Microbial Mats: A Bioreactor of Lithification

Microbial Mats: A Bioreactor of Lithification Microbial mats: a bioreactor of lithification and an indicator of Earths evolution Introduction Microbial mat is a general term that is used to describe a variety of microbial communities that are found at interfaces between different types of material, mostly on submerged or moist surfaces such as estuarine environment and salt marshes (Krumbein et al., 1977; Nicholson et al., 1987). Bacteria and archaea are two main microbes forming the layers. Microbial mats contain a variety of different but essential trophic groups including primary producers, consumers, and decomposers. This is why even though microbial mats are, to an extreme extent, geographically small, they are ecosystems from an ecological perspective. Microbial mats are dynamic ecosystems in which a wide range of metabolic processes take place. Inside this tiny ecosystem, different physical and chemical environments are distinguished by a variety of gradients, include but not limited to light, oxygen and sulfide (Visscher and van den Ende, 1994). The gradients may not be always constant. For example, oxygen concentration may have varied from diurnally to seasonally. In some aquatic systems, it will drop from supersaturated to undetectable within a few centimeters. The light penetration depth is fluctuated because of change of seasons or just with cloud covering. All these temporal environmental oscillations mentioned above, will result in coupled reactions, that are critical to the biogeochemical cycle, like reduction and oxidation of elements such as carbon and sulfur. Therefore, heterogeneity of microbe habitat is a common character that exhibits among all microbial mats.   Microbial mat ecosystems can be viewed as a semiclosed system which require little more than sunlight to function, as such it is efficient in all kinds of reactions and element cycling. The relatively simple but functional structures make it, to a certain extent, easy to reach equilibrium and mass balances. Generally, microbial mats tend to have high rates of oxygenic photosynthesis, aerobic respiration, sulfate reduction, and sulfide oxidation (Canfield and Des Marais, 1993; Revsbech et al., 1986), when compared to other benthic ecosystems. A classical view of a microbial mat (Figure 1)(Visscher et al., 2000) is that a fixed sequence of microbial groups exists: starting with oxygenic cyanobacteria as a surface community, underlain by oxygenic phototropic bacteria and sulfate-reducing bacteria as subsequent layer (Krumbein, 1983). This view, however, was later questioned and revised. Structure and the layers are not a result of different metabolic reaction types, on the contrary, they might be found in association with the cyanobacterial layer. Some research showed that the sulfur reducing bacteria was also found in the surface layer (Frà ¼nd and Cohen, 1992; Visscher et al., 1992). Microbial mats and mineral interaction In microbial ecosystems, when the precipitation rate of minerals is faster than that of dissolution, lithification will occur. Precipitations mediated by microbial mats is not limited to carbonates but also constituted by other minerals, such as gypsum and anhydrite (Ehrlich, 1998). Among all these precipitation types, carbonate precipitation is perhaps the most important process as it is directly related to the global carbon cycling. Therefore, in this section, a main focus will be put on sedimentary biofilms in hypersaline environments to help with the interpretation of the rock record. 2.1 Stromatolites and carbonate precipitation Stromatolites are lithifying organic sedimentary structures formed by microorganisms (Figure 2). Carbonate precipitation activities of microbial mats are trapped and recorded in stromatolites layered structures. As such, microbial mats can be viewed as bioreactors (Dupraz et al., 2004a). The stromatolites structure is characterized as an alternating soft and hard layers whose heights ranges from a few centimeters to two meters. The evolutionary processes of stromatolites remain largely uncharacterized (Zavarzin, 2002). There are two major hypotheses. Des Marais (1997) speculated that microbial lithification is a result of by-product of microbial metabolism. On the other hand, McConnaughey and Whelen (1997) suggested that this could be directly related to the consequence of microbes harvesting energy from protons released during calcium carbonate precipitation. However, regardless of origin, stromatolites have thrived for a long history that could be seen as a major evolutionary advan ce for us to study the Earths early history and global biogeochemical cycles. Cyanobacteria have played a crucial role in carbonate precipitation as shown in Figure 3. Two microbially as well as physicochemically controlled factors determine carbonate precipitation: the saturation index (SI) and exopolymeric substances (Lozano-Garcà ­a et al.). SI = log(IAP/Ksp), where IAP denotes the ion activity product (i.e. [Ca2+]*[CO2-]) and Ksp, the solubility product of the corresponding mineral (10-6.37 for calcite at 25 °C, 1bar atmospheric pressure and 35 PSU salinity (Zeebe and Wolf-Gladrow, 2001)). If IAP > Ksp, the solution is supersaturated, and when SI > 0.8, calcite carbonate tends to precipitates (Kempe and Kazmierczak, 1994). Or else, calcite carbonate will dissolve. The [CO32-] depends on the carbonate equilibrium, which comprises three species as followed: H2CO3, HCO3 and CO32-. In another word, pH is influencing the precipitation. Therefore, before investigating how microbial metabolism affect the CaCO3 precipitation, understanding production and consum ption of inorganic carbon and the environmental pH change is a prerequisite. EPS act as a chelator for cations and the template for crystal nucleation (Costerton et al., 1995; Decho, 2000). It is constantly modified by including but not limited to UV radiation, pH and microbial degradation (e.g. through hydrolysis, decarboxylation). 