DID THE ALKALINE GUT OF TERMITES CREATE LATOSOLS (ULTISOLS and OXISOLS), BAUXITES, AND LATERITES?

by Charles Weber, MS

ABSTRACT

It is proposed here that the laterization of tropical soils, whereby silica is leached out to leave iron and aluminum hydroxides behind, is caused by a high alkalinity past the mid gut of soil eating termites. It is suggested that this alkalinity first arose primarily in order to make phosphate available to the termites and it resulted in laterites, bauxites, glauconite, and diatom rise especially in early Cretaceous

DISCUSSION

The humus eating termites, which make up over half of the termite species [Brune & Kuhl, 1996], must have to solve the problem of binding of phosphorus by iron and aluminum in soil. They may have done this by creating an alkaline medium in their gut. The humus eating termites create an alkaline pH of 11 to 12.5 just past the mid gut-hindgut junction (the P1 segment) [Bignell, 1998 (there are diagrams of the various gut designs of termites here)]. It is said that this is the highest pH in the biological world. It was obtained using micro electrodes, so is probably accurate [Brune & Kuhl, 1996]. This is 100,000 times as many hydroxyl ions as in a neutral solution, the equivalent of a solution of potassium lye. To put this in perspective, a tenth normal solution of potassium carbonate has a pH of 11.6, a tenth normal solution of potassium silicate has a pH of 12.6, saturated lime (calcium hydroxide) has a pH of 12.4, and a one hundredth normal solution of potassium hydroxide (potassium lye) has a pH of 12. A 200 mM (one fifth normal) solution of potassium carbonate mimics the pH of the gut's P1 region [Kappler].

They must be doing it by removing all the carbonate and plant acid anions and leaving behind the potassium ions or by removing potassium from rectum of the hind gut, which has a pH of 5.0, and transferring it to the P-1 segment. Potassium is the dominant cation in the gut fluid of Zootermopsis. Such a high pH would tend to displace phosphate from the iron [Dixon p414] and aluminum and make it available to be absorbed. Brune and Kuhl (1996) suggest that the reason for the high pH is to enable termites to digest soil bacteria and/or to make polyphenolic compounds soluble and unable to bind peptide nitrogen compounds. This is plausible since wood eating termites have a high pH also and Lepidoptera and Diptera larvae that eat leaves and detritus have a high pH also [Brune and Kuhl, 1996]. Apparently termites can degrade lignin somewhat [Brune, Miambi, & Breznak, ]. This may add to the desirability of a high intestinal pH to termites, but I suggest the main imperative is to make phosphate soluble in eaters of soil humus. Sodium hydroxide is the best extracting medium of phosphate from soil [Cade-Menum ] and the hydroxide ion is a strong competitor for aluminum compared to others even phosphate and fluoride [Dixon p366]. Phosphate adsorbed on goethite (Fe OOH) has a steep decline after a pH of 8.0 to almost zero [Dixon p414].

