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Plant Hormones and Worm casts or vermicomposts

Summary from Blakemore (2002) COSMOPOLITAN EARTHWORMS CD (Ref.)
Plant growth promoting substances (e.g. vitamins, plant hormones, enzymes and amino
acids) have been detected in earthworm extracts (e.g. Gavrilov, 1962; Nielson, 1965;
Graff & Makeschin, 1980; Dell'Agnola & Nardi, 1987 – as summarized in Blakemore,
1994). Earthworms have been found to stimulate soil enzymes, such as glucosidase and
phosphatase (possibly of microbial origin) which influence availability of plant nutrients
(e.g. Ross & Cairns, 1982; Tiwari et al., 1989). Nielson (1965) identified indole
compounds in extracts of several lumbricids while Springett & Syers (1979) suggested
that auxin-like substances are present in casts. Microbially derived plant hormones have
also been isolated from earthworm casts (Tomati et al., 1988). Production of specific
plant hormones by earthworms adds weight to the arguments for their co-evolution.
Plant growth is regulated by hormones which act at low concentrations to modify
the metabolic or developmental processes of the plant.
There are five main classes of plant hormone:
Auxins - there is only one naturally occurring auxin: indole-3-acetic acid (IAA) which is
chemically related to the amino acid tryptophan. Auxin alone promotes cell enlargement.
Synthetic auxins are aromatic compounds that affect plant growth in the same way as
auxin. The biggest use of these auxin-like compounds is as herbicides, eg. 2,4-D and
MCPA. Other examples of synthetic auxins are naphthalene acetic acid (used to control
fruit set and sucker growth on trees after pruning); and Indole butyric acid (used as
rooting compound).
Cytokinins – several naturally occurring forms, all related to the nucleotide adenine.
Only in combination with auxins do cytokinins promote cell division, growth and tissue
differentiation. Synthetic cytokinins include benzyladenine and kinetin which are used in
tissue culture media and for growth control in fruit.
Ethylene – is a hydrocarbon gas derived from the amino acid methionine. This gas is
used commercially for ripening fruit, especially bananas. Synthetics include ethephon
that can be applied as a spray for various applications. Ethylene is known to control
differentiation of root hair cells, and is also implicated in plant defense mechanisms,
possibly acting as a trigger that leads to disease suppressive soils (Penmetsa and Cook,
1997).
Abscisic acid – ABA is one of two compounds (the other is xanthoxin) related to
carotenoids. Canot be synthesized.
Gibberellins – GAs are the largest group with over 70 compounds, an example is
Gibberellic acid (GA3). They are used commercially to break dormancy and to promote
set of grapes. Gibberellins promote growth, so “anti-gibberellins” are used as retardants;
dwarf varieties of plants lack gibberellins.

Table 1. mauritii on aging under constant moisture & darkness at room temperature (26° C) _______________________________________________________________________ _ Weeks after casting Loss of activity compared to original level (%) Cytokinins Auxins _______________________________________________________________________ _ 0 0 0 1 2 1. cytokinins were benzyladenine equivalents. and that the concentrations of these diminished as the casts aged (Table 2). auxins were indole acedic acid equivalents. proteins and enzymes when compared to the soil matrix. Loss of plant growth promoter activities in paper pulp wormcasts of L. Rate of plant growth promoter in casts of two tropical earthworm species _______________________________________________________________________ _ Species Rate of production (nanograms day-1 worm-1) Cytokinins Auxins _______________________________________________________________________ _ Wormless control soil nil nil Lampito mauritii 4. excavatus. Common estimates of annual productions of surface casts in temperate regions of 4-250 t ha-1 are lower than some reports . 1995) found that plant hormone-like compounds were present in the casts of two earthworm species (Table 1).5 2 4 4 3 9 7 4 15 18 5 26 21 6 35 34 7 47 41 10 69 58 _______________________________________________________________________ _ Worm casts. mauritii and 6 for P.3 54 Perionyx excavatus 78. Considerable rates of deposition have been reported and several workers have also found casts to be enriched in plant available macronutrients and micronutrients. Table 2.Krishnamoorthy & Vajranabhaiah (1986. as reported in Ishmail.1 316 _______________________________________________________________________ _ Note: number of observations were 5 for L.