2.2 Microbial mats and lithification Contemporary microbial mats, vertically laminated ecosystems, resemble the layered sedimentary structures of stromatolites. As such, they have been attracting extensively research interests for being analogues for stromatolites. Shown in Figure 3, there are 6 different functional groups of microbes exist in microbial mats. From top of the figure to the bottom are: Cyanobacteria act as primary producers, which are believed to affect the trapping and biding of sediments; Aerobic heterotrophic bacteria, which gain energy from oxygen respiration and organic carbon; Anoxygenic phototrophs, mainly purple and green bacteria, which using HS- for photosynthesis; Sulfate reducing bacteria (SRB), which respiring organic carbon with SO2- while producing HS; Sulfide oxidizing bacteria (SOB), chemolithoautotrophs that oxidize HS with oxygen or nitrate while fixing CO2; fermenters, using organic carbon or sulfur compounds as electron donor and acceptor. However, this view of the mat composition is facing challenge because nucleic acid sequences will undoubtedly reveal more diverse and complex community structures than the simple classified ones. Cyanobacteria is more like an important mediator of biogeochemical cycle of the mats ecosystem. It produces oxygen for the whole system to be functional (Fenchel, 1998). As mentioned before, the mat ecosystem is very efficient and productive. The relatively high photosynthetic rates, which shows a diurnal fluctuation, will reach its peak in the afternoon. Aerobic heterotrophs respire during the daytime when there is abundant oxygen, thereby creating an anoxia environment at twilight. Fermenters degrade complex organic molecules into smaller ones and benefit the SRB. SOB and anoxyphototrophs have contributed less to carbon fixation comparing with cyanobacteria and the role of fermentation remains ambiguous. All these activities above have resulted in steep vertical geochemical gradients with extreme diel fluctuations (Figure 3). To understand the role of microbial mats in precipitation and dissolution, it is important to determine both the abundance and metabolic activity of these key functional groups. Because the quality and quantity of EPS are largely determined by the metabolic activity of the community. In the previous researches, several microbial mat systems have been found to produce carbonate phases: travertine in hot springs in Yellowstone (Fouke et al., 2000), dolomite in Lagoa Vermelha, Brazil (Vasconcelos and McKenzie, 1997) and Salt Pan, Bahamas (Dupraz et al., 2004a). However, there are still mats that will no lithify or fossilize. So here comes the question, what determines the lithification potential? A previous study, using a combination of geological and microbial techniques, of lithifying microbial mat systems in hypersaline lake system was carried on in Salt Pan in Eleuthera, Bahamas (Dupraz et al., 2004a). The lake is not deep with an average depth less than 60cm. From the shoreline towards the center of the lake, a gradient from lithifying mats to jellylike soft mats exists (Figure 4). The shallow water column was found to contain cyanobacterial pigments that efficiently quench the sunlight. Not surprisingly, the photosynthesis, aerobic respiration, sulfate reduction are generally higher and geochemical gradients are steeper in the shallower lithifying mats. Moreover, EPS is easily destructed by strong UV radiation in shallower mats. This process helps with removing inhibition of precipitation by releasing more Ca2+ into the environment. The combination of these processes benefits carbonate precipitation. 2.3 Microstructure of precipitation and EPS UV radiation will cause browning reactions, dehydration and alkalinity. However, EPS production in stromatolite mat can prevent damages such as desiccation of the mat, retains essential nutrients, and provides water channels for transporting metabolites and signaling compounds (Costerton et al., 1995; Decho, 2000). Decho, A.W. et al. (Decho et al., 2005) had shown that EPS production in a stromatolite mat accounted only for 8% of 14HCO3uptake during the light, and a rapid turnover followed during the dark. They concluded that despite the fast rate of production, the net EPS production was low. The production and consumption are in equilibrium. Once being hydrolyzed, EPS components were readily consumed by the mat community, particularly anaerobes instead of aerobes. This is somehow surprising that when Schizothrix EPS, xanthan, or sugar and amino acid monomers and polymers that comprise EPS were supplied in mats, stimulation of anaerobic heterotrophic activity stimulation was greater than aerobic heterotrophs activity (Decho et al., 2005; Visscher et al., 2000). The combined action of fermentative organisms and SRB could be responsible for this high consumption rate. Oxygen levels are influenced by the rapid and extensive diurnal fluctuations as well as cloud cover and O2-consuming cell clusters in the EPS can produce anoxic microenvironments, therefore, the anaerobic pathway plays an important role in microbial EPS degradation. EPS can not only release Ca2+and HCO3 during microbial alteration, but also influence chemical gradients, which will in turn affect the mineral phases. The EPS matrix preferably slows down the mobility of hydrated Mg2+, therefore, temporarily increase relative abundance of Ca2+(Figure 5). The delay of Mg diffusion would lead to a decrease of the Mg2+:Ca2+ ratio of mineral products forming inside the EPS (Verrecchia et al., 1995). As mentioned above, changes in the amount or type of EPS could influence the rate of precipitation or types of crystals formed. 2.4 Microbial metabolism and saturation index Simple redox reactions form the basis of microbial metabolism. These metabolic reactions often involve C and either O, S or N (Figure 3;(Fenchel, 1998)). Daytime and nighttime metabolism of the six key functional groups is typically different, especially when it is influenced by oxygen and sunlight. Chemical alterations of the microenvironment that result from different metabolic reactions might change the alkalinity and thus facilitate carbonate precipitation or dissolution (Visscher and Stolz, 2005). Microbial mats have a high metabolic activities, thus it is not surprising that the rapid SI changes, despite the internal buffering capacity of the carbonate system, would result in a chemical alteration of the microenvironment. High rates of cyanobacterial photosynthesis cause a rapid depletion of CO2, which challenge the resilience or reestablishment of the carbonate equilibrium, and the increasing alkalinity will results in carbonateprecipitation through removal of the Hthat is pro duced in the latter reaction. It should be noted that in these reactions, organic carbon is assumed to be CH2O and different outcomes are expected with different organic compounds. For example, CO2produced bythe decomposition of carboxylic acids, will potentially increase the carbonate alkalinity by CO2degassing(Visscher et al., 1992). As such, this could probably explain why heterotrophic aerobes have been shown to precipitate carbonate. Microbial mats as an indicator of sulfur evolution The sulfur cycle has evolved over the long history of the Earth, with the concentration and the isotopic fractional abundance much different between Precambrian and contemporaneous environment (Cameron, 1982). The surface environment of the early Earth was basically reducing. Little atmospheric oxygen existed. Even though it is still under debate how the oxygen was produced at first, a majority of researchers believe that the history of atmospheric oxygen and seawater sulphate are closely linked (Habicht and Canfield, 1996; Ohmoto et al., 1993; Walker and Brimblecombe, 1985). Sulphate in Archaean and early Proterozoic sediment was found to be consistent in 34S depletion, which is similar to meteorites and mantle-derived igneous rocks (Cameron, 1982; Monster et al., 1979). Moreover, sulphate level was found to positively influence the rate of 34S depletion as lower levels sulphate (

Sexual Ethics Essay

1. There are a number of dilemmas in sexual ethics such as homosexuality, marriage and divorce and pornography. For instance homosexuality is a major problem within sexual ethics because homosexual sex cannot lead to reproduction. However sexual ethic it differs between denominations compared to government law. Therefore these issues are a concern to religious belief; different faiths have different view on homosexuality or marriage and divorce.Many religious beliefs do not encourage homosexuality in fact it is severely discouraged for example in the Islamic faith they do not allow this because there is no doubt that in Islam homosexuality is considered ‘sinful'. Homosexuality as far as Islam is concerned is a profound mistake (as are all sins if they are not intending to do wrong). For instance a quote from the Qur’an says â€Å"What! Of all creatures do ye come unto the males, and leave the wives your Lord created for you? Nay, but ye are forward folk. † Qur'an 26:165.The references relate to gay sexual activities; lesbian practices are not mentioned in the Qur'an, neither is it mentioned in the bible. Christians are divided over the issue of homosexuality. Due to natural law homosexual sex cannot lead to reproduction therefore the marriage would be considered invalid, however then a marriage between an older couple or couples who infertile would also be invalid, therefore all homosexual acts and relationships are sinful so many believe homosexuality is contrary to God’s will.However the Roman Catholic Church teaches that homosexual should be treated with respect, compassion and sensitivity and no discrimination, they believe the feeling aren’t wrong but the actions are sinful because they contrary to the will of God. The scriptures of the bible suggest that the only sexual relationship that God approves of is one between married heterosexual partners. The creation narrative describe woman as being created especially for man: â€Å"a helpful suitable for him† (genesis 2:18). The Old Testament said â€Å"If a man lies with a man as one does with a woman, both of them have done what is detestable.They must be put to death† (Leviticus 20:13) however there is still no mention on lesbian practices. Marriage is an important part of the Christian life a the purpose of Christian marriage is faithfulness to one another, many Christians teach than within marriage there is a natural hierarchy that reflects the relationship of Christ and the church. The Roman Catholic Church says â€Å"wives, in the same way be submissive to your husband†¦ Husband, in the same way be considerate as you live with your wives, and treat them with respect† (1 peter 3:1, 7).Whereas the Church of England argues that the relationship is based upon Christ’s headship over the church and is about love and sacrifice, not domination and power. This argument originally started as the bible said â€Å"a man will leave his father and mother and be united to his wife and they will become one flesh† (genesis 2:24). On the other hand, in the Islamic faith marriage is only a way to make sex and having children legal and lawful, the belief also states in the eyes of Allah both members of the relationship are treated equally even though they have separate duties. Whereas the Christian views on divorce are that marriage is for life.People take vows for better or worse so if they meant that then there should be no divorce. For instance the Roman Catholic Church believe there shouldn’t be any divorce unless the marriage has not been consummated or they are willing to get a divorce with the permission from the pope. Whereas the Church of England except divorce but some more traditional vicars don’t allowed divorces to remarry in church, the bible believes divorce is not favoured by God: â€Å"Therefore what God has joined together, let man not separate (Matthew 19:6) which means God joins them so the law cannot separate them.Even Jesus forbids divorce: â€Å"Anyone who divorces his wife and marries another woman commits adultery. And if she divorces her husband and marries another man, she commits adultery† (Mark 10:11-12) However in Matthews gospel it say â€Å"anyone who divorces his wife, except for marital unfaithfulness, and marries another woman commits adultery† (Matthew 19:9). Alternatively divorce in Islam is only when the male partner says Talaq 3 times can divorce the couple, however this banned in some countries which results in going to arbitration council and judicial intervention before the divorce is granted. . Due to the loss of faith in many denomination and cultures today, we as people no longer look to religion to guide our sexual ethics as we once did. It is no longer relevant because many people believe that anymore should be able to marry anyone they want, for example same-sex partners. The debate is still being waged be tween the church and government to offer this union. Many people believe if two people are happy together and want to join themselves in holy matrimony, then they should be allowed.Marriage has also undergone many changes as people are less likely to have a â€Å"traditional† wedding instead they are more likely to enjoy extended co-habitation. Also the concept of marriage has change because it is more of a legal act rather than a declaration of love to God. Whereas divorce is seen by the eyes of most people is all about the law rather than church. Church seems to have very little authority when a marriage has got to the divorce stage.A minority of people do still take guidance from the church and believe the once they have entered into holy matrimony then it is for life just as God ordained. In conclusion, like most of the civilised world I believe that the religious beliefs about sexual ethics are no longer relevant because civilisation has changed so much and people are wi lling to accept all that was once considered taboo by the church and are willing to open their mind about the change in religious beliefs and sexual ethics.

Introducing Term Paper

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Civil Disobedience And The Civil Rights Movement - 881 Words

In Thoreau s essay Civil Disobedience he makes the point that bystanders are just as bad as criminals and that people should stand against unjust crimes even if it means going against the law. And to some extent I do agree because in the past people have broken unjust laws and have created change. A well-known example would be when Rosa Parks sat on the bus in the White-only seating area, which lead to important events that helped push the Civil Rights movement forward. But I think that it depends on which laws they choose to break and how far they choose to go with it. Honestly though, I would not want to break the law even if it were unjust unless it was greatly affecting my family or the people I care about because I would rather not risk going to jail or losing everything that I have. There s also the fear of losing respect and not being able to find a job. In Henry David Thoreau s essay Civil Disobedience, he says, I saw to what extent the people among whom I lived could be trus ted as good neighbors and friends; that their friendship was for summer weather only... (Civil Disobedience, 12). After going to jail he had lost the people around him, they no longer treated him the same. He did not commit a violent crime, he just refused to pay his taxes and was labeled a criminal. If his friends could treat him differently because of that, then imagine how employers would look at him. But, if there were ever a time when I needed to break a law to protect my familyShow MoreRelatedCivil Disobedience And The Civil Rights Movement867 Words   |  4 PagesDuring the Civil Rights Movement, King and many of his followers and fellow activists deeply followed the path of non-violent protest, otherwise known as civil disobedience. 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