metabolites as all microbes obtain them in finished, prefabricated form from their external environment. But, however, they do differ with respect to procurement of their organic metabolites (called “food”) i.e. the source of carbon (sources of carbon normally also contribute both oxygen and hydrogen). On the basis of their difference in procurement of organic metabolites i.e. the source of carbon the microorganisms are broadly categorized as autotrophs and heterotrophs. Autotrophs (Gr. auto = self, trophe = nourishing) are those that manufacture (synthesize) their organic metabolites (food) from CO; supplied from external environment as raw material and an organic complex already existing in their cell with the help of their enzymic equipment. That is, the autotrophs use CO, as their sole or principal source of carbon. This mode of food-procurement (nutrition) is called autotrophy or autotrophic nutrition. Heterotrophs (Gr. hetero = other, trophe = nourishing), however, cannot manufacture (synthesize) organic metabolites (food) on their own like autotrophs rather they must obtain (absorb) organic metabolites (food) in prefabricated form from their external environment. That is, the carbon already prepared in organic form by other living organisms. This mode of food-procurement (nutrition) is called heterotrophy or heterotrophic nutrition, To produce the food either by synthesis (autotrophy) or by obtainment (absorption) in pre-fabricated form (heterotrophy), all microorganisms need energy This energy comes either from sun (photo; light-energy) or from break-down of chemicals (chemo; chemical-energy). In this way, taking source of energy into account, the microorganisms can be categorized as phototrophs (that use light as their energy source) and chemotrophs (that obtain energy from the oxidation of chemical compounds, either organic or inorganic). In addition to the sources of carbon and energy, the microorganisms also require sources of electrons for growth to take place. Microorganisms can be categorized as lithotrophs and organotrophs on the basis of their electron sources. Lithotrophs (i.e. “rock-eaters”) are those that use reduced inorganic substances as their electron source, whereas organotrophs extract electrons from organic compounds. The above mentioned relationships of the various nutritional forms can be tabulated as given in Table 10.4. TABLE 10.4 Nutritional forms based on sources of carbon, energy and electrons Carbon Sources ‘Autotrophs . CO, sole or principal source of carbon. Heterotrophs .. ‘educed, preformed, organic molecules from other organisms. Energy Sources Phototrophs. Light. Chemotrophs xidation of organic or inorganic compounds, Electron Sources Lithotrophs Reduced inorganic molecules. - Organotrophs . ganic molecules, Taking the sources of carbon, energy and electrons together, the nutritional forms of microorganisms can be classified as given in Table 10.5. Scanned with CamScannerA summarised account of nutrition and the various major nutritional forms of ganisms are given with examples in Fig. 10.1. TABLE 10.5 jonal forms of microorganisms defined in terms of energy, electrons and carbon sources microors Major nu! Nutritional Forms “Sources of energy, electrons and carbon 1, Photolithoautotroph: Light energy (photolithoautotrophy) Inorganic electron donor CO) carbon source themical energy source (inorganic) 2, Chemolithoautotrophs (chemolithoautotrophy) Inorganic electron donor CO; carbon source 3, Photoorganoheterotrophs.. _.Light energy (photoorganoheterotrophy) Organic electron donor Organic carbon source (COz may also be used) 4, Chemoorganoheterotrophs ... _.Chemical energy source (organic) (chemoorganoheterotrophy) Organic electron donor Organic carbon source 10.3.1. Photolithoautotrophs ‘This group includes photosynthetic microalgae, (purple sulphur bacteria and green sulphur bacteria). : In photosynthetic microalgae and cyanobacteria the external energy-source is light. One or more varieties of chlorophyll are present to trap the solar energy. Such microorganisms are, therefore, largely green. The hydrogen-source of all photoautotrophic microalgae and cyanobacteria is environmental water (H,0) which is split into oxygen and hydrogen with the help of light energy. The oxygen is released as byproduct and the free hydrogen liberated, as a result of photolysis of water reduces carbon dioxide (the inorganic raw material) to carbohydrate (organic metabolite food) which is used as food. This pattern of food production may be symbolized as follows : cyanobacteria, and photosynthetic bacteria Light —— Chlorophyll ——> Energy O (by product) H,O a CO; “> Carbohydrate