Uptake of silica byTrifolium incarnatum in relation to the concentration in the external solution and to transpiration

Summary The uptake of monosilicic acid by crimson clover (Trifolium incarnatum L.) was investigated using solution cultures in which the level ranged from 0.4 to 60 ppm SiO₂, and soils in which the level in solution ranged from 7 to 67 ppm SiO₂. With increasing levels of silica in the external solution there were systematic increases in uptake, but the quantities of silica in the tops were always less than those which were theoretically carried to the roots in the mass flow of water. The silica content of the roots was higher than in the corresponding tops and seemed to be largely associated with the epidermis. These findings and the observation that the concentration of monosilicic acid in the xylem sap is lower than that in the external solution, are regarded as evidence that the plant excludes a proportion of the monosilicic acid from the transpiration stream. This exclusion is attributed to a barrier in the root through which monosilicic acid passes at a slower rate than water.The distribution of silica among the parts of the tops was unaffected by the level of monosilicic acid in the external solution and, in turn, by the quantity which entered the tops. The chemistry of silica and the pattern of its distribution in the tops suggest that the monosilicic acid which has moved across the root then moves concomitantly with water in the transpiration stream and that silica is deposited in greatest quantities in those parts from which water is lost in greatest quantities.     

The effect of sesquioxides on the availability of molybdenum toTrifolium subterraneum

Summary Freshly precipitated ferric and aluminium oxides were added separately to soil and their effects on the availability of both native and added molybdenum were determined using subterranean clover (Trifolium subterraneum L.) as the experimental plant. Both oxides lowered the molybdenum content of the clover and, although it had a smaller specific surface area, ferric oxide had a greater effect than aluminium oxide. In a parallel study with similarly treated soil, the amounts of molybdenum extracted by 0.01M CaCl₂, 0.82M NaHCO₃ andM NH₄OAc were also lowered by both oxides, and ferric oxide again had the greater effect. Extraction with 0.275M NaHC₂O₄, as proposed elsewhere for assessing available molybdenum, did not show these differences.     

Studies of silica in the oat plant

Summary Oat plants,Avena sterilis L., were grown on soils in which the concentration of monosilicic acid in the soil solution, that is the level of supply of silica, ranged from 7 to 67 ppm SiO₂. Analyses at intervals throughout the growing period showed that the level of supply affected the amount and concentration of silica in the plant but not the pattern of its distribution among the parts.At maturity the caryopsis contained only 0.5 to 0.8 per cent of the total silica in the tops while the other parts of the inflorescence contained 40.7 to 41.3 per cent. The leaves (blade and sheath) contained 42.5 to 45.0 per cent of the total silica and the stems contained 7.8 to 10.9 per cent; the remainder was present in small sterile tillers.The concentration of silica in the dry matter was highest in the palea, lemma glumes, awn, and leaves. Among the leaves, the flag leaf had the highest silica content, both in terms of concentration in the dry matter and amount per leaf. The distribution of silica along a leaf followed a hyperbolic curve, the concentration being highest at the apex and lowest at the base of the blade.The chemistry of silica and the pattern of its distribution in the tops suggest that monosilicic acid and water move concomitantly in the transpiration stream and that solid silica is deposited in greatest quantities in those parts and regions from which water is lost in greatest quantities.     

The effect of lime on phosphorus adsorption and barley growth in three acid soils

For three acid soils from Santa Catarina, Brazil, lime application and time of incubation with lime had little effect on the adsorption of added phosphorus. In two soils with high contents of exchangeable aluminium, solution P and isotopically exchangeable P were decreased by incubating with lime for 1 month: phosphorus was probably adsorbing on freshly precipitated aluminium hydrous oxides. In one soil with less exchangeable aluminium, P in solution was increased by liming. After 23 months lime increased solution and exchangeable P possibly due to crystallization of aluminium hydrous oxides reducing the number of sites for P adsorption. All these changes were however small. In a pot experiment, lime and phosphorus markedly increased barley shoot and root dry matter and P uptake. Although liming reduced P availability measured by solution P, isotopically exchangeable P and resin extractable P, it increased phosphorus uptake by reducing aluminium toxicity and promoting better root growth. The soil aluminium saturation was reduced by liming, but the concentration of aluminium in roots changed only slightly. The roots accumulated aluminium without apparently being damaged.     

