Mineral Nutrients I. Introduction A. Definition B. Evidence 1. Julius Sachs Experiment Fig 37.7 Julius Sachs 1860’s C. Plant Mineral Composition 1. Incorporation a. As is= Some minerals can be used as is: e.g. K+ ions for guard cell regulation b. Combined = Some minerals have to be incorporated into other compounds to be useful: e.g. Fe+ in the cytochrome complex of the light reactions c. Altered = Some mineral compounds have to be altered to be useful: NO3- must be converted to NH4+ inside the plant d. Water i. 80–85 % of an herbaceous plant is water. ii. Water supplies most of the hydrogen and some oxygen incorporated into organic compounds by photosynthesis. iii. But > 90% of the water absorbed is lost by transpiration. iv. Water’s primary function is to serve as a solvent. v. Water also is involved in cell elongation and turgor pressure regulation 2. Dry weight a. 95% “organic” – C, H, & O from air & water, assimilated by photosynthesis b. 5% inorganic minerals II. Categories A. Essential Nutrients Nutrients that are required for a plant to grow from a seed and to complete its life cycle. 1. Types: a. Macronutrients Elements required by plants in relatively large amounts. i. Elements CHOPKNS ii. Functions Ca Mg Category Form Available Uses Carbon CO2 Organic compounds Hydrogen H20 Organic compounds Oxygen CO2 (air), O2 (soil) Organic compounds Phosphorus H2PO4-, HPO42- Nucleic acids, Phospholipids, ATP Potassium K+ Water Balance (stomata), Protein Synthesis Nitrogen NH4+, NO3- Proteins, Nucleic acids, Hormones, Chlorophyll Sulfur SO42- Proteins Calcium Ca2+ Cell walls & Membranes, Enzyme Activation Magnesium Mg2+ Chlorophyll, Enzyme Activation Information taken from Table 37.1 b. Micronutrients These elements are required by plants in relatively small amounts (<0.1% dry mass). i. Elements Fe, B, Cl, Mo, Cu, Mn, Ni, & Zn ii. Functions Category Form Available Uses Chlorine Cl- Required for photosystem II to split water and water balance Iron Fe3+ or Fe2+ Component of cytochromes and enzyme activation Manganese Mn2+ A.A. formation, enzyme activation, and split water in PS II Boron H2BO3- Cofactor in chlorophyll synthesis, involved in carbohydrate transport and nucleic acid synthesis, and role in cell wall function Zinc Zn2+ Cofactor in chlorophyll synthesis and enzyme activation Copper Cu+ or Cu2+ Involved with re-dox and lignin biosynthesis enzymes Nickel Ni2+ Cofactor in nitrogen metabolism enzymes Molybdenum MoO4-2 Essential for mutualistic relationship with nitrogen fixing bacteria and cofactor for nitrogen reducing enzymes Information taken from Table 37.1 III. Mineral Deficiencies A. Dependent on: 1. the role of the nutrient in the plant 2. its mobility B. Immobile Nutrients 1. Once they have been incorporated into plant tissue, they remain (can’t return to phloem). 2. Boron, calcium, and iron 3. Growth = normal until the mineral is depleted from soil; new growth suffers deficiency and thus youngest tissues show symptoms first. WHY? C. Mobile Nutrients 1. can be translocated by phloem to younger (actively growing) tissue. 2. Cl, Mg, N, P, K, and S 3. When mineral is depleted, nutrients translocated to younger tissue. 4. Thus older tissues show deficiency & then die What is the adaptive value of nutrient mobility? D. Criteria 1. Not common in natural populations. Why? Plants have adapted to soil components. 2. Common in crops & ornamentals. Why? Human selection is for biggest, fastest plants. Need more nutrients than the soil provides. Crop growth depletes the soil because no organic matter is returned. 3. Deficiencies of N, P, and K are the most common. 4. Shortages of micronutrients are less common and often soil type specific. 5. Overdoses of some micronutrients can be toxic. E. Symptoms 1. Chlorosis – leaves lack chlorophyll: yellow, brittle, papery. Typically lack of N or Fe. 2. Necrosis – the death of patches of tissue 3. Purpling – deficiency of N or P, causes accumulation of purple pigments 4. Stunting – lack of water, N Soils I. Soil Formation A. Forces II. Soil Horizons B. Characteristics A. Names Fig 37.2 III. Orders A. Definition B. Primary C. Locations IV. Soil Properties A. Chemistry 1. Minerals 2. Nitrogen–fixing bacteria 3. Mycorrhizal fungi 4. Water 5. Oxygen (Gases) B. Composition 1. Chemistry – determines which minerals are present and available, thus affecting plant community composition 2. Physical nature – affects porosity, texture, density of soil, which affects #1 3. Soil organisms – decomposition & mineral return. Interact with roots to make nutrients available Nitrogen! The only mineral that the plant can ONLY get from reactions mediated by soil organisms. C. Texture 1. Soil is created by weathering of solid rock by: water freeze/thaw, leaching of acids from organic matter, carbonic acid from respiration + water. 