Scanned with CamScannerINORGANIC METABOLITES NUTRIENTS JAUL INoRGANIG METABOUTES| ORGANIC METABOLITES Mics General Microbiology LIVING BODY OF MICROORGANISMS SUPPLIED BY ENVIRONMENT oe (Foo0 MANUFACTURED FROM {F000 ABSORBED FROM INORGANIG RAW MATERIAL PREFABRICATED ORGANIC Guiote ‘SOURCES). ‘SouRce) Sources —> ; AUTOTROPHS HETEROTROPHS PHOTOLITHOAUTOTROPHS | PHOTOORGANOHETEROTROPHS | usit—>] — (PHOTOAUTOTROPHS) (PHOTOHETEROTROPHS) z PURPLE SULPHUR BACTERIA, GREEN PURPLENON.SULPHUR BACTERIA 2 ‘SULPHUR BACTERIA, CYANOBACTERIA & 1y [MICROALGAE (EXCEPT COLOURLESS ONES) Sounces : CHEMOLITHOAUTOTROPHS | CHEMOORGANOHETEROTROPHS (CHEMOAUTOTROPHS) (CHEMOHETEROTROPHS) 2 2 2 CHEMICALS—>] SULPHUR BACTERIA, RON BACTERIA, | SAPROPHYTIC BACTERIA, SYMBIOTIC z HYDAOGEN BACTERIA, NITRIFYING. BACTERIA, FUNGI PROTOZOA, é BACTERIA ‘COLOURLESS MICROALGAE ELECTRON LITHOTROPHS ORGANOTROPHS ounces | (REDUCED INORGANG MOLECULES) (ORGANIC MOLECULES) Fig. 10.1. Schematic representation of nutrition and major nutritional forms of microorganisms:"* The hydrogen-source of photoautotrophic bacteria is not water, and oxygen is never a byproduct of photosynthesis. These bacteria are adapted to live in sulphur springs and other sulphurous regions where. hydrogen sulphide (HS) is normally available. This compound generally serves as the hydrogen-source. Two families of bacteria belong to photoautotrophic group, the purple sulphur bacteria (e.g., Chromatium, Thiospirillum) and the green sulphur bacteria (e.g., Chlorobium, Chloropseudomonas, Chlorobacterium). These bacteria Possess a variety of special chlorophyll known as bacteriochlorophyll. It is green but its colour is masked by the additional yellow carotenoids which are also present. For the photoautotrophic bacteria, therefore, the special pattern of photosynthesis becomes : Scanned with CamScannerSS Microbial Nutrition 193 Light ——> Bacteriochlorophyll ——» Energy ‘S(elemental sulphur) H, Hy co, Carbohydrate Elemental sulphur is the byproduct. It is stored inside the cells in the purple sulphur bacteria, and is excreted from the cells in the green sulphur bacteria. 10.3.2. Chemolithoautotrophs (chemoautotrophs) This is a group of nonphotosynthetic autotrophic microorganisms consisting entirely of bacteria. They cannot use light, and their external energy sources in food manufacture are a variety of inorganic metabolites absorbed from the environment. In the most cases, these metabolites are combined with molecular oxygen in the cells, resulting in release of energy (exothermic reaction) and a variety of inorganic byproducts. Water and carbon dioxide are the inorganic raw materials in subsequent food manufacture. The concept of chromoautotrophy (chemolithotrophy) was formulated by Winogradsky. By studying Beggiatoa, he demon- strated for the first time that a living organism could oxidize H,S to elemental sulphur and then to SOZ-. This process of manufacturing food is called chemosynthesis. The general pattern is as follows. Inorganic Metabolites —0_, Inorganic Byproducts Energy (by product) H,0 HH, co, ———— Carbohydrate ‘Among the best known chemoautotrophic microorganisms are the sulphur-oxidizing bacteria, the iron-oxidizing bacteria, the nitrifying bacteria, and the hydrogen-oxidizing bacteria. 10.3.2.1, Sulphur-oxidising Bacteria Sulphur-oxidising bacteria are those chemoautotrophs that oxidize sulphur compounds as electron donors and energy-releasing compounds. The sulphur compounds used by them are H,S, elemental sulphur (S), S;03 and SO3. The best studied sulphur-oxidising chemoautotroph is the genus Thiobacillus that contains several gram-negative and rod-shaped bacteria such as T. thioparus , T. denitrificans, T. neopolitanus, T. thioxidans and T. intermedius. Other sulphur-oxidising chemoautotrophs are Acromatium, Beggiatoa, Thiothrix, Thioploca, Thiomicrospira, Thiosphaera, Thermothrix and Thiovulum. The general Pattern of chemoautotrophy used by these bacteria is the following. ~~~Scanned with CamScanner194 General, H,S, S, 8,0 7,S05 +0 _, so} energy 40 (by product) HO ‘HITE co, Carbohydrate Several species of Thiobacillus are acidophilic as they generate large amounts of sulphuric acid during the reactions. One such species, 7. ‘ferrooxidans, can also grow chemoautotrophically by the oxidation of ferrous iron and is a major biological agent for the oxidation of this metal. Achromatium commonly occurs in freshwater sediments containing sulphide. Cells of Achromatium store elemental sulphur, like Chromatium (a photoautotrophic bacterium), internally inside the granules that later disappear 2$ sulphur is baidised to sulphate. Most species of Beggiatoa