Like the phosphate, the silicate of the soils would also tend to be displaced to form sodium silicate. The fore gut and rectum are acid. So the silicate must become the hydroxide and become very small colloids before it is excreted. Silicon hydroxide formed should be much more soluble than parent materials, and may even be able to move down through the soil in the form of small colloids, which are probably formed when the potassium silicate reaches the hind gut, especially if protected by organic anions. This may be the reason why tropical soils have the silica leached out of them. It could also explain the formation of silicretes and opals deep in Australian Tertiary savanna soils, silica for which arise from dissolution at the top [Thiry p733]. Weathering and leaching proceeds to greater depths when rainfall is concentrated in a short period [Dixon p361]. This, then, would explain the laterization of tropical soils. Schaefer has linked termites to latisols in South America because of the microcrystalline similarity to soil of termite mound material [Schaefer]. It would also explain the large deposits of bauxite (aluminum hydroxide) characteristic of some of them from the appropriate parent materials. Bauxite tends to be low in iron when subjected to constant rainfall [Dixon p361]. Bauxite probably remains intact in the middle soil profile because it is friable and therefore not suitable for cementing together sand particles. The termites bore down past the bauxite to the lower part of the profile to where the kaolin clay is in order to build their nests and runways in Australia, probably because the bauxite is friable and does not make strong construction. So it appears that some species move silicon up to the surface of the soil. These soils used to be called latisols, but are called oxisols and ultisols by the USDA and ferralsols or acrisols by the UN these days. There would be plenty of time for the leaching to take place because humivorus (eat humus) termites do not usually make above ground runways [Kooyman] as compared to the short time that erosion susceptible Amitermitinae (now called Termitinae) soils probably linger. It could also account for the source of the silicate to form the deposits of marine glauconite (green sand or iron silicate) laid down starting around early Cretaceous. and account for the rise in diatom diversity that commenced then [Miller 2005, diagram on p1294]. Diatoms are algae with a silica skeleton. The sediments in the North American interior sea way averaged 81% silica in the (Campanian) [Young]. Most of it was thought to be from diatoms, but some from sponges and radiolarians. It is primarily in the form of cristabolite (64%).

It is possible that some of the ancient soil eating roaches and wood roaches had the beginnings of such an attribute and thus account for the red beds which started in early Permian in South Africa, and in mid Permian in Europe, North America and Argentina [Veevers, et al]. In the Permian it was not necessarily humus eating wood roaches, but could have been humus eating cockroaches. The anaerobic reducing conditions in part of the termite gut may have assisted this process by reducing the iron in the minerals to the ferrous form [Vu], which oxidized to the ferric form upon reaching the soil, and could conceivably have contributed to red bed formation even with less of an alkaline gut. The soils of early Triassic were said to have no modern parallels, although some Madagascar soils may be fairly close. They were extremely low in organic matter and had no detritus. There was much less podzolization (or lacked podosols), although iron did migrate down to precipitate as iron carbonate [Retallack 1997]. Laterites or soils high in iron oxides are thought to have formed as early as the Triassic in north Australia [Twidale p170]. Since termites probably existed by that time [Emerson 1955 p476], ascribing laterites to termites is tentatively supported by paleontology. Laterites were widespread by the Cretaceous.

It has been proposed that the roots of plants remove silicon from the B horizon (soil below the top soil) of the soil and add it to the A horizon to form kaolin from the gibbsite clay [Lucas]. This is plausible, but I suspect that they were able to use more efficiently subsoil silicon because it had already been solubilized by the termites. It used to be proposed that tropical silica dissolved in the tropics because of high soil temperatures. However, silica solubility is independent of temperature between 0 and 200 degrees centigrade between a pH between 2 and 9.5 [Stever]. Do not feel that termites are not numerous enough to have had such an effect. Even in today's world kept in check by ants they transpire 2% of the world's carbon dioxide and 4% of the world's atmospheric methane [Sanderson], even though they are largely confined to tropical areas, and they were almost certainly much more numerous in the past. They are responsible for eating almost as much vegetation in their areas as vertebrates [Wood & Sands p280] and are the chief consumers in southwest USA range lands [Whiteford]. There can be over a kilogram of termites in a square meter in some areas [Eggleton].

REFERENCES are below

LINKS to other effects of termites and roaches on soil and vegetation.

Did the Wood Roach Cause the Permian - Triassic Coal Hiatus?: Digestion of cellulose by the wood roach may have reduced soil litter and enabled the rise of conifers in the Permian.
The Battles of Termites with Ants: The ability of Amitermitinae to smother plants with phosphorus rich runways may have caused a phosphorus famine in the Cretaceous.
Cretaceous Termites and Soil Phosphorus: Removal of soil phosphorus by erosion of termite runways may have resulted in changes in vertebrate bone evolution and explain animal sizes and shapes in our world.
Evolution of Angiosperm Trees: Angiosperm trees migrated across Western Pacific atolls and were made additionally successful by termites.