2. Topsoil is a mixture of weathered rock particles & humus (decayed organic matter). 3. Texture: sand silt clay Large, spaces for water & air Small, more SA for retaining water & minerals V. Topsoil A. Characteristics 1. Biotic = Bacteria, fungi, insects, protists, nematodes, & Earthworms! Create channels for air & water, secrete mucus that binds soil particles 2. Humus: reservoir of nutrients from decaying plant & animal material 3. Bacterial metabolism recycles nutrients B. Nutrient Availability 1. Cations in soil water adhere to clay particles (negatively charged surface) 2. Anions do not bind; thus they can leach! (NO3, HPO4, SO4) 3. Cations become available for root uptake by cation exchange – H+ displaces cations on the soil particle surface 4. H+ from carbonic acid – formed from water + CO2 released from root respiration 5. Humus – negatively charged & holds water & nutrients. Thus very important in the soil! Fig 37.3 C. Soil pH 1. Low pH (acidic) = high H+ concentration a. More cations released b. Too much acid – cations leach…..mineral deficiency 2. High pH (basic) a. Not enough H+ for cation release….mineral deficiency VI. Soil conservation A. Factors Affecting 1. Natural systems: decay recycles nutrients 2. Agricultural systems: crops harvested, depleting soil of nutrients & water 3. Fertilizers: N:P:K a. Synthetic: plant-available, inorganic ions. Faster acting. i. Problem: ii. leaching, acidifying the soil b. Organic: slow release by cation exchange, holds water, thus less leaching B. Phytoremediation 1. Use of plants to extract toxic metals from soil 2. Benefits: easier to harvest the plants than to remove topsoil! VII. NITROGEN A. Why so important? 1. Air is 80% Nitrogen, but….. 2. Macronutrient that is most often limiting. Why? Is almost always taken up as anions (NO3-). 3. What’s it used for? Proteins (AAs), nucleic acids, chlorophyll production, and ??? The Nitrogen Cycle N2 N2 fixation Denitrification Uptake NO3 Organic N NH4 Leaching B. Nitrogen Cycle 1. Steps: a. N fixation – conversion of N2 to NH3 b. Ammonification – conversion of NH3 or organic N into NH4+ c. Nitrification – conversion of NH4+ to NO3d. N reduction – conversion of NO3- back to NH4+ within plant. e. N assimilation – incorporation of NH4+ into AAs, nucleic acids, lignin, others(?) of the plant All steps within the soil are mediated by bacteria!!!! Fig 37.10 a. Nitrogen Fixation Process This process is catalyzed by the enzyme nitrogenase, requires energy (ATP), and occurs in three ways: i. Lightening – converts N in air to inorganic N that falls in raindrops ii. Non-symbiotic – certain soil bacteria iii. Symbiotic with Legumes Legumes: peas, beans, alfalfa The legume/bacteria interaction results in the formation of nodules on roots Plant – gets ample inorganic N source Bacteria – gets ample carbon source Fig 37.12 Fig 37.11 iii. Fixation in Non-legumes Here in the NW: alder Azolla (a fern) contains a symbiotic N fixing Cyanobacteria useful in rice paddies. Plants with symbiotic N fixers tend to be first colonizers. Why? b. Ammonification i. conversion of NH3 or organic N into NH4+ c. Nitrification i. Unfortunately NH4+ is a highly desirable resource for free–living bacteria, oxidizing it to NO3-. ii. Consequently the predominant form of N available to roots is NO3-. d. Nitrate Reduction i. NO3- must be reduced back to NH4+ in order to be incorporated into organics. ii. This process is energetically expensive but required. e. Nitrogen Assimilation i. The actual incorporation of NH4+ into organic molecules in the plant body. ii. This process is similar to that of an electron transport chain. iii. Reduced N passes through a series of carriers that function repeatedly but in the long run are unchanged. iv. Usually occurs within the roots. 2. Loses a. Leaching – loss of NO3- by soil water movement b. Denitrification – conversion of NO3- back to N2 c. ??? C. Nutritional Adaptations a. Parasitic plants i. Extract nutrients from other plants Ex. Mistletoes on Douglas Fir & Ponderosa pine Ex. Indian pipe – parasite on trees via mycorrhizae Fig 37.15 http://www.nofc.forestry.ca/publications/leaflets/mistletoe_e.html http://cals.arizona.edu/pubs/diseases/az1309/ b. Carnivorous plants i. Digest animals & insects – why? Grow in soils lacking an essential nutrient ii. Motor cells! iii. Trap insects & secrete digestive juices Ex. Venus flytrap, pitcher plant, Darlingtonia Figure 37.16 c. Mycorrhizal relationships i. Fungus & plant roots ii. Fungus gets carbos iii. Plants get greater SA for water & phosphorus uptake iv. Almost all plant species! v. 2 types: Ectomycorrhizae – hyphae form dense sheath over root; extend into cortex & out into soil. Thickened roots of woody plants Endomycorrhize – microscopic, more common. Fig 37.13 Learning power will supplant physical power.