Sodium in the mineral nutrition ofAvena Sativa

Summary In pot experiments with oats on sandy soil poor in sodium and potassium a study was made of the effect of the addition of Na₂SO₄ and K₂SO₄ on the mineral composition of the straw.The addition of sodium at 200 ppm Na significantly increased the phosphorus content of the straw and there was a slight further increase with 200 ppm Na plus 200 ppm K, but the phosphorus content never exceeded 0.13 per cent.Sodium and potassium caused a highly significant decrease in the silica content of the straw. Decreased lodging in soils inundated with sea water can therefore not be ascribed to an increase of silica content.The calcium and the cellulose contents of the straw were both significantly reduced by sodium and potassium.     

The relative content of manganese in plants

Summary Oats, vetch, and rye were grown as successive crops in pots of a manganese-deficient soil to which -MnO₂ had been added in increasing amounts. Manganese deficiency of oats was overcome with 32 ppm of added manganese. Additions of 10, 18 and 32 ppm caused a decrease in concentration of maganese in the plants. Vetch and rye were unaffected by manganese deficiency and kept on accumulating manganese with increasing additions to the soil. Of the three plants vetch showed the greatest uptake of manganese, both with and without added manganese.     

The properties of a biologically formed manganese oxide,its availability to oats and its solution by root washings

Summary Manganese oxide, produced byCorynebacterium sp. in liquid medium was found to be amorphous, probably hydrated and was readily reduced by neutral quinol. Preparations of the oxide had values of n in the formula MnO_n which ranged from 1.76 to 1.88. The oxide was completely available to oats grown in sand culture but only slightly available in a manganese deficient soil. Plants grown under sterile conditions on agar slopes were able to obtain manganese from manganese oxide, indicating that the roots and not associated micro-organisms, were responsible for the solution process. Root washings of oat plants contained substances which dissolved manganese oxides and the activity of these substances increased with increasing acidity. The possible importance of these substances in making soil manganese available to plants is discussed.     

Si amelioration of Al toxicity in barley (Hordeum vulgare L.) growing in two Andosols

The influence of silicon on aluminium toxicity in barley (Hordeum vulgare L. cv. Shunrai) was studied in two Andosols. Silicon sources were a solution of sodium metasilicate with pH adjusted to 5.0, silica gel, and an industrial waste, porous hydrated calcium silicate. The waste is produced in large amounts in the manufacturing processes of autoclaved light concrete, and has been used as a silicon source for rice plants. The addition of the waste increased the concentration of Si in the soil solution, soil pH and amelioration of aluminium toxicity was observed. The addition of silica gel and sodium metasilicate solution to both soils increased significantly (p<0.05) the Si concentration of the soil solutions, but no amelioration of aluminium toxicity was observed. An amelioration of aluminium toxicity by the waste porous hydrated calcium silicate was probably due to the increase in soil pH rather than to the increase of silicon concentration in the soil solution.     

The influence of lead,cadmium, and nickel on the growth of ryegrass and oats

Summary A 2⁴ factorial experiment was conducted under greenhouse conditions. Factors and levels in the experiment were soil pH at 4.5 and 6.4, and cadmium, lead, and nickel added to the soil to provide soil concentrations of 50, 250, and 50 ppm, respectively, above background levels. Two species were grown in succession in the same experimental pots. Ryegrass (Lolium hybridum Hausskn. cv. Tetrelite) was harvested three times and then one crop of oats (Avena sativa L. cv. Garry) was grown and harvested. Plant tissue concentrations of cadmium, lead, and nickel were monitored throughout the experiment.Addition of cadmium to the soil lowered the dry matter yields in all three ryegrass harvests and also reduced the yield of oat grain. The application of lead nitrate to the soil enhanced the yield of ryegrass obtained at the first harvest and also increased the yield of oat grain. Nickel, added to the soil at 50 ppm, was relatively innocuous to ryegrass and oats.The presence of added cadmium, lead, or nickel to the soil resulted in enhanced tissue concentrations of these metals in both ryegrass and oats. This effect was particularly enhanced by a soil pH of 4.5. A single exception to this observation was that lead was not detected, under any of the conditions of this experiment, in oat grain.The presence of lead in the soil enhanced cadmium concentrations in ryegrass tissues. The presence of cadmium in the soil decreased lead tissue concentrations in ryegrass and oat straw. Soil pH interacted with both nickel and lead in the second ryegrass harvest with subsequent changes in tissue cadmium concentrations. At a soil pH of 4.5, the presence of added lead or nickel to the soil significantly increased the ryegrass tissue cadmium concentration beyond that observed at a soil pH of 6.4 with or without enhanced nickel or lead concentrations.     

Identification of the silicon form in xylem sap of rice (Oryza sativa L.)

Rice (Oryza sativa L.) is a typical silicon (Si)-accumulating plant, but the mechanism responsible for the translocation from the root to the shoot is poorly understood. In this study, the form of Si in xylem sap was identified by (29)Si-nuclear magnetic resonance (NMR) spectroscopy. In rice (cv. Oochikara) cultured in a monosilicic acid solution containing 0.5 mM Si, the Si concentration in the xylem reached 6 mM within 30 min. In the (29)Si-NMR spectra of the xylem sap, only one signal was observed at a chemical shift of -72.6 ppm, which is consistent with that of monosilicic acid. A (1)H-NMR study of xylem sap did not show any significant difference between the wild-type rice and mutant rice defective in Si uptake, and the components of the xylem sap were not affected by the Si supply. The Si concentration in the xylem sap in vitro decreased from an initial 18 mM to 2.6 mM with time. Addition of xylem sap to a solution containing 8 mM Si did not prevent the polymerization of silicic acid. All these results indicate that Si is translocated in the form of monosilicic acid through the xylem and that the concentration of monosilicic acid is high in the xylem only transiently.     

Solubility and dissolution of iron oxides

In most soils, FeIII oxides (group name) are the common source of Fe for plant nutrition. Since this Fe has to be supplied via solution, the solubility and the dissolution rate of the Fe oxides are essential for the Fe supply. Hydrolysis constants and solubility products (K_sp) describing the effect of pH on FeIII ion concentration in solution are available for the well-known Fe oxides occurring in soils such as goethite, hematite, ferrihydrite. K_sp values are usually extremely low ((Fe³⁺)·(OH)³=10^–37–10^–44). However, for each mineral type, K_sp may increase by several orders of magnitude with decreasing crystal size and it decreases with increasing Al substitution assuming ideal solid solution between the pure end-members. Based on such calculations a poorly crystalline goethite with a crystal size of 5 nm may well reach the solubility of ferrihydrite. The variations in K_sp are of relevance for soils because crystal size and Al substitution of soil Fe oxides vary considerably and can now be determined relatively easily.The concentration of Fe²⁺ in soil solutions is often much higher than that of Fe(III) ions. Therefore, redox potential strongly influences the activity of FeII. At a given pH and E_h, the activity of FeII is higher the higher K_sp of the FeIII oxide and thus also varies with the type of Fe oxide present.Besides the solubility, it is the dissolution rate which governs the supply of soluble Fe to the plant roots. Dissolution of Fe oxides takes place either by protonation, complexation or, most important, by reduction. Numerous dissolution rate studies with various FeIII oxides were conducted in strong mineral acids (protonation) and they have shown that besides the Fe oxide species, crystal size and/or crystal order and substitution are important determinative factors. For example, in soils, small amounts of a more highly soluble meta- or instable Fe oxide such as ferrihydrite with a large specific surface (several hundred m²g^–1) may be essential for the Fe supply to the plant root. Its higher dissolution rate can also be used to quantify its amount in soils. Ferrihydrite can be an important component in soils with high amounts of organic matter and/or active redox dynamics, whereas highly aerated and strongly weathered soils are usually very low in ferrihydrite. On the other hand, dissolution rates of goethites decrease as their Al substitution increases.Much less information exists on the rate of reductive and chelative dissolution of Fe oxides which generally simulate soil conditions better than dissolution by protonation. Here again, type of oxide, crystal size and substitution are important factors. Organic anions such as oxalate, which are adsorbed at the surface, may weaken the Fe³⁺-O bonds and thereby increase reductive dissolution. As often observed in weathering, the dissolution features of the crystals appear to follow zones of weakness in the crystal.     