LINKS TO MARS GEOLOGY
The Canyons of Mars as Erosion by Rivers of Silicone Dust
---- For a hypothesis that explains the large volcanoes of Mars and the bulges associated with them. They are proposed as the disruption from the antipode (opposite side of a sphere) of a huge meteorite or comet impact.

GLOBAL WARMING
Climate warming as caused by denudation of soil.

GEOLOGY
----For a site that proposes a thin plate hypothesis to explain the plates in the crust of the earth, see this site. It has a link that explains the formation of ocean trenches.
----For a site that proposes disruption of the crust on Earth at the antipode (opposite side of a sphere) to produce massive lava flows.

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You may obtain a book about potassium nutrition at this site, along with the table of contents and first chapter. It discusses how food processing, diuretics, diarrhea, enemas, laxatives, corticosteroids, poisons, and disease states cause a deficiency and how potassium will cure heart disease, rheumatoid arthritis, gout, and hypertension. It also discusses procedures to cope with too high a blood potassium and abnormal potassium in diabetes.

Copper Response in Rheumatoid Arthritis: Nutrition and physiology of copper, especially relating to hemorrhoids, aneurysm, herniated discs, anemia, emphysema, and gray hair.
The Purpose of Cortisol: Cortisol is presented as an immune hormone used to defend against diarrhea
Cashew Nuts to Cure Tooth Abscess: Anacardic acids in raw cashew nuts may cure tooth abscesses and possibly gram positive diseases such as acne and leprosy.
Observations on Diabetes: Diabetes may be caused by a poison in food.
A review of what is known about chronic fatigue syndrome.Cronic Fatigue Syndrome
Fluoride in city water will cause fluorosis discoloration of teeth, weakened bones, damage to the kidneys and immune system, bone cancer and, worst of all, damage to the nerves resembling Alzheimer’s disease.

The Eve Controversy: A proposal as to why the human species seems to be derived from a single couple.

REFERENCES

------Abe, T., Bignell, D.E. and Higashi, M. (eds.) 2000. Termites: Evolution, Sociality, Symbioses, Ecology. Kluwer Academic Publishers, Dordrecht. 466 pp

------Bignell DE Anderson JH 1988 Determination of pH and oxygen status in the guts of lower and higher termites. Journal of Insect Physiology 26; 183-188

------ Bignell, D.E. Eggleton, P. 1995. On the elevated intestinal pH of higher termites (Isoptera: Termitidae). Insectes Sociaux 42, 57-69.

------Breznak, J.A.; Brune, A. 1994. Role of microorganisms in the digestion of lignocellulose by termites. Annu. Rev. Entomol. 39:453-487.

------Brune A, Miambi E, & Breznak JA 1995 Roles of oxygen and intestinal microflora in the metabolism of lignin derived phenyl propanoids and othermonoaromatic compounds by termites. Applied Environmental Microbiology 61; 2688-2695.

------Brune A & Kuhl M 1996 pH profiles of the extremely alkaline hind guts of soil - feeding termites (isoptera: Termitidae) determined by micro pH electrodes. Journal of Insect Physiology 42: 1121-7.

------Cade-Menum BJ Liu CW Nunlist T McColl JG 2002 Soil and litter phosphorus – 31 nuclear magnetic resonance spectroscopy. Extractants, metals, and phosphorus relaxation times. Journal of Environmental quality 31 457-465.

------- Dixon JB Weed SB 1989 Minerals in Soil Environments, 2nd edition. Number one in the Soil Science Society of America book series. Soil Science Society of America, Madison, Wisconsin.

------- Donovan, S.E; Eggleton, P.; Bignell, D.E. 2001. Gut content analysis and a new feeding group classification of termites. Ecol. Entomol. 26:356-366.

-------- Dow, J.A.T. 1992. pH gradients in lepidopteran midgut. J. Exp. Biol. 172:355-375.

------Eggleton P Bignell DE Sands WA Mawdsley NA Lawton JH Wood TG Bignell NC 1996 The diversity, abundance and biomass of termites under differing levels of disturbance in the Mbalmago Forest Preserve, southern Cameron. Philosophical Transactions of the Royal Society of London, series B 351; 51-68.

