experiment Station. 145. Retrieved 24 November 2024.
59. Eyheraguibel, Boris; Silvestre, Jérôme; Morard, Philippe (July 2008). "Effects of humic substances
derived from organic waste enhancement on the growth and mineral nutrition of maize" (https://hal.sci
ence/hal-00940093/file/Eyheraguibel_10804.pdf) (PDF). Bioresource Technology. 99 (10): 4206–12.
Bibcode:2008BiTec..99.4206E (https://ui.adsabs.harvard.edu/abs/2008BiTec..99.4206E) .
doi:10.1016/j.biortech.2007.08.082 (https://doi.org/10.1016%2Fj.biortech.2007.08.082) .
PMID 17962015 (https://pubmed.ncbi.nlm.nih.gov/17962015) . Retrieved 17 November 2024.
60. Zandonadi, Daniel Basilio; Santos, Mirella Pupo; Busato, Jader Galba; Peres, Lázaro Eustáquio Pereira;
Façanha, Arnoldo Rocha (2013). "Plant physiology as affected by humified organic matter" (https://doi.
org/10.1590%2FS2197-00252013000100003) . Theoretical and Experimental Plant Physiology. 25 (1):
12–25. doi:10.1590/S2197-00252013000100003 (https://doi.org/10.1590%2FS2197-0025201300010
0003) .
61. Nardi, Serenella; Ertani, Andrea; Francioso, Ornella (February 2017). "Soil–root cross-talking: the role of
humic substances" (https://www.academia.edu/102119488) . Journal of Plant Nutrition and Soil
Science. 180 (1): 5–13. doi:10.1002/jpln.201600348 (https://doi.org/10.1002%2Fjpln.201600348) .
hdl:2318/1731194 (https://hdl.handle.net/2318%2F1731194) . Retrieved 17 November 2024.
62. Shoba, V. N.; Chudnenko, Konstantin V. (August 2014). "Ion exchange properties of humus acids" (http
s://www.researchgate.net/publication/269385340) . Eurasian Soil Science. 47 (8): 761–71.
doi:10.1134/S1064229314080110 (https://doi.org/10.1134%2FS1064229314080110) . Retrieved
24 November 2024.
63. Olness, Alan; Archer, David (February 2005). "Effect of organic carbon on available water in soil" (http
s://fr.articles.sk/book/54264627/14eda9) . Soil Science. 170 (2): 90–101.
Bibcode:2005SoilS.170...90O (https://ui.adsabs.harvard.edu/abs/2005SoilS.170...90O) .
doi:10.1097/00010694-200502000-00002 (https://doi.org/10.1097%2F00010694-200502000-0000
2) . S2CID 95336837 (https://api.semanticscholar.org/CorpusID:95336837) . Retrieved
24 November 2024.
64. Kikuchi, Ryunosuke (February 2004). "Deacidification effect of the litter layer on forest soil during
snowmelt runoff: laboratory experiment and its basic formularization for simulation modeling" (http
s://fr.articles.sk/book/16655436/acdc92) . Chemosphere. 54 (8): 1163–69.
Bibcode:2004Chmsp..54.1163K (https://ui.adsabs.harvard.edu/abs/2004Chmsp..54.1163K) .
doi:10.1016/j.chemosphere.2003.10.025 (https://doi.org/10.1016%2Fj.chemosphere.2003.10.025) .
PMID 14664845 (https://pubmed.ncbi.nlm.nih.gov/14664845) . Retrieved 24 November 2024.
65. Caesar-Tonthat, Thecan C. (August 2002). "Soil binding properties of mucilage produced by a
basidiomycete fungus in a model system" (https://fr.articles.sk/book/20537646/09bc94) .
Mycological Research. 106 (8): 930–37. doi:10.1017/S0953756202006330 (https://doi.org/10.1017%2
FS0953756202006330) . Retrieved 24 November 2024.
66. Zhu, Rui; Wu, Min; Yang, Jian (February 2011). "Mobilities and leachabilities of heavy metals in sludge
with humus soil" (https://fr.articles.sk/book/14270592/0155c3) . Journal of Environmental Sciences.
23 (2): 247–54. doi:10.1016/S1001-0742(10)60399-3 (https://doi.org/10.1016%2FS1001-0742%281
0%2960399-3) . Retrieved 24 November 2024.
67. Ludwig, J. W.; Harper, John L. (July 1958). "The influence of the environment on seed and seedling
mortality. VIII. The influence of soil colour" (https://fr.articles.sk/book/57099612/546f22) . Journal of
Ecology. 46 (2): 381–89. doi:10.2307/2257402 (https://doi.org/10.2307%2F2257402) . Retrieved
1 December 2024.
68. Amelung, Wulf; Bossio, Deborah; De Vries, Wim; Kögel-Knabner, Ingrid; Lehmann, Johannes;
Amundson, Ronald; Bol, Roland; Collins, Chris; Lal, Rattan; Leifeld, Jens; Minasny, Budiman; Pan, Gen-
Xing; Paustian, Keith; Rumpel, Cornelia; Sanderman, Jonathan; Van Groenigen, Jan Willem; Mooney,
Sacha; Van Wesemael, Bas; Wander, Michelle; Chabbi, Abbad (27 October 2020). "Towards a global-
scale soil climate mitigation strategy" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7591914) .
Nature Communications. 11 (1): 5427. Bibcode:2020NatCo..11.5427A (https://ui.adsabs.harvard.edu/a
bs/2020NatCo..11.5427A) . doi:10.1038/s41467-020-18887-7 (https://doi.org/10.1038%2Fs41467-0
20-18887-7) . ISSN 2041-1723 (https://search.worldcat.org/issn/2041-1723) . PMC 7591914 (http
s://www.ncbi.nlm.nih.gov/pmc/articles/PMC7591914) . PMID 33110065 (https://pubmed.ncbi.nlm.ni
h.gov/33110065) .
69. Tang, Chunyu; Li, Yuelei; Song, Jingpeng; Antonietti, Markus; Yang, Fan (25 June 2021). "Artificial
humic substances improve microbial activity for binding CO2" (https://www.ncbi.nlm.nih.gov/pmc/arti
cles/PMC8387571) . iScience. 24 (6): 102647. Bibcode:2021iSci...24j2647T (https://ui.adsabs.harvar
d.edu/abs/2021iSci...24j2647T) . doi:10.1016/j.isci.2021.102647 (https://doi.org/10.1016%2Fj.isci.2
021.102647) . ISSN 2589-0042 (https://search.worldcat.org/issn/2589-0042) . PMC 8387571 (http
s://www.ncbi.nlm.nih.gov/pmc/articles/PMC8387571) . PMID 34466779 (https://pubmed.ncbi.nlm.ni
h.gov/34466779) .
External links