Shipitalo et al. which tend to neutrality. Nevertheless. 1985). Most studies have involved lumbricids. the Philippines. 1988). USA.600 t ha-1 yr-1. Australia. 1977. casts are used in India. 620 g.0 mm thick annually (Darwin. 40 and 60 kg K 2O ha-1 ) compared to soil only. Reddy surmised that the effects of the casts on the growth of rice may be due to the presence of “plant growth substances” identified by others as indole compounds also “yield-influencing substances” which could be secreted into the casts. Surface casting varies with species. Barois. 1881.000 t ha-1 yr-1.from the tropics. in turn increase the plant growth. 1987). Lunt & Jacobson (1944) established that worm casts are enriched in N. Reddy (1988) tested the effects of vermicasts or vermicompost (the product. ref2. 1992). along with worm biomass. season and soil type. Highest estimates in the tropics.5 t ha-1 casting in pastures in South Australia is exceptionally low and may be seasonal. chemical analysis of casts showed that they were richer in various plant nutrients . Lavelle. reported in Edwards & Lofty (1977: 144). In addition. China. ref3). K. 1985. and. Lee. by Lavelle (1988) and Lavelle et al.100 and 2. P. Significant increases in plant growth were recorded after four months. of vermicomposting operations) on growth of rice in pots: Rates were 310 g. Edwards & Lofty. and 940 g of casts per 2 kg soil (to give the equivalents to 20. Lavelle. Cuba and Mexico at least (ref1. (1988) give maximum sub-surface adjusted casting rates in the order of 280950 t ha-1 yr-1. Despite having higher nutrients. (Note: higher figures for cast production in the tropics of 2. (1989) are of over 1. Many subsequent studies have shown that secretion of CaCO3 or NH3 in the gut affects the pH of casts.2 to 2. Ca and Mg. for example. Lal. It is pertinent that most earthworms work below ground where their casting may be many times greater (Lavelle. Mansell et al. For example. but the common ranges above represent deposition of continuous layers of soil from about 0. relatively little is known about the cast compositions in other groups (cf. and that the amounts of exchangeable cations released in casts are often more than double the rates in the substrate (Lee. 1988). (1981) detected only slight increases in yields with cast material when added at "normal" rates of application (equivalent to average casting rates from the field). Barley’s (1956c) estimate of 2. are overcalculations from the original sources by a factor of ten!). 1881. of ranges of about 60-600 t ha-1 (Darwin. 1978. direct contribution of casts to improved plant growth may be moderate.

. friable Dichogaster saliens often inhabiting small chimneys Spenceriella (= Anisochaeta) minor casts not obvious Pontoscolex corethrurus medium sized globular or ribbon casts especially when soil wet. Pont.20).21): Table 4. burrows also along edges.21. Description of surface casts and soil matrices of dismantled cores. Spp Drawida barwelli Appearance of casts. A. Also in India. soils in the field. matrix ramified with re-filled burrows suggesting sub-surface casting Dichogaster affinis fine granular casts (0. there were significant increases in the colonisation of soil by nitrogen-fixing bacteria and mycorrhizal fungi. Higher levels of total nitrogen in the plot with added vermicompost was attributed to higher populations of nitrogen-fixing bacteria. taprobanae occasional surface casts Eudrilus eugeniae copious regular pellets 2-3x1 mm especially around sides. and Dichogaster spp. corethrurus. Jambhekar from Matarasha Agricultural Bioteka) that grapes fertilized with 2 tons of vermicompost per acre (ca. minor.5-1. large diameter burrows Species casting profusely on the surface were Eu. higher than yields under conventional fertilization and there were additional improvements in soils. surface pitted and many large ramifying burrows (2-9mm) Po. californica in either or both Narayen and Samford soils (Table 4. Native and exotic species casts were characterized by Blakemore (1994: tab. Polypheretima elongata. S.0 mm). (1992). Species that appeared to cast mainly below ground were Polypheretima elongata.3.3. eugeniae in all three soils. using mulches of vermicompost derived from grape marc or pomace led to 300% increase in grape yields within six months. found that when vermicompost was applied to a rice paddy as an organic fertiliser amendment. India (by H. 4. roots grow up into casts Apporectodea trapezoides extensive globular surface casts Metaphire californica patchy subrounded up-wellings Fletcherodrilus unicus distinctive elongate pellets 2-6x2 mm Eukkeria saltensis fine granules Digaster brunneus none. Kale et al.3.. 5 t/ha) per year for five years increased yields to 15 tons per acre. P. soils permeated with burrows Polypheretima elongata large globular (30 mm) and long threads. 1999). A report by Logsdon (1994) makes passing reference to a study in Pune. Australian trials. reported by Buckerfield & Webster (1998a. trapezoides and M.compared to the underlying soil.