The effect of manganese-oxidizing bacteria and pH on the availability of manganous ions and manganese oxides to oats in nutrient solutions

Summary The uptake of Mn from manganous ions (Mn-ions) and pyrolusite (MnO₂) by three week-old oat plants (Avena sativa L.) grown in nutrient solutions controlled at pH values between 6 and 8, was almost completely inhibited by suspensions of Mn- oxidizing bacteria over a three day uptake period.Grey speck symptoms of Mn deficiency developed in oats grown for 10 days with Mn bacteria in a nutrient solution that had received 1 ppm Mn ions and was controlled at pH 6.3. Rape plants (Brassica napus L.) absorbed appreciable amounts of Mn from treatments similar to those that inhibited Mn uptake by oats.Treatments which decreased or prevented biological oxidation of Mn ions favoured the uptake of Mn by oats from Mn ions, MnO₂ and bacterial Mn-oxide. Acid conditions (pH 5.0) always increased Mn uptake. This was due in part to inhibition of bacterial oxidation and to an increase in the ability of the plants to obtain Mn from Mn oxides.Uptake of Mn is explained on the basis of the rates of two opposing processes; the rate of release of Mn from oxides and the rate of biological oxidation of Mn ions. The results are discussed in relation to the availability of Mn in soils.     

Review of methodologies for extracting plant-available and amorphous Si from soils and aquatic sediments

There is a variety of methodologies used in the aquatic sciences and soil sciences for extracting different forms of Si from sediments and soils. However, a comparison of the published extraction techniques is lacking. Here we review the methodologies used to extract different Si fractions from soils and sediments. Methods were classified in those to assess plant-available Si and those to extract Si from amorphous silica and allophane. Plant-available Si is supposed to comprise silicic acid in soil solution and adsorbed to soil particles. Extraction techniques for plant-available Si include extractions with water, CaCl₂, acetate, acetic acid, phosphate, H₂SO₃, H₂SO₄, and citrate. The extractants show different capabilites to desorb silicic acid, with H₂SO₃, H₂SO₄ and citrate having the greater extraction potential. The most common extractants to dissolve amorphous silica from soils and aquatic sediments are NaOH and Na₂CO₃, but both also dissolve crystalline silicates to varying degrees. In soils moreover Tiron is used to dissolve amorphous silica, while oxalate is used to dissolve allophanes and imogolite-type materials. Most techniques analyzing for biogenic silica in aquatic environments use a correction method to identify mineral derived Si. By contrast, in the soil sciences no correction methods are used although pedologists are well aware of the overestimation of amorphous silica by the NaOH extraction, which is most commonly used to extract silica from soils. It is recommended that soil scientists begin to use the techniques developed in the aquatic sciences, since it seems impossible to extract amorphous Si from soils completely without dissolving some of the crystalline silicates.     

Characterization of Gaseous Ozone Decomposition in Soil

Laboratory scale batch experiments were performed to investigate the decomposition characteristics of gaseous ozone in porous media. The decomposition rates of gaseous ozone in several solid media were determined, and the relationship of moisture content with sorbed ozone molecules was evaluated. Ozone decomposition in control and glass beads packed columns followed second-order reaction kinetics, while ozone consumption in a sand-packed column demonstrated first-order kinetics with a rate constant of 0.0109 min^?1 and half-life of 1.0 h. The presence of typical metal oxides in the soil resulted in ozone consumption rates in the following order: hematite (Fe₂O₃) > silica-alumina (SiO₂Al₂O₃) > alumina (Al₂O₃) > silica (SiO₂). Ozone decomposition was highly dependent upon soil moisture content. Over 90% of the total ozone mass decomposed in the field soil with moisture content at less than 1 wt%, whereas as low as 5–15% of the total ozone mass degraded with moisture content at more than 2 wt%. In conclusion, ozone decomposition in soils was primarily controlled not only by soil organic matter but also by reactive metal oxides on the soil surface. These two factors were shown to be highly dependent upon soil moisture content.     

Kinetics and reactions of hydrogen sulphide in solution of flooded rice soils

Summary The sulphide-ion electrode was used to study the kinetics and reactions of free hydrogen sulphide in solution of flooded rice soils. The observed sulphide potential obeyed the Nernst equation over a range of sulphide-ion concentration from 10^-1 to 10^-19 M. Peak H₂S concentrations were lowest in neutral soils high in iron and manganese; moderately high in soils low in iron or high in organic matter; and highest in acid sulphate soil low in iron. Harmful concentrations of H₂S may be present in acid sulphate and acid soils low in iron during the first few weeks after flooding. The concentrations in acid sulphate soils can be drastically lowered by liming. There was thermodynamic evidence for the presence of FeS and ZnS in the solutions of most soils.     

The evaluation of plant-available sulphur in soils. II

Summary The availability to oats of adsorbed sulphate in soils and of sulphate impurity in calcium carbonate was studied in pot-culture experiments.When calcium carbonate was added to soils with pH values ranging from 5.7 to 7.4 the uptake of sulphur by oats was increased, due probably to enhanced mineralization of soil organic sulphur. When the calcium carbonate contained sulphate impurity the uptake of sulphur was further increased by an amount comparable with the release of sulphate which could be expected from a reaction of the calcium carbonate with the exchangeable hydrogen of the soil. Sulphate in excess of this amount appeared to be largely unavailable. Uptake of sulphur by oats from calcareous sands containing large amounts of insoluble sulphate associated with calcium carbonate also suggested that soil sulphur in this form had very low availability to plants.Substantial increases in the amounts of sulphur extracted by reagents commonly used for the determination of adsorbed sulphate in soils occurred when soils were airdried at about 20°C. Decreases in adsorbed sulphate in soils following the growth of oats in pot culture confirmed that adsorbed sulphate is readily available to plants.     

The influence of heavy metals upon the growth of sitka-spruce in South Wales forests

Summary Experiments on sitka-spruce seedlings grown in acidic peaty gley soils under green-house conditions, where the soils where doped with increasing amounts of Cd, Cu and Pb up to maximum levels of metal added of 16 ppm, 32 ppm and 400 ppm respectively, showed that the levels of Cd and Pb in shoots and roots increased with increasing levels in the soil, whereas levels of copper appeared to be independent. The addition of these three metals to the soils did not influence the uptake of other heavy metals, or of the nutrients potassium or calcium. Increases in the shoot cadmium levels significantly reduced the yields of the plant shoots. However, the plant yields were only affected by the highest level of lead that was added to the soil (400 ppm Pb) and unaffected by all the copper treatments (0–32 ppm Cu in the soil). The lengths of the sitka-spruce roots were reduced when cadmium and lead levels in the soil exceeded certain threshold concentrations (2.5 ppm total Cd, where 0.3 ppm was extractable with 0.5 M acetic acid; and 48 ppm total Pb, where 1.7 ppm was extractable). However, root lengths were not reduced by copper. This was probably related to the fact that copper appears to be relatively unavailable in the type of soil used, as only 1.1. ppm Cu was extractable from a total of 32 ppm Cu added. Root branching was apparently reduced by increases in the soil levels of cadmium, copper and lead. The roots of some control plants had symbiotic mycorrhizal associations (4 out of 19 plants), whereas the roots of all the plants grown in the soils with added heavy metals did not develop these.     

Stimulation by nickel of soil microbial urease activity and urease and hydrogenase activities in soybeans grown in a low-nickel soil

Summary The concentration of nickel in some soils may be insufficient to meet the requirements of enzymes such as urease in soybeans and hydrogenase in Rhizobium. In an initial evaluation of nickel availability, several soils were examined for nickel content and microbial urease activity. Total and extractable nickel were determined by atomic emission spectrometry. Purified glucose and urea were added to soils to stimulate microbial growth and urease activity, the latter of which was monitored by the rate of decomposition of¹⁴C urea. Nickel also was added to some samples to determine if the indigenous supply was limiting. In one low-nickel soil (total Ni 13 ppm) urease activity increased 150% in response to additional nickel, while other soils (total Ni 22–3491 ppm) failed to respond to nickel. However, additional nickel did stimulate urease activity (up to 109%) in 3 out of 10 soils to which purified CaCO₃ was added. Presumably the rise in pH associated with this treatment decreased nickel availability. Additions of Co, Mn, Fe, or Cu had no consistent effect on urease activity, thus indicating that the response to Ni was specific. Nickel fertilization increased leaf urease and nodule hydrogenase activity of soybeans grown in low-nickel soil, however, yield was not improved. These results may have practical implications in the nutrition of plants and micro-organisms that metabolize H₂ and urea.     

Effects of Plant Residue and Salinity on Fractions of Cadmium and Lead in Three Soils

Bioavailability and mobility of heavy metals (HMs) in soils are determined by their partitioning between solution and solid-phase and their further redistribution among solid-phase components. A study was undertaken to determine the effects of organic matter (OM) and salinity on cadmium (Cd) and lead (Pb) distribution among soil fractions. Three agricultural soils were treated with 20 mg Cd/kg as Cd (NO₃)₂·4H₂O, 150 mg Pb/kg as Pb (NO₃)₂, 20 g/kg alfalfa powder, and 50 mmol/kg of NaCl, and then incubated at 60% water holding capacity (60% WHC) and constant temperature (25°C) for 12 weeks. Various fractions of Cd and Pb were extracted from the soils after 2 and 12 w of incubation using a sequential extraction technique. Results showed that in the early stage of incubation (2 w), added Pb were found mainly in the specifically sorbed (SS) and amorphous Fe oxides (AFeO) fractions and added Cd found in SS and Mn oxides (MnO) fractions. Addition of 2% OM decreased the exchangeable (EXC) Pb fraction almost in all soils, whereas it had a different effect on the EXC Cd fraction depending on soil pH. Addition of NaCl increased the EXC Cd fraction in two soils, but it did not alter Pb fractions. At the end of the incubation period, Pb decreased in the EXC and MnO fractions except in the neutral soil and Cd decreased mainly in the SS fraction.     

Water near its Supercritical Point and at Alkaline pH for the Production of Ferric Oxides and Silicates in Anoxic Conditions. A New Hypothesis for the Synthesis of Minerals Observed in Banded Iron Formations and for the Related Geobiotropic Chemistry inside Fluid Inclusions

An alternative hypothesis for the origin of the banded iron formations and the synthesis of prebiotic molecules is presented here. I show the importance of considering water near its supercritical point and at alkaline pH. It is based on the chemical equation for the anoxic oxidation of ferrous iron into ferric iron at high-subcritical conditions of water and high pH, that I extract from E-pH diagrams drawn for corrosion purposes (Geophysical Research Abstracts Vol 15, EGU2013–22 Bassez 2013, Orig Life Evol Biosph 45(1):5-13, Bassez 2015, Procedia Earth Planet Sci 17, 492-495, Bassez 2017a, Orig Life Evol Biosph 47:453-480, Bassez 2017b). The sudden change in solubility of silica, SiO₂, at the critical point of water is also considered. It is shown that under these temperatures and pressures, ferric oxides and ferric silicates can form in anoxic terrains. No Fe^II oxidation by UV light, neither by oxygen is needed to explain the minerals of the Banded Iron Formations. The intervention of any kind of microorganisms, either sulfate-reducing, or Fe^II-oxidizing or O₂-producing, is not required. The chemical equation for the anoxic oxidation of ferrous iron is applied to the hydrolyses of fayalite, Fe₂SiO₄ and ferrosilite, FeSiO₃. It is shown that the BIF minerals of the Hamersley Group, Western Australia, and the Transvaal Supergroup, South Africa, are those of fayalite and ferrosilite hydrolyses and carbonations. The dissolution of crustal fayalite and ferrosilite during water-rock interaction needs to occur at T&P just below the critical point of water and in a rising water which is undersaturated in SiO₂. Minerals of BIFs which can then be ejected at the surface from venting arcs are ferric oxide hydroxides, hematite, Fe^III-greenalite, siderite. The greenalite dehydrated product minnesotaite forms when rising water becomes supersaturated in SiO₂, as also riebeckite and stilpnomelane. Long lengths of siderite without ferric oxides neither ferric silicates can occur since the exothermic siderite formation is not so much dependent in T&P. It is also shown that the H₂ which is released during hydrolysis/oxidation of fayalite/ferrosilite can lead to components of life, such as macromolecules of amino acids which are synthesized from mixtures of (CO, N₂, H₂O) in Sabatier-Senderens/Fischer-Tropsch & Haber-Bosch reactions or microwave or gamma-ray excitation reactions. I propose that such geobiotropic synthesis may occur inside fluid inclusions of BIFs, in the silica chert, hematite, Fe^III-greenalite or siderite. Therefore, the combination of high-subcritical conditions of water, high solubility of SiO₂ at these T&P values, formation of CO also at these T&P, high pH and anoxic water, leads to the formation of ferric minerals and prebiotic molecules in the process of geobiotropy.