can obtain energy from the oxidation of inorganic sulphur compounds ba Tek emer of Calvin xl thus require organic compounds as carbon source, Such a nutritional lifesty! called mixotrophy. Beggiatoa, the filamentous gliding sulphur-oxidising bacteria, occurs in nature mainly in HS -rich sulphur springs, decaying seeweed beds, mud layers of lakes, and waters polluted with sewage. In these habitats the filaments of Beggiatoa are usually filled with sulphur granules. An interesting habitat of Beggiatoa is the rhizosphere of plants (rice, cattails, and other swamp plants) living in flooded, and hence anoxic, soils. Such plants pump oxygen down into their foots so a sharply defined oxic/anoxic boundry develops between the root and the soil. Beggiatoa (and probably other sulphur-oxidising bacteria) develop at this boundry, and plays a beneficial role for the plant by oxidising (as thus detoxifying) the H,S. 10.3.2.2. Iron-oxidising Bacteria Iron-oxidising bacteria cause aerobic oxidation of iron from ferrous (Fe%) to ferric (Fe) state that yields energy. Ferric (Fe*) compounds are soluble, whereas the ferric (Fe?) compounds, eg. ferric hydroxide (Fe(OH),] are insoluble, The insoluble ferric hydroxide precipitates in water and imparts reddish colour to it. However, the general pattern of Pitritional lifestyle of iron-oxidising bacteria can be represented as : co) Soluble ferrous (Fe2*) compounds — Insoluble ferric (Fe*) compounds Energy O (by product) H,0 Cette CO, Carbohydrate Thiobacillus ferroxidans, Leptospirillum, Gallionella, Ferrobacillus, Leptothrix, and Cladothrix are the representatives of iron-oxidising bacteria, The best studies ones are 7. ferrooxidans and Leptospirillum ferrooxidans, which are very common in acid polluted ‘environments such as coal-mining dumps. Other iron-oxidising bacteria are commonly found in water-logged soils, bogs, and anoxic lake sediments. One of the most common forms of iron in nature is pyrite (FeS,) , which is formed from the reaction of sulphur with ferrous sulphide (FeS) to form a highly insoluble crystalline to Scanned with CamScannerMicrobial Nutrition - 195 s very common in bituminous coals and in many ore bodies. The bacterial 1 ardation of pyrite is very significant in the development of acidic conditions in mining operations. Additionally, oxidation of pyrite by iron-oxidising bacteria is of considerable ~ Significance inthe process called microbial leaching of ores. \ 10.3.2.3. Nitrifying Bacteria | Nitrifying bacteria are those chemoautotrophic bacteria that grow at the expense of reduced” inorganic nitrogen compounds. No nitrifying bacterium is known that carries out the complete oxidation of ammonia (NH,) to nitrate (NO 3); thus, nitrification in nature results from the sequential action of two separate groups of nitrifying bacteria, the ammonia- oxidising bacteria (the nitrosifyers) and the nitrite-oxidising bacteria or true nitrifying (nitrate -producing) bacteria. Nitrifying bacteria are widespread structure, and i in soil and water and occur in highest numbers in those environments where considerable amounts of ammonia is present (e.g., sites where extensive protein decomposition takes place and in swage treatment facilities). Nitrifying bacteria flourish especially in lakes and streams that receive inputs of sewage or other waste waters | because these are frequently rich in ammonia. Most of the nitrifying bacteria are obligate chemoautotrophs (chemolithotrophs). Species of Nitrobacter are an exception as they may grow chemoheterotrophically (chemoorganotrophically) on acetic acid or pyruvic acid as sole carbon and energy source. ‘Ammonia-oxidising bacteria (the nitrosifyers) The ammonia-oxidising bacteria or nitrosifying bacteria typically have genus names beginning in “Nitroso”. Nitrosomonas, Nitrosococcus, Nitrosospira, Nitrosolobus, and Nitrosovibrio are the major genera of nitrosofyers. These bacteria possess a key enzyme (ammonia monooxygenase) that oxidises ammonia (NH;) to hydroxylamine (NH,OH), which then is oxidised to nitrite (NO) by nitrosifying bacteria. The general pattern of nutritional | lifestyle of nitrosifyers is as follows : | ‘Ammonia (NH;) —P-— Nitrite (NO3) Energy 0 (by product) HO ee co, Carbohydrate Nitrite-oxidising bacteria (the true nitrifying bacteria) ‘The nitrite-oxidising bacteria (the true nitrifying bacteria or nitrate-producing, bacteria) have genus names begining in “Nitro”. Nitrobacter, Nitrospira, Nitrococcus, and Nitrospira are the main representative genera of nitrate-producing bacteria. These bacteria oxidize nitrite (NO;) produced by nitrosifyers to nitrate (NCS). The general pattem of nutritional lifestyle of nitrite -oxidising bacteria is as follows: 10.3.2.4, Hydrogen-oxidising Bacteria Hydrogen-oxidising bacteria represent the group of bacteria that oxidise H, (the sole electron donor) and reduce O; (the electron acceptor) via “knallgas” reaction, the reduction of eee etn Scanned with CamScanner196 Nitrite (NO3) ao (Nitrite (NO5) Enersy 6 (by product) H,O H, co, FS carbonyarate O, with H). This reaction yields energy (-273 kJ/reaction), which is used in CO, fixation. The best studied genera of this These bacteria are both gram-positive and gram-negative. 4 group of bacteria are Ralstonia, Pseudomonas, Paracoccus, and Alkaligenes; others are ‘Acidovorax, Aquaspirillum, Hydrogenophaga, Hydrogenobacter, Bacillus, Aquifex, and Mycobacterium. All hydrogen-oxidising bacteria possess one or more hydrogenase enzymes that function to bind hydrogen (Hz) and use it either to generate ATP or for reducing power for chemoautotrophic growth. Most hydrogen-oxidising bacteria flourish best under microaerobic conditions when growing chemoautotrophically (chemolithotrophically) on hydrogen because hydrogenases are oxygen-sensitive enzymes. Typically, oxygen levels of about 5-10% support best growth of these bacteria. The general pattern of the nutritional life- style of chemoautotrophic (chemolithotrophic) hydrogen-oxidizing bacteria is as follows: Hydrogen (H,) a) H,0 Energy (byproduct) H co, ae ‘Almost all hydrogen-oxidising bacteria are facultative chemoautotrophs, i.e, they can also grow chemoheterotrophically (chemoorganotrophically) with organic compounds as energy source. This means that the hydrogen-oxidising bacteria have ability to switch between chemoautotrophic and chemoheterotrophic (chemoorganotrophic) modes of metabolism and generally do so in nature whenever required. This is a major distinctio a oxidising bacteria and many sulphur-oxidising bacteria or ni A latter two groups are obligate chemoautotrophs their growth fails to occur in the absence of the inorganic energy source). 10.3.3. Photoorganoheterotrophs (Photoheterotrophs) Some bacteria use light as energy source with organic compounds as the carbon source 10 grow. These bacteria are called photoorganoheterotrophs (photoheterotrophs) and belong '0 urple nonsulphur bacleria.. The Tatter have Geen called “nonsulphur” because it the group of pi was originally thought that they were unable to use sulphide as an electron donor for the reduction of CO, to cell material. Recent studies clarified that most species of Hoomr or Scanned with CamScannerand Microbiol Nutrition 00 can use sulphide although the level of sulphide utilised by them is quite low than that by purple sulphur bacteria. Most purple nonsulphur bacteria have additional ability to grow aerobically in darkness utilising organic compounds as electron donor. It is their great ability to practice photoheterotrophy that likely accounts for their competitive success in nature. Purple nonsulphur bacteria are typically nutritionally diverse with respect to photoheterotrophy as they can use fatty, organic, or amino acids; sugars; alcohols; and even aromatic compounds like benzoate as carbon sources. Purple nonsulphur bacteria possessing additional ability to photoorganoheterotrophy (e.g, Rhodopseudomonas, Rhodospirillum. Rhodobacter, Rhodovulum, Rhodopila, Rhodobaca, Rhodocyclus, Rhodoferax, Phaeospirillum, Roseospira, Roseospirillum, Rubrivivax, Rhodoplanes, Rhodobium, Rhodomicrobium) are considered to be intermediate, between photolithoautotrophs and chemoorganoheterotrophs. The general pattern of nutritional Tifestyle of these bacteria can be represented as : ‘Light —> Bacteriochlorophyll——> Energy Organic byproduct Organic compounds H, Carbohydrate 10.3.4. Chemoorganoheterotrophs Majority of heterotrophic microorganisms belong to this nutritional category. Since they cannot synthesize their own food (organic substances) they obtain it directly from external environment using chemical energy-source. A clear-cut distinction between the carbon-source and the energy-source, characteristic of the three preceding nutritional forms, loses its clarity in the context of chemoorganoheterotrophs. In the latter, both carbon and energy can usually be derived from the metabolism of a single organic compound. The chemoorganoheterotrophs include protozoa, fungi (including slime molds), and the great majority of bacteria. The chemoorganoheterotrophic form of nutrition can further be divided into two categories on the basis of the physical state in which organic nutrients enter the cell. These categories are: holotrophic nutrition and absorptive nutrition. Scanned with CamScanner