Edwards and Lofty. Wang et al. several helical burrows were found in the cores on dismantling).3. 1988b). 1992). (1991) showed that when earthworms were excluded. absence of an earthworm fauna is often accompanied by undesirable accumulation of surface litter and root mats. 1988) but is reported for some species (Baylis et al. 1983. From the opposite position. Through feeding. Burrows and roots. Parker.. Conversely.22 and 4. argue that improved root growth does not always translate into increased crop yield. Several aspects and examples of the effects of burrows on root growth are summarised in Logsdon & Linden (1992) who. brunneus (despite mortality of this latter species. Several researchers have noted root proliferation in burrows which they usually attribute to the combined effects of following paths of least resistance and of better access to nutrient and moisture gradients in the drilosphere (e. an obvious build-up of the organic matter occurred in the litter layer.g. 1992). Although earthworms obtain much of their nutrition via dead root material and root exudates (Lee & Pankhurst. 1985. (1970) reported that the successful introduction of lumbricids to pasture soil in NSW greatly reduced root mats and stimulated cycling of N locked in this organic layer. 1978. This aptitude varies according to climatic conditions and the precise nature of the organic reserves (Martin et al. and extend as deep as 15 m (Blakemore.. Cortez & Bouché. and have been ecologically categorised according to. 1992). In temperate regions. Earthworm species vary in. 1983. Parmelee et al. 1991). Chemical analyses of samples of air-dried surface casts and topsoils for the Narayen clay and Samford sandy soils were presented in Blakemore (1994: tabs. 1985. Barley & Kleinig (1964) and later Noble et al. 1986. 1965. (1990) and Clements et al. Root mats and organic matter. 1989. 1986. Earthworms are able to exert considerable pressure when probing compacted soils (McKenzie & Dexter. van Rhee. 2000e). 1987). Burrow diameters may range from less than 1 mm to greater than 10 mm. 1959.3.taprobanae and Dg. Ehlers et al.. mixing and stimulation of microflora. 4. Therefore.23). 1988a. Dexter. they have profound implications not only for aeration and water conductivity but also for channelling of plant roots. 1980. 1977. earthworms in the humid tropics . their burrowing and feeding strategies (Lee. nevertheless. earthworms regulate the rates of mineralisation and immobilisation of soil organic matter (SOM). Springett. Additional observations were that casts in the denser clay soils were more aggregated and water-stable than those in sand. rhizophagy is uncommon (Lavelle.

although varying with species and region. Lavelle & Martin. 1992. Mesofauna and microorganisms. some of which increase logarithmically after passing through an earthworm's gut (Lee & Ladd. The trophic. Microfauna populations are also affected.g. 1984. with bacteria the least important (Lee & Ladd. algae less so. 1988). other microorganisms and actinomycetes. Lee & Pankhurst. Senapati (1992) found that mainly plant parasitic nematodes were reduced whereas microbivore nematodes increased. Miles (1963) found that Eisenia fetida cultured in sterile soil to which soil fungi and bacteria were added. quoted in Lee. regulate the rate of mineralization of nutrients from SOM reserves in different but balanced ways in both temperate and tropical environments. Some microorganisms are reduced.. 1992). involving gut symbioses.were found to have contrasting effects on SOM dynamics: accelerating mineralisation (of labile SOM) in the short-term due to digestion. Both external and internal relationships exist. symbiotic. 1985) reported nematode populations reduced by 37-66% when earthworms were introduced to soils. As for Collembola. have been proposed for several members of tropical and temperate species (e. Possibly the greatest influences are on protists. Mechanisms of mutualistic digestion in earthworms. but when soil protozoa were added. 1989. microarthropods. are killed during gut transit. especially (bacterivorous) protozoa and fungi. suggesting they are a major part of the earthworm diet. Edwards & Fletcher.. 1991. particularly microflora. failed to grow. microorganisms and fungi are complex (Lee & Ladd. Experimental evidence shows that certain organisms. 1988). Thus earthworms activities. Marinissen & Bok (1988) ascertained that presence of earthworms favoured survival of both larger species and individuals. whereas others proliferate. 1992). Edwards & Fletcher. Barois & . and synergistic relationships with beneficial or pathogenic nematodes. Martin et al. 1985. for example Yeates (1981. the worms grew to maturity. 1984. Martin. suggesting that at least in this species protozoa are an important part of the diet. competitive. but arresting the decomposition (of recalcitrant SOM) in the long-term by protecting nutrients within water-stable casts (Lavelle et al.

Martin.kr/~aeml/research_3_2004. Barois.ac. symbiotic N-fixing actinomycete) in one species (Reddell & Spain. It is further reported that soil borne diseases are less prevalent on organic . and higher soil counts of total microbes. but that parasitic nematodes were not affected. (2004) and Kwang-Hee Shin et al. (2004) isolated nearly 200 colonies of aerobic and anaerobic bacteria from the gut of E. gracilis.htm). Fungal pathogens Rosellinia necatrix.. 1988.Lavelle... have both been reduced by vermicompost (Stephens et al.gist. Ozawa & Risal. Martin et al. 1982). 1988.). (1992) reported increased colonization of plants by mycorrhizae. 1992. there are papers of culturing diazotrophic bacteria from the intestines of pheretimoids (e.. Lavelle.. 1991). 1987. 147-152 http://env1. vesicular-arbuscular mycorrhizas (VAM) were transmitted in casts of 13 species of earthworms. 1986. 1991a. Chem. and Horn et al. Mendez et al. the potential dual benefit of reduced severity of Rhizoctonia and "take-all" pathology in wheat combined with increased root colonization with symbiotic Rhizobium and Pseudomonas through earthworm activities were found (Dr P. comm.M Stephens pers. Moreover. and Plasmodiophora brassicae. reported in Lavelle et al. 2004). following application of vermicompost. clubroot of brassicas. 1987. as was Frankia (a nodulating. The intestinal caeca of such pheretimoid species probably serve to culture symbiotic gut flora and microbes (perhaps analogous to complex typhlosoles in other groups). 47: 137-142. and although I can find few references to the specific microbes involved. Barois et al. Beneficial. especially N-fixers and spore formers. Inoculation of sorghum seeds with Azospirillum brasiliense and earthworm casts increased growth of sorghum (Savalgi & Savalgi. (2005) described new N2O-producing bacteria from A.. Risal & Ozawa. 1999). (1993) found that Phytophthora and Fusarium fungi were suppressed by vermicompost. 1987). Experimental effects of vermicompost on some plant pathogens by Szczech et al. 2003).g. the potential for free N-fixing in the gut has been demonstrated in some earthworms (Barois et al. caliginosa gut. 1988) and beneficial organisms (Buckalew et al. white root rot of apples.. Recently. 1991b). Enhanced dispersal and viability of spores and propagules of fungi and microorganisms in the presence of earthworms have been reported for both pathogenic (Edwards & Fletcher. 1993. Recently Hyun-Jung Kim et al. 1987. Kale et al. Barois (1992) discusses the mutualistic microbiology of Amynthas corticis and A. 2002. Trigo & Lavelle. fetida (Agric. Biotechnol.

1980.R.N. PhD thesis Online. Gavrilov. Proc. Plant growth promoting substances (e. 1988). 1997. 1999). An assessment of plant growth promoter levels in the casts. Nielson (1965) identified indole compounds in extracts of several lumbricids.. Biological activity of earthworm casts. Chochin. and Cook.g. Sci. In Thampan. while Springett & Syers (1979) suggested that auxin-like substances are present in casts. 1989).). Sci. . R. 95: 341-351. Sultan. A report of an Indian study in Ishmail (1995) was of plant hormone-like compounds (Cytokinins were benzyladenine equivalents. (ed). Salmonella sp. References – other references may be sourced from these citations. A two-year project in Florida has further found that vermicomposting using earthworms to process sewage sludge (biosolids) was effective in reducing pathogen levels (viz.V. Pp 77-100. maruitii and P.. (1986).V.g. Cosmpolitan Earthworms CD. 1965. and helminth ova) to meet EPA Class A requirements (Eastman. Graff & Makeschin. Microbially derived plant hormones have also been isolated from earthworm casts (Tomati et al. Release of metabolites that stimulate plant growth has been proposed by several authors to explain some of the effects of earthworms on plant growth. (1997). excavatus that diminished in concentration as casts aged over 10 weeks. Jan 24.). vitamins. D.K. A. “Organic Agriculture”. Ismail.g. (Anim. Ross & Cairns. fecal coliform. Krishnamoorthy. enzymes and amino acids) have been detected in earthworm extracts (e. Penmetsa.farms that rely on the kinds of natural microbial activities invoked above (Workneh et al. Dell'Agnola & Nardi. Production of specific plant hormones by earthworms gives credance to arguments for plant/worm co-evolution. A legume ethylene-insensitive mutant hyperinfected by its rhizobial symbionts. Nielson. S. (Ref. R. 1987). Tiwari et al.. Peekay Tree Crops Development Foundation. enteric virus.. plant hormones. Plant growth factors. Blakemore (1994). 1994). Auxins were indole acedic acid equivalents) in casts of L. 1982. such as glucosidase and phosphatase (possibly of microbial origin) which influence availability of plant nutrients (e.. 1993 referred to in Hoitink & Grebus. Blakemore (2008). P. Earthworms have been found to stimulate soil enzymes. Indian Acad. and Vajranabhaiah. (1995) Earthworms in soil fertility management.. 1962. Science.

(1988). Pottinger (eds.A. The hormone-like effect of earthworm casts on plant growth.. (1979). A. E. Crosby & R. Springett. Biology and Fertility of Soils.P. J. Grappelli.)." Government Printer. "Proceedings of the 2nd Australasian Conference on Grassland Invertebrate Ecology. & Galli. 44-47. N. The effect of earthworm casts on ryegrass seedlings. . Pp.K.Z. & Seyers. 5: 288-294. In: T.Tomati. U. J.K. Wellington.