------Emerson AE 1955 Geographical origin and dispersions of termite genera. Fieldiana: Zool. 37; 465-521.

------- Ji, R.; Kappler, A.; Brune, A. 2000. Transformation and mineralization of synthetic 14C-labeled humic model compounds by soil-feeding termites. Soil Biol. Biochem. 32:1281-1291.

------Kappler A Brune A 1999 Influence of gut alkalinity and oxygen status on mobilization and size class distribution of humic acids in the hind gut of soil feeding termites. Applied Soil Ecology 13; 219-229.

------Koenig, H. (ed.) 2006. Intestinal Microorganisms of Termites and other Invertebrates. Soil Biology, vol. 6 (series ed. A. Varma). Springer-Verlag, Berlin, Heidelberg.

------Kooyman C Onck RFM 1987 Distribution of termite (isoptera) species in Southwestern Kenya in relation to land use and the morphology of their galleries. Biology and Fertility of Soils 3; 69-73.

------Lucas Y Luizao FJ Chauvel A Rouiller J Nahon D 1993 The relation between biological activity of the rain forest and mineral composition of soils. Science 260; 521-523.

------Miller KG Kominz MA Browning JV Wright JD Mountain GS Katz ME Sugarman PJ Cramer BS Christie-Blick N Pekar SF 2005 The Phanerozoic record of global sea level change. Science 310; 1293-1298.

------ Noirot, C. 1992. From wood- to humus-feeding: an important trend in termite evolution. In: Billen, J. (ed.), Biology and Evolution of Social Insects. Leuven University Press, Leuven, Belgium, pp. 107-119

. ------Retallack G 1997 Paleosols in the upper Narrabeen group of New South Wales as evidence of early Triassic paleoenvironments without exact modern analogs (review) Australian Journal of Earth Sciences 44; 185-281.

------Sanderson MG 1996 Biomass of termites and their emissions of methane and carbon dioxide, a global data base. Global Biogeography 10; 543-557.

----- Schaefer CER 2001 Brazilian latosols and their B horizon microstructure as long-term biotic constructs. Aust. J. Soil Res., 2001, 39, 909–926.

------Stever R 1962 Silica solubility, 0-200 degrees C and diagenesis of siliceous sediments. Journal Geol. 70; 127-150.

------ Thiry M Milnes AR Rayot V Simon-Coincon R 2006 Interpretation of palaeoweathering features and successive silicifications in the Tertiary regolith of inland Australia. Journal of Geological Society of London 163; 723-736.

------Twidale CR 1994 Gondwana (late Jurassic and Cretaceous) paleosurfaces of the Australian craton. Palaeogeography, Palaeoclimatology, 112; 157-186.

------Veevers JJ Conaghan PJ Shaw SE 1994 Turning point in Pangean environmental history at the Permian/Triassic (P/Tr) boundary. Geological Society of America Special Paper 187 to 196, Klein GD (ed) Geological Society of America, Boulder, Colorado.

------Vu AT Cao-Nguyen N Leadbetter JR 2004 Iron reduction in the metal-rich guts of wood-feeding termites. Geobiology 2 (4); 239–247.

------Whitford WG 1991 Subterranean termites and long term productivity of desert rangelands. Sociobiology 19; 235-243.

------Wood TG & Sands WA 1978 The role of termites in ecosystems. in; Production Ecology of Ants and Termites. Brian MV ed. Cambridge University Press.

------Young HR Moore PR 1994 Composition and depositional environment of the siliceous Odanah member (Campanian) of the Pierre Shale in Manitoba. In: Shur GW Ludigson GA Hammond RH, eds. Perspectives on the Eastern Margin of the Cretaceous Western Interior Basin. Special Paper 287 Geological Society of America, inc., Boulder Colorado.

Charles Weber - USA telephone = 828 692 5816 email = isoptera at att.net



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------- This article updated in Aug. 2010