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We will study effects of stresses on plants and see where it leads us
Course Plan We will study effects of stresses on plants and see where it leads us Phytoremediation Plant products Biofuels Effects of seed spacing on seed germination Effects of nutrient deprivation Effects of stresses Climate/CO2 change Non-coding RNAs Biotechnology Plant movements: flytraps, mimosa, soybeans Carnivorous plants Stress responses/stress avoidance Plant signaling (including neurobiology) Flowering? Hormones? Plant pathology? Plant tropisms and nastic movements Root growth responses Metal toxicity? Circadian rhythms? Effects of magnetic fields? Effects of different colors of light on plant growth?
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We will study effects of stresses on plants and see where it leads us
Learn about plant stress Pick some stresses to study Decide which plants to study and how to assay the stress effects What to measure? Biomass Yield Proteins, pigments Secondary products Responses to stimuli Solar tracking Leaf closing Tropisms Flowering
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Plant Stress Won Senator Proxmire’s “Golden Fleece” award for wasteful government spending Water? Nutrients? Environment? Temp? pCO2? Pollution? Ozone, other gases? Herbicides, eg Round-Up, Atrazine? Light? Insects and other herbivores? Pathogens = bacteria, viruses, fungi
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Mineral Nutrition Studied by soil-free culture in nutrient solutions:
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Mineral Nutrition Soil-free culture Sand culture Hydroponics: immerse roots in nutrient solution Slanted film maintains [nutrients] & O2 Aeroponics sprays nutrient solution on roots
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Mineral Nutrition Macronutrients CHOPKNS CHOPNS make biopolymers
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Mineral Nutrition Macronutrients CHOPNS make biopolymers N deficiency mobile in plant, so old leaves go yellow first
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Mineral Nutrition P deficiency mobile in plant, so shows first in old leaves which turn dark green or purple
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Mineral Nutrition S deficiency mobile in plant, so shows first in old leaves which turn yellow
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Mineral Nutrition K deficiency mobile in plant, so shows first in old leaves which go yellow at edges.
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Mineral Nutrition Ca deficiency immobile in plant, so shows first in young leaves which show blotches of necrotic tissue Crucial role in water- splitting!
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Mineral Nutrition Fe deficiency immobile in plant, so shows first in young leaves which turn yellow while veins stay green
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Mineral Nutrition Mg deficieincy mobile in plant, so shows first in old leaves which brown off at edges
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Mineral Nutrition Mg deficieincy mobile in plant, so shows first in old leaves which brown off at edges One/ chlorophyll and also cofactor for many enzymes
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Mineral Nutrition Micronutrients: BNaCl B: cell elongation. NA metabolism immobile in plant, so shows first in young leaves
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Mineral Nutrition Micronutrients: BNaCl B: cell elongation. NA metabolism Na: PEP regeneration in C4, K substitute mobile in plant, so shows first in old leaves
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Mineral Nutrition Micronutrients: BNaCl B: cell elongation. NA metabolism Na: PEP regenerationin C4, K substitute Cl: water-splitting, osmotic balance mobile in plant, so shows first in old leaves
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Mineral Nutrition Micronutrients: BNaCl others include Cu, Zn, Mn B: cell elongation. NA metabolism Na: PEP regeneration, K substitute Cl: water-splitting, osmotic balance Cu: cofactor immobile in plant, so shows first in young leaves
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Mineral Nutrition Micronutrients: BNaCl others include Cu, Zn, Mn Mn: cofactor for many enzymes, especially water splitting enzyme Immobile in plants so shows first in young leaves
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Mineral Nutrition Commercial fertilizers mainly supply NPK
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Plant food 2.6% ammoniacal nitrogen 4.4% nitrate nitrogen
9% phosphorus 5% potassium calcium (2%) magnesium (0.5%) sulfur (0.05%) boron (0.02%) chlorine (0.1%) cobalt (0.0015%) copper (0.05%) iron (0.1%) manganese (0.05%) molybdenum (0.0009%) nickel (0.0001%) sodium (0.10%) zinc (0.05%)
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Nutrient excesses More infrequent since plants control nutrient uptake
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Nutrient excesses More infrequent since plants control nutrient uptake Na stress: frequently occurs in areas that have been irrigated or are near oceans
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Effects of excess Sodium
Reduced turgor due to lower Ys Reduced growth rate Plasmolysis
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Effects of excess Sodium
Reduced turgor due to lower Ys Enzyme inhibition [Na] > 100 mM inhibit enzymes of all spp Even spp that live in 5 M NaCl
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Effects of excess Sodium
Reduced turgor due to lower Ys Enzyme inhibition Accumulation in old leaves to toxic levels
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Mechanisms of Sodium Tolerance
Exclusion negative correlation with HKT1 (unloads Na from xylem)and salt tolerance in Arabidopsis Accumulation of Na+ in leaf promotes Na tolerance!
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Mechanisms of Sodium Tolerance
Exclusion Compartmentalization in vacuoles Some plants make salt glands that excrete salt
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Mechanisms of Sodium Tolerance
Exclusion Compartmentalization in vacuoles Osmotic tolerance Mechs not fully understood; sometimes involve accumulation of organics such as trehalose in vacuole Reason weak HKT1 promotes Na tolerance?
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Mineral Nutrition Soil nutrients Amounts & availability vary PSU extension Text
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Mineral Nutrition Soil nutrients Amounts & availability vary Many are immobile, eg P, Fe
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Mineral Nutrition Soil nutrients Amounts & availability vary Many are immobile, eg P, Fe Mobile nutrients come with soil H2O
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Mineral Nutrition Soil nutrients Amounts & availability vary Many are immobile, eg P, Fe Mobile nutrients come with soil H2O Immobiles must be “mined” Root hairs get close
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Mineral Nutrition Immobile nutrients must be “mined” Root hairs get close Mycorrhizae get closer
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Mineral Nutrition Immobile nutrients must be mined Root hairs get close Mycorrhizae get closer Solubility varies w pH
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Mineral Nutrition Solubility varies w pH 5.5 is best compromise
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Mineral Nutrition Solubility varies w pH 5.5 is best compromise Plants alter roots to aid uptake
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Mineral Nutrition Nutrients in soil Plants alter roots to aid uptake Also use symbionts Mycorrhizal fungi help: especially with P
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Mineral Nutrition Also use symbionts Mycorrhizal fungi help: especially with P P travels poorly: fungal hyphae are longer & thinner
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Mineral Nutrition Also use symbionts Mycorrhizal fungi help: especially with P P travels poorly: fungal hyphae are longer & thinner Fungi give plants nutrients
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Mineral Nutrition Also use symbionts Mycorrhizal fungi help: especially with P P travels poorly: fungal hyphae are longer & thinner Fungi give plants nutrients Plants feed them sugar
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Mineral Nutrition Also use symbionts Mycorrhizal fungi help: especially with P P travels poorly: fungal hyphae are longer & thinner Fungi give plants nutrients Plants feed them sugar Ectomycorrhizae surround root: only trees, esp. conifers
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Mineral Nutrition Ectomycorrhizae surround root: only trees, esp. conifers release nutrients into apoplast to be taken up by roots
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Mineral Nutrition Ectomycorrhizae surround root: trees release nutrients into apoplast to be taken up by roots Endomycorrhizae invade root cells: Vesicular/Arbuscular Most angiosperms, especially in nutrient-poor soils
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Mineral Nutrition Endomycorrhizae invade root cells: Vesicular/Arbuscular Most angiosperms, especially in nutrient-poor soils May deliver nutrients into symplast
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Rhizosphere Endomycorrhizae invade root cells: Vesicular/Arbuscular Most angiosperms, especially in nutrient-poor soils May deliver nutrients into symplast Or may release them when arbuscule dies
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Rhizosphere Endomycorrhizae invade root cells: Vesicular/Arbuscular Most angiosperms, especially in nutrient-poor soils Deliver nutrients into symplast or release them when arbuscule dies Also find bacteria, actinomycetes, protozoa associated with root surface = rhizosphere
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Rhizosphere Also find bacteria, actinomycetes, protozoa associated with root surface = rhizosphere Plants feed them lots of C!
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Rhizosphere Also find bacteria, actinomycetes, protozoa associated with root surface = rhizosphere Plants feed them lots of C! They help make nutrients available
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Rhizosphere Also find bacteria, actinomycetes, protozoa associated with root surface = rhizosphere Plants feed them lots of C! They help make nutrients available N-fixing bacteria supply N to many plant spp
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Rhizosphere N-fixing bacteria supply N to many plant spp Most live in root nodules & are fed & protected from O2 by plant
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Nutrient uptake Most nutrients are dissolved in water
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Nutrient uptake Most nutrients are dissolved in water Enter root through apoplast until hit endodermis
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Nutrient uptake Most nutrients are dissolved in water Enter root through apoplast until hit endodermis Then must cross plasma membrane
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Selective Crossing membranes A) Diffusion through bilayer
B) Difusