Alkaline and Saline Soils •Saline soils occur in soils with pH>8.5 •Ca2+, Mg2+, K+ and Na+ do not produce acid upon reacting with water •The do not produce OH- ions either, but in soils with pH>8.5, there are higher concentrations of carbonate and bicarbonate anions (due to dissolution of certain minerals) CaCO3 Ca2+ + CO32or CO32- + H2O HCO3- + OHHCO3- + H2O H2CO3 + OHH2CO3 H2O + CO2(gas) NaCO3 2Na2+ + CO32- •pH rises more for most soluble minerals (eg. NaCO3) •pH rise is limited by the common ion effect Micronutrient deficiencies in saline soils •Fe and Zn deficiencies are common because their solubility is extremely low in alkaline conditions •Addition of inorganic fertilizer may not improve this deficiency as they quickly become tied up in insoluble forms •Chelate compounds are often applied to soils (Fe associated with organic compounds) • Under high pH, B tightly adsorbs to clays in an irreversible set of reactions. In sandy soils, B content is generally low under any pH level, especially in wet environments due to leaching (problematic in wet or dry environments, but less so in between). Effect of soil pH on nutrient content and soil microorganisms •Phosphorus is often deficient in alkaline soils, because it is tied up in insoluble calcium or magnesium phosphates [eg. (Ca3(PO4)2 and Ca3(PO4)2] •Some plants excrete organic acids in the immediate vicinity of their roots to deal with low P Other notes of interest: •Ammonium volatilization is commonly problematic during nitrogen fertlization on alkaline soils (changes to gas – lost to atmosphere) •Molybdenum levels are often toxic in alkaline soils of arid regions Salinization The process by which salts accumulate in the soil Soil salinity hinders the growth of crops by lowering the osmotic potential of the soil, thus limiting the ability of roots to take up water (osmotic effect). Plants must accumulate organic and inorganic solutes within their cells. + Specific ion effect: Na+ ions compete with K Soil structure breaks down, leading to poor oxygenation and infiltration & percolation rates •36% of prairie farmland has 1-15% of its lands affected by salinization and 2% has more than 15% of its lands affected. •Most prairie farmland (61% in Manitoba, 59% in Saskatchewan, and 80% in Alberta) has a low chance of increasing salinity under current farming practices. Conservation farming practices to control soil salinity •Reducing summerfallow •Using conservation tillage •Adding organic matter to the soil •Planting salt-tolerant crops (eg., canola and cabbage) Conditions promoting salinization: •the presence of soluble salts in the soil •a high water table •ET >> P These features are commonplace in: •Prairie depressions and drainage courses •At the base of hillslopes •In flat, lowlying areas surrounding sloughs and shallow water bodies. •In areas receiving regional discharge of groundwater. Signs of Salinization A. Irregular crop growth on a solonetz Source: Agriculture and Agri-food Canada Whitish crust of salts exposed at the surface (B,C) Aerial photo of saline deposits at Power, Montana D. Presence of salt streaks within soils E. Presence of salt-tolerant native plants, such as Red Sapphire Human activities can lead to harmful effects of salinization, even in soils of humid regions Effect of road salt on Maple leaves (a) (b) Calcium carbonate accumulation in the lower B horizon The white, rounded "caps" of the columns are comprised of soil dispersed because of the high sodium saturation Salinization in response to conversion of natural prairie to agriculture Measuring the electrical conductivity (EC) of a soil sample in a field of wheatgrass to determine the level of salinity. A portable electromagnetic (EM) soil conductivity sensor used to estimate the electrical conductivity in the soil profile Effect of salinity on soybean seedlings Influence of irrigation technique on salt movement and plant growth in saline soils The Importance of Soil Nitrogen Amino acids Enzymes Proteins that catalyze chemical reactions in living organisms Nucleic Acids A nucleic acid is a complex, high-molecularweight, biochemical macromolecule composed of nucleotide chains that convey genetic information. Nitrogen Deficiency •Pale, yellowish-green colour due to low chlorophyll content •Older leaves turn yellow first and may senesce prematurely •Spindly stems or few stems •Low protein, but high sugar content (not enough N to combine with carbon chains to produce proteins) •Low shoot:root ratio •Rapid maturity Nitrogen Deficiency “Chlorosis” Yellowing of older foliage Restricted growth Few stems or spindly stems Nitrogen oversupply •Lodging with wind or heavy precipitation due to excessive growth •Delayed maturity •Susceptibility to fungal diseases •Reduced flower production •Poor fruit flavour •Low vitamin and sugar content of fruits and vegetables Nitrogen Forms Reduced NH4+ N2 N 2O NO ammonia molecular N nitrous oxide nitric oxide Oxidized NO2- NO2- NO3- nitrite nitrogen dioxide nitrate Nitrogen Fixation The nitrogen molecule (N2) is very inert. Energy is required to break it apart to be combined with other elements/molecules. Three natural processes liberate nitrogen atoms from its atmospheric form •Atmospheric fixation by lightning •Biological fixation by certain microbes — alone or in a symbiotic relationship with plants •Industrial fixation Atmospheric fixation by lightning •Energy of lightning breaks nitrogen molecules. •N atoms combine with oxygen in the air forming nitrogen oxides. •Nitrates form in rain (NO3-) and are carried to the earth. •5– 8% of the total nitrogen fixed in this way (depends on site) Industrial Fixation •Under high pressure and a temperature of 600°C, and with the use of a catalyst, atmospheric nitrogen and hydrogen (usually derived from natural gas or petroleum) is combined to form ammonia (NH3). •Ammonia can be used directly as fertilizer, or further processed to urea and ammonium nitrate (NH4NO3). Biological Fixation Performed mainly by bacteria living in a symbiotic relationship with plants of the legume family (e.g., soybeans, alfalfa), although some nitrogen-fixing bacteria live free in the soil. •Biological nitrogen fixation requires a complex set of enzymes and a huge expenditure of ATP. Although the first stable product of the process is ammonia, this is quickly incorporated into protein and other organic nitrogen compounds. Carried out by Rhizobium bacteria in a SYMBIOTIC relationship. Host provides carbohydrates for energy; Rhizobium supplies plant with fixed nitrogen. Nitrogen mineralization 95-99% of N is in organic compounds, unavailable to higher plants, but protected from loss 1. Soil microbes attack these organic molecules, (proteins, nucleic acids, amino sugars, urea), forming amino compounds 2. The amine groups are hydrolyzed, with N released as NH4+ (ammonium ions; See pg. 548) 3. Oxidation of NH4+ to NO2- and NO3The reverse process (incorporation of NO3- or NH4+ into soil micro-organisms) is called immobilization Nitrification •Bacteria of the genus Nitrosomonas oxidize NH3 to nitrites (NO2−). •Bacteria of the genus Nitrobacter oxidize the nitrites to nitrates (NO3−). Soil Organic Nitrogen Organic (as opposed to mineralized) nitrogen has variable structure (still poorly understood) Most SON uptake occurs after mineralization of SON to NO3- or NH4+ Plants may also take up SON directly, or the N can be assimilated by mychorrizal associations Ammonium fixation by clay minerals Occurs more in the subsoil than in the topsoil Ammonium may become ‘fixed,’ or entrapped within Cavities of the crystal structure of vermiculites, micas and smectites Ammonia volatilization NH4+ + OH- H2O + NH3(gas) Occurs more in soils with high pH, especially when drying and when temperatures are high Soil colloids (clay and humus) inhibit ammonia volatilization through adsorption Nitrate leaching Negatively-charged nitrate ions are not adsorbed by colloids, so they move freely with drainage water Result: (i) impoverishment of soil N; (ii) environmental problems (eutrophication) especially in heavily irrigated zones with N-fertilizer application or manure Denitrification •Denitrification reduces nitrates to nitrogen gas, thus replenishing the atmosphere. Performed by bacteria in anaerobic conditions. They use nitrates as an alternative to oxygen for the final electron acceptor in the respiration process. Nitrogen Storage in Soils • Current levels of nitrogen in soils reflect the accumulation of N in the organic fraction over long periods of time. • Only about 1.5-3% of the N stored is used on an annual basis. • Over long time frames N is stable in natural ecosystems (dynamic equilibrium established between losses and additions) Soil Phosphorous and Potassium Why is phosphorus so important? 1. Essential component of ATP (adenosine triphosphate) Molecular currency of intercellular energy transfer Used as energy source during photosynthesis and cellular respiration Consumed by many enzymes in metabolic reactions and during cell division. * Notice the 5 N and 3 P in the ATP molecule 2. Incorporated into nucleic acids DNA and RNA Genetic instructions for the development and functioning of all living organisms Sugar-phosphate backbone 3. Phospholipid bilayer Cell membranes, composed of a phospholipid bilayer, control what goes into and out of a cell Phospholipid Active transport across the cell membrane requires ATP Phospholipid bilayer The Importance of Phosphorus P is involved in: P deficiency: Photosynthesis Nitrogen fixation Flowering Fruiting & fruit quality Maturation Root growth Tissue strength Stunting Thin stems Bluish-green leaves Delayed maturity Sparse flowering Poor seed quality •Similarly to nitrogen deficiency, the older leaves are often first affected •P deficiency is often difficult to diagnose as visual changes are subtle The Phosphorus Problem in Soil Fertility 1. The total P content of soils is low. 200-2000 kg/ha in uppermost 15 cm (topsoil) 2. Phosphorus compounds found in soils are often highly insoluble 3. When soluble sources are added (fertilizers and manure) they often become fixed into insoluble compounds • 10-15% of P added is taken up by crop in year of application • Overfertilization for decades has led to saturation of the P-fixation capacity (large P reserves in N. American soils) • In contrast, P deficiency is a serious problem in sub-Saharan Africa (removal repeatedly has exceeded addition) N, P and K Fertilizer Use in USA Figure 14.1 Impact of Phosphorus on Environmental Quality 1. P deficiency: Land degradation Little P is lost in natural ecosystems as P cycles between living biomass and soils Once cleared for agriculture: (i) Soil erosion loss (ii) Biomass removal • P-supplying capacity decreases, even if total P is sufficient • Nodulation is affected by P-deficiency, thereby promoting N-deficiency Most problematic in most highly weathered soils • Warm, moist environments of the tropics • Oxisols & Andisols • Low availability of P when in association with Fe & Al • Lots of P needs to be applied to Andisols (Fig 14.20) Combined P & N deficiency limits biomass and promotes further erosion Water Quality Degradation due to Excess P (and N) Point sources Sewage outflows (phosphates in soaps) Industries Non-point sources Runoff water Eroded sediment from soils in affected watershed “Too much of a good thing” The Phosphorus Cycle Phosphorus in the soil solution •Very low concentrations (0.001 to 1 mg/L) •Roots absorb phosphate ions, HPO42- (alkaline soils) and H2PO4- (acid soils) Uptake by Roots •Slow diffusion of phosphate ions to root surfaces •Mychorrizal hyphae extend outward several cm from root surface •P can then be incorporated into plant tissues (Fig 14.9) •Soil P replenished by plant residues, leaf litter, and animal waste •Soil microorganisms can temporarily incorporate P into their cells •Some soil P gets tied up in organic matter (storage & future release) Available P seldom exceeds 0.01% of total soil phosphorus Forms of Soil Phosphorus Organic phosphorus Calcium-bound phosphorus (alkaline soils) Iron-bound phosphorus (acid soils) Aluminium-bound phosphorus (acid soils) •Low solubility – not readily available for plant uptake •P is slowly released from each of these types of compounds •Leaching loss is low, but can play a role in eutrophication •Unlike N, P is not generally lost in a gaseous form Gains and Losses •Losses from plant removal, erosion of P-containing soil particles and dissolved P in surface runoff water •Gains from atmospheric dust are very limited, but a balance is established in most natural ecosystems Leaching of P after saturation of fixed pool Figure 14.22 Potassium •The nutrient third most-likely to limit productivity •Present in soils as K+ ion (not in structures of organic compounds) •Soil cation exchange and mineral weathering dominate its exchange and availability (as opposed to microbiological processes) •Causes no off-site environmental problems •Igneous rocks are a good source – alkaline soils keep it. •Activates certain enzymes. •Regulates stomatal opening •Helps achieve a balance between negatively and positively charged ions within plant cells. •Regulates turgor pressure, which helps protect plant cells from disease invasion. •Promotes winter-hardiness and drought-tolerance Potassium deficiency •Leaves yellow at tip (chlorosis) and then die (necrosis) The leaves, therefore, appear burnt at the edges and may tear, leaving a ragged edge •White, necrotic spots may appear near leaf edges •Oldest leaves are most affected The Potassium Cycle •High concentrations in micas and feldspars K between 2:1 crystal layers becomes available •Returned to soil through leaching from leaves and from plant residue decomposition •Some loss by eroded soil particles and leaching •Replenishment required in most agroecosystems (1/5 of plant K is typically removed in product). Excess in plants can cause a dietary imbalance in ruminants Calcium Vast reserves in calcareous (chalk) soil. •Calcium is a part of cell walls and regulates cell wall construction. •Cell walls give plant cells their structural strength. •Enhances uptake of negatively charged ions such as nitrate, sulfate, borate and molybdate. •Balances charge from organic anions produced through metabolism by the plant. •Some enzyme regulation functions. Magnesium Reserves in magnesium limestone. Magnesium is the central element within the chlorophyll molecule. It is an important cofactor the production of ATP, the compound which is the energy transfer tool for the plant. Sulphur Found in rocks and organic material. Sulphur is a part of certain amino acids and all proteins. It acts as an enzyme activator and coenzyme (compound which is not part of all enzyme, but is needed in close coordination with the enzyme for certain specialized functions to operate correctly). It is a part of the flavour compounds in mustard and onion family plants. Boron Boron is important in sugar transport within the plant. It has a role in cell division, and is required for the production of certain amino acids, although it is not a part of any amino acid. Manganese Manganese is a cofactor in many plant reactions. It is essential for chloroplast production. Copper Synthesis of some enzymes important in photosynthesis Copper is a component of enzymes involved with photosynthesis. Iron Iron is a component of the many enzymes and light energy transferring compounds involved in photosynthesis. Zinc Zinc is a component of many enzymes. It is essential for plant hormone balance. Molybdenum Molybdenum is needed for the reduction of absorbed nitrates into ammonia prior to incorporation into an amino acid. It performs this function as a part of the enzyme nitrate reductase. Molybdenum is also essential for nitrogen fixation by nitrogenfixing bacteria in legumes. Responses of legumes to Molybdenum application are mainly due to the need by these symbiotic bacteria.