Biogeochemical phenomena induced by bacteria within sulfidic mine tailings

Summary Mill tailings resulting from mining and metallurgical processes are usually disposed of into open-air impoundments, where they become subjected to chemical or microbial leaching. At the surface of the tailings, where oxic conditions prevail, acidophilic bacteria, such as thiobacilli, can oxidize sulfidic minerals (e.g. pyrite and pyrrhotite) and generate acidic metal-rich leachates as by-products of their metabolism. This, combined with chemical oxidation, leads to acid mine drainage (AMD). Biomineralization, whereby a proportion of the metal leachate is precipitated, can also occur in the oxidized tailings, often as a result of a close metal-bacteria interaction. Iron-rich precipitates are usually found on bacterial cell walls, and are thought to serve as nucleation sites for further mineralization within the tailings impoundments. As depth increases in mine tailings, oxygen depletion and the presence of water-saturated pores usually lead to anoxic conditions. Under such redox and chemical conditions, populations of sulfate-reducing bacteria (SRBs) can colonize the tailings. As a result of their metabolic activity, sulfate is reduced to hydrogen sulfide, which in turn can react with dissolved metals to form metal sulfide precipitates. Microbial sulfate reduction also generates alkalinity, although chemical dissolution of carbonate and oxide minerals probably also play an important role in the generation of alkaline conditions in mine tailings.     

Significance of microbial communities and interactions in safeguarding reactive mine tailings by ecological engineering

Pyritic mine tailings (mineral waste generated by metal mining) pose significant risk to the environment as point sources of acidic, metal-rich effluents (acid mine drainage [AMD]). While the accelerated oxidative dissolution of pyrite and other sulfide minerals in tailings by acidophilic chemolithotrophic prokaryotes has been widely reported, other acidophiles (heterotrophic bacteria that catalyze the dissimilatory reduction of iron and sulfur) can reverse the reactions involved in AMD genesis, and these have been implicated in the "natural attenuation" of mine waters. We have investigated whether by manipulating microbial communities in tailings (inoculating with iron- and sulfur-reducing acidophilic bacteria and phototrophic acidophilic microalgae) it is possible to mitigate the impact of the acid-generating and metal-mobilizing chemolithotrophic prokaryotes that are indigenous to tailing deposits. Sixty tailings mesocosms were set up, using five different microbial inoculation variants, and analyzed at regular intervals for changes in physicochemical and microbiological parameters for up to 1 year. Differences between treatment protocols were most apparent between tailings that had been inoculated with acidophilic algae in addition to aerobic and anaerobic heterotrophic bacteria and those that had been inoculated with only pyrite-oxidizing chemolithotrophs; these differences included higher pH values, lower redox potentials, and smaller concentrations of soluble copper and zinc. The results suggest that empirical ecological engineering of tailing lagoons to promote the growth and activities of iron- and sulfate-reducing bacteria could minimize their risk of AMD production and that the heterotrophic populations could be sustained by facilitating the growth of microalgae to provide continuous inputs of organic carbon.     

Dust emission of chromium from chromite ore processing residue disposal areas in the vicinity of Krasnogorskii Village in Chelyabinsk oblast

The chromium content of soil, dust, and plants near an open-pit mine used to store self-decomposing chromite ore processing residue (COPR) of the Chelyabinsk Electrometallurgical Combine was studied by synchrotron radiation X-ray fluorescence analysis (SRXFR).     

Transmission Electron Microscopic Observation of Mercury-Bearing Bacterial Clay Minerals in a Small-Scale Gold Mine in Tanzania

Mercury is a toxic substance that is widely distributed throughout the hydrosphere, biosphere, and lithosphere. Mine waste environments and mine waters support a wide diversity of microbial life. The microbial ecology of environments where mine waters are polluted with heavy metals is poorly understood. Here, we describe the features of bacteria in mercury-contaminated gold panning ponds in a small-scale gold mine (Geita) near Lake Victoria, Tanzania using energy filtering transmission electron microscopy (EF-TEM) and scanning transmission electron microscopy equipped with energy dispersive X-ray spectroscopy (STEM-EDX). Most bacteria in the panning pond showed thick exopolysaccharides (EPSs), and many clay minerals attached onto the surface of EPSs. The clay minerals and EPSs might act as protective layers for the bacteria against toxic materials. The clay minerals were composed of smectite, halloysite, and kaolinite associated with calcite and goethite. Scanning electron microscopy equipped with energy dispersive X-ray spectroscopy indicated that the bulk soil samples contained abundant Si, Al, K, Ca, and Fe with heavy metals such as Au, Ti, and Ag. The results indicate that Hg pollution from panning ponds is caused by not only volatilization of Hg from Au-Hg amalgams, but Hg is also released into the air as dust mixed with dry fine clays, suggesting high long-term environmental risks. Mercury-resistant bacteria associated with clay minerals may have a significant effect on the weathering processes of the ore during long-term bioremediation. The clay mineral complexes on the surface of bacterial cell walls are a stimulator for Hg-resistant bacterial growth in mud ponds contaminated with the Au-Hg materials.     

Acid-Generating and Leaching Potential of Soils in a Coal Waste Rock Pile in Northeastern China

The present study involved the assessment of potential generation of acid drainage and also metal leaching from an abandoned large (175 m) low sulfur waste rock pile—the dominant mine waste at the site—at the Haizhou coal mine. Laboratory-based static and column leaching tests on waste rock samples were conducted. The static tests were done for 8 composite samples collected from different parts of pile. A column study was performed using mixture of waste rock soil samples to assess metals attenuation process of background soils in vicinity of pile when rainwater reacts with low-sulfide waste rocks. Total concentration of major elements in solid samples was determined using dispersive X-ray fluorescence spectrometry (XRF) and chemical analysis of leachate were measured by inductively coupled plasma optical emission spectrometry (ICP-OES), ion chromatography and titration methods. The inverse geochemical modeling using PHREEQC was applied to explain possible mass transfer processes between column leachates of waste pile and background soils. Static tests (including acid–base accounting (ABA) and net acid generation (NAG)) and mineralogical information suggest that the waste rock is non-acid generating. The large amount of aluminosilicate minerals and probably trace amounts of carbonates with respect to low sulfur content of waste rock pile represents a potentially large source of neutralization potential according to static test results. It was also found that presence of inherent neutralizing materials in waste rock and also in surrounding background soils provide sufficient neutrality and possibly immobilize the trace and heavy metal contents of waste rocks and potentially protects water resources.     

INVESTIGATION OF A DIATOM POPULATION FROM MINE TAILINGS IN NYE COUNTY,NEVADA, U. S. A.1

A diatom population consisting of 31 taxonomic entries was found in tailings from an abandoned mine near Ichthyosaur Paleontologic State Monument, Nye County, Nevada, U.S.A. The diatom population of the mine tailings resembled those found in dry lakes of the vicinity which are similar to some soil diatom populations. The mine tailings population was very similar to some Quaternary diatom assemblages in the vicinity and in other parts of the conterminous United States. The diatom population indicated that alkaline and oligohalobous conditions were prevalent with associated “indifference” with respect to the current spectrum.     

Effects of different soil ameliorants on karee trees (Searsia lancea) growing on mine tailings dump soil-part I: pot trials

Rehabilitation of mine tailings dams is often a challenge due to a lack of nutrients and a poor humus reservoir prevailing in tailings soils. This is especially true for establishing longer lived species such as trees. For these reasons the effects of different soil ameliorants (woodchips compost, vermicompost, mature sewage sludge), added to the root system of Karee (Searsia lancea) saplings were tested in pot trials. Those pots were filled with platinum and gold tailings substrate as well as red clay soil, respectively. For three months plants remained in a greenhouse and were subsequently moved to a test field outside. Throughout the test period regular chl a fluorescence measurements were taken and subjected to JIP-test quantifying changes in photosynthetic vitality status. Additionally, growth measurements and one-off leaf analysis were carried out. Test plants growing on mine tailings experienced an up to 35% higher average photosynthetic vitality (PI(ABS)) and improved nutrient supply, when treated with mature sewage sludge. Consequently, sewage sludge treated plants showed a higher biomass build-up rate and an up to 55% higher diameter growth, compared to control. In summary the experiments present a low cost alternative for reforestation enterprises on platinum and gold tailings dams in South Africa.     

Can Physiological Endpoints Improve the Sensitivity of Assays with Plants in the Risk Assessment of Contaminated Soils?

Site-specific risk assessment of contaminated areas indicates prior areas for intervention, and provides helpful information for risk managers. This study was conducted in the Ervedosa mine area (Bragança, Portugal), where both underground and open pit exploration of tin and arsenic minerals were performed for about one century (1857 – 1969). We aimed at obtaining ecotoxicological information with terrestrial and aquatic plant species to integrate in the risk assessment of this mine area. Further we also intended to evaluate if the assessment of other parameters, in standard assays with terrestrial plants, can improve the identification of phytotoxic soils. For this purpose, soil samples were collected on 16 sampling sites distributed along four transects, defined within the mine area, and in one reference site. General soil physical and chemical parameters, total and extractable metal contents were analyzed. Assays were performed for soil elutriates and for the whole soil matrix following standard guidelines for growth inhibition assay with Lemna minor and emergence and seedling growth assay with Zea mays. At the end of the Z. mays assay, relative water content, membrane permeability, leaf area, content of photosynthetic pigments (chlorophylls and carotenoids), malondialdehyde levels, proline content, and chlorophyll fluorescence (F_v/F_m and Φ_PSII) parameters were evaluated. In general, the soils near the exploration area revealed high levels of Al, Mn, Fe and Cu. Almost all the soils from transepts C, D and F presented total concentrations of arsenic well above soils screening benchmark values available. Elutriates of several soils from sampling sites near the exploration and ore treatment areas were toxic to L. minor, suggesting that the retention function of these soils was seriously compromised. In Z. mays assay, plant performance parameters (other than those recommended by standard protocols), allowed the identification of more phytotoxic soils. The results suggest that these parameters could improve the sensitivity of the standard assays.     

Anthyllis vulneraria/Mesorhizobium metallidurans, an efficient symbiotic nitrogen fixing association able to grow in mine tailings highly contaminated by Zn, Pb and Cd

The excessive concentrations of toxic heavy metals in mine tailings and their very low N content make soil reclamation strategies by phytostabilization difficult. Our objective was to test if the symbiotic association between the legume Anthyllis vulneraria subsp. carpatica and the bacteria Mesorhizobium metallidurans originating from highly polluted mine tailings is able to increase N concentration in soils with contrasting Zn, Pb and Cd contents. Plants of A. vulneraria subsp. carpatica from a mine site and of a non-metallicolous subsp. praeopera from non-polluted soil were inoculated with a metallicolous or a non-metallicolous compatible Mesorhizobium spp. and grown on low and high heavy metal-contaminated soils. In contaminated soil, many nodules were observed when the metallicolous A. vulneraria was inoculated with its rhizobium species M. metallidurans, whereas the non-metallicolous A. vulneraria died after a few weeks regardless of the rhizobium inoculant. Eighty percent of the total nitrogen was derived from biological nitrogen fixation through the association between metallicolous A. vulneraria and the rhizobium grown on metal-enriched soil. The ability of the metallicolous A. vulneraria to develop a high nitrogen fixing potential opens new possibilities for promoting a low-maintenance plant cover and for stabilizing the vegetation in high heavy metal-contaminated soils.     

Characterization of Blue Agave Bagasse (BAB) as Raw Material for Bioethanol Production Processes by Gravimetric,Thermal, Chromatographic,X-ray Diffraction,Microscopy, and Laser Light Scattering Techniques

A detailed characterization of the main types of blue agave bagasse (BAB) obtained from the four largest tequila factories in the State of Jalisco (Mexico) is presented here. After milling/sieving the agave bagasses, two particle size fractions were identified, one rich in fibers and the other consisting of dust/fine particles. Both fractions were analyzed to determine the content of cellulose, hemicellulose, lignin, organic-soluble compounds, absorbed remaining sugars, minerals, and organic matter. After detailed analyses of both fractions by wet, thermal (thermo-gravimetric analysis (TGA)/differential thermo-gravimetric analysis (DTA)), and other methods (high-performance liquid chromatography (HPLC), microscopy, particle size by laser diffraction light scattering, and crystallinity by X-ray diffraction), a moderate-to-intensive method was devised for further processing the fibrous fraction, which had a high crystalline cellulose content, as well as for its subsequent enzymatic saccharification under well-defined moderate conditions. Alternative processing options were also devised for the dust/fine particle fraction, which has a moderate crystalline cellulose that is rich in adsorbed sugars and that has a high mineral matter content.     

Trace Element Concentrations in Soil,Sediments, and Waters in the Vicinity of Geita Gold Mines and North Mara Gold Mines in Northwest Tanzania

The “Geita Gold Mine” (GGM) and “North Mara Gold Mines” (NMGM) have developed large quarries in the middle of fertile agricultural lands. Possible hazardous impact on the natural recourse has warranted a study on the trace element concentrations in soils, sediments, and natural waters. Generally, the study shows a great variation in type of elements and intensity of their spill to soils, sediments, and waters. We found indications of acid mine drainage (AMD) of trace metals near the GGM waste rock tailings, but the data are inconclusive. The environmental impact at NMGM was, on the other hand, more extensive. Severe trace element contamination of sediments and waters near one of the mining facilities managed by NMGM is connected to an accidental acid spill in 2009. However, we found strong indication that, in addition to the accident, leakage of alkaline wastewater into surface soils and free water took place from a large wastewater reservoir. We found very high concentrations, especially of As in sediments and water samples downside this reservoir. Water from several sites contained As concentration in the free water at more than one order of magnitude higher than the WHO drinking water recommendations. The chemical speciation of water samples indicated high fraction of Free Ion Activity (FIA) in several samples and the change in pH had a significant impact on the FIA. Due to extensive alkaline and acidic runoff from these sites, more information about hydrological transport routes and the chemical speciation of the free water is of great importance for assessing potential risk of these mines.     

华南铅锌尾矿生态恢复的理论与实践

介绍了近年来有关华南地区5个铅锌尾矿生态恢复研究领域的主要研究成果。内容包括：铅锌尾矿的理化性质和影响植物定居的限制因子;尾矿酸化的预测与控制;尾矿的基质改良;尾矿自然定居植物的生态对策;重金属耐性植物的筛选与耐性机理;豆科植物在废弃地植被恢复中的作用;尾矿湿地系统的重建及其废水处理效率;土壤种子库在尾矿生态恢复中的作用;尾矿植被恢复的野外中试研究等。同时,也讨论了尾矿废弃地生态恢复的未来研究方向。     

Arsenic contamination of soils and sediments from tailings in the vicinity of Myungbong Au mine,Korea

Abstract

Geochemical characteristics of As contamination in the Myungbong gold mine area in Korea were investigated, and the mine tailings were found to contain high concentrations of As (2,500–6,420 mg/kg). The mine tailings are an obvious candidate for the source of contamination in the study area. From the sequential, and oxalate, extraction analyses the majority of the As in the tailings was found to be associated with Fe oxides produced when sulfide minerals are oxidised. Mineralogical examinations showed the formation of poorly crystalline Fe oxides on a rim of pyrite, onto which the As was retained. The sediments of two ponds affected by the tailings also contained significant levels of As. High concentrations of dissolved As were also found in pond and stream waters. It has been suggested that the mobility of the As was increased by the reductive dissolution of Fe oxides under reducing conditions in the pond. Agricultural fields, located beneath the tailing dumps, have been contaminated by the continuous use of contaminated stream water and groundwater for irrigation.     

Microbial distribution and diversity in saturated, high pH, uranium mine tailings, Saskatchewan, Canada

Microbiological analyses were conducted on core samples collected along a vertical profile (0-66 m below surface) from the tailings management facility (TMF) at the Rabbit Lake uranium mine in northern Saskatchewan, Canada. Bacterial numbers in the core materials were similar to surrounding soils and surface waters, regardless of the seemingly unfavorable pH (mean=9.9) and temperature (approximately 0 degrees C) in the TMF. The greatest number of viable cells (105 CFU/g) was detected at the interface between the tailings and overlying standing water, below which cell counts decreased rapidly with depth. Whole-community metabolic profiles for samples from the different depths grouped into 3 clusters; however, these groups could not be positively correlated with sampling depth, temperature, redox potential, pH, or ore-mill feed. Flow-cell studies demonstrated microbial communities in the tailings surface water could develop biofilms and maintain cell activity at both pH 10 and 7, and altering the pH between these 2 values had little effect on biofilm viability. These results demonstrate the resilience and adaptive nature of naturally occurring microbial communities and signify a potential role of microbial activity in the long-term geochemical evolution of the TMF.     

Dust as a Nutrient Source for Fynbos Ecosystems,South Africa

Abstract Fynbos is the main vegetation of the Cape Floristic Region, a biodiversity hotspot that occurs in southwestern South Africa. A major question concerning the fynbos ecosystem is how it supports abundant and diverse vegetation on soils derived from nutrient-poor bedrock. In addition to marine aerosols (recycled sea salts), geochemical analyses reported here suggest that dust (aeolian) deposition represents a significant source of nutrients (for example, K, Ca and Zn) to the fynbos ecosystem. Headwater portions of the Boontjies River sub-catchment near the Cederberg Mountains support mountain fynbos communities that are entirely underlain by the Peninsula Formation, a quartz arenite with greater than 98 wt% SiO₂. Fynbos soils in these areas are composed of quartzose sand with 3–6 wt% kaolinitic clay and 1–2 wt% organic carbon. The minor amount of feldspar and mica minerals in the bedrock (0.5 wt% Al₂O₃) suggests an aeolian source for much of the clay minerals in the soil. The isotope composition of soluble Pb and Sr from fynbos vegetation and soils indicates a mixture of anthropogenic and terrigenous sources, most likely from washout of combusted petrol and dust from the arid interior particularly in association with Berg Wind events. Approximate mass balance calculations indicate that washout of aerosols provides an important source of nutrients such as Ca, K, P, Fe, Mn and Zn which the fynbos ecosystem is highly effective in retaining.     

Microbial communities in a porphyry copper tailings impoundment and their impact on the geochemical dynamics of the mine waste

The distribution and diversity of acidophilic bacteria of a tailings impoundment at the La Andina copper mine, Chile, was examined. The tailings have low sulfide (1.7% pyrite equivalent) and carbonate (1.4% calcite equivalent) contents and are stratified into three distinct zones: a surface (0-70-80 cm) 'oxidation zone' characterized by low-pH (2.5-4), a 'neutralization zone' (70-80 to 300-400 cm) and an unaltered 'primary zone' below 400 cm. A combined cultivation-dependent and biomolecular approach (terminal restriction enzyme fragment length polymorphism and 16S rRNA clone library analysis) was used to characterize the indigenous prokaryotic communities in the mine tailings. Total cell counts showed that the microbial biomass was greatest in the top 125 cm of the tailings. The largest numbers of bacteria (10(9) g(-1) dry weight of tailings) were found at the oxidation front (the junction between the oxidation and neutralization zones), where sulfide minerals and oxygen were both present. The dominant iron-/sulfur-oxidizing bacteria identified at the oxidation front included bacteria of the genus Leptospirillum (detected by molecular methods), and Gram-positive iron-oxidizing acidophiles related to Sulfobacillus (identified both by molecular and cultivation methods). Acidithiobacillus ferrooxidans was also detected, albeit in relatively small numbers. Heterotrophic acidophiles related to Acidobacterium capsulatum were found by molecular methods, while another Acidobacterium-like bacterium and an Acidiphilium sp. were isolated from oxidation zone samples. A conceptual model was developed, based on microbiological and geochemical data derived from the tailings, to account for the biogeochemical evolution of the Piuquenes tailings impoundment.     

Culturable and molecular phylogenetic diversity of microorganisms in an open-dumped,extremely acidic Pb/Zn mine tailings

A combination of cultivation-based and molecular-based approaches was used to reveal the culturable and molecular diversity of the microbes inhabiting an open-dumped Pb/Zn mine tailings that was undergoing intensive acid generation (pH 1.9). Culturable bacteria found in the extremely acidic mine tailings were Acidithiobacillus ferrooxidans, Leptospirillum ferriphilum, Sulfobacillus thermotolerans and Acidiphilium cryptum, where the number of acidophilic heterotrophs was ten times higher than that of the iron- and sulfur-oxidizing bacteria. Cloning and phylogenetic analysis revealed that, in contrast to the adjacent AMD, the mine tailings possessed a low microbial diversity with archaeal sequence types dominating the 16S rRNA gene library. Of the 141 clones examined, 132 were represented by two sequence types phylogenetically affiliated with the iron-oxidizing archaea Ferroplasma acidiphilum and three belonged to two tentative groups within the Thermoplasma lineage so far represented by only a few environmental sequences. Six clones in the library were represented by the only bacterial sequence type and were closely related to the well-described iron-oxidizer L. ferriphilum. The significant differences in the prokaryotic community structures of the extremely acidic mine tailings and the AMD associated with it highlights the importance of studying the microbial communities that are more directly involved in the iron and sulfur cycles of mine tailings.     

Stabilization of Oxidic Tailings and Contaminated Soils by Calcium Oxyphosphate Addition: The Case of Montevecchio (Sardinia,Italy)

The effectiveness of an in situ heavy metals fixation technique aimed at converting contaminants to low solubility and low bioavail-ability forms, eliminating the risk posed by oxidic tailings and contaminated soils, was investigated. Calcium oxyphosphate salt (Ca(H₂PO₄)₂·H₂O) was used as a stabilizing agent for oxidic tailings and contaminated soils originating from Montevecchio, Sardinia, Italy. Stabilization was effected by mixing the contaminated soil or oxidic tailing sample with calcium oxyphosphate salt at various doses. The effectiveness of stabilization was evaluated by USEPA TCLP standard toxicity testing. Complementary EDTA extraction tests and biological tests using beans Phaseolous vulgaris as plant indicator were carried out. The toxicity of Pb and Cd was reduced below TCLP regulatory limits at calcium oxyphosphate doses higher than 0.7 and 0.2% w/w for soils and tailings, respectively. Lead solubility according to the EDTA test decreased with phosphate dose for both materials tested. Lead uptake by plant leaves and roots from the soil sample decreased with the phosphate addition, while Cd uptake remained almost constant. An adverse effect on plant growth and Zn uptake was observed for calcium oxyphosphate dose up to 1.1% w/w. Based on the results, a remediation scheme for oxidic tailings and contaminated soils is proposed.     

Skeleton bones in museum indoor environments offer niches for fungi and are affected by weathering and deposition of secondary minerals

Large skeleton specimens are often featured as iconic open displays in Natural History Museums, for example, the blue whale ‘Hope’ at the Natural History Museum, London. A study on Hope's bone surface was performed to assess the biodeterioration potential of fungi. Fungi were isolated, and a fungal internal transcribed spacer (ITS) clone library survey was performed on dust and bone material. Mineral particles derived from bone and dust were analysed using energy dispersive X-ray spectroscopy, variable pressure scanning electron microscopy (SEM) and high vacuum SEM. Results showed that bone material, although mainly mineral in nature, and therefore less susceptible than organic materials to biodeterioration phenomena in the indoor environments, offers niches for specialized fungi and is affected by unusual and yet not so well-documented mechanisms of alteration. Areas of bone surface were covered with a dense biofilm mostly composed of fungal hyphae, which produced tunnelling and extensive deposition of calcium and iron-containing secondary minerals. Airborne halophilic and xerophilic fungi including taxa grouping into Ascomycota and Basidiomycota, capable of displacing salts and overcome little water availability, were found to dominate the microbiome of the bone surface.     

Mine tailing disposal sites: contamination problems,remedial options and phytocaps for sustainable remediation

Mining practices and the absence of proper mine land reclamation has led to heavy metal contaminated sites with serious impact on the ecosystems and risk for human health. The origin of the contamination is often associated to mine tailing deposits because they are a source of the acid mine drainage (AMD). These areas are devoid of vegetation due to the harsh soil conditions that prevent the rooting of plant species. The remediation of these areas followed by revegetation is necessary to suppress the generation of the AMD and its negative effects on the ecosystems. Conventional remediation technologies for heavy metal contaminated sites are usually not applicable because of the high cost associated with chemicals and energy requirements, as well as the long treatment time to remediate large areas. In this study, the use of phytocapping for the remediation of mine tailing deposits and abandoned mine areas is reviewed. Phytocapping is cost effective, environmentally friendly and has multifunctional role against various problems of mine tailings: it provides erosion control, landscape rehabilitation, enhances the soil properties for further colonization of other more demanding vegetal species, reduces the leachability of metals downwards the groundwater, and favors the immobilization of metals forming less bioavailable species. The most critical step in phytocapping is the developing of the first vegetative cover because of the biotoxicity of the mine soil and mine tailings. Several amendment materials can be used to ameliorate soil conditions creating a favorable environment for the rooting of plants, as well as serving as a source of nutrients. Local plant species with fast growing are preferable because their adaptation to the soil and climate conditions favors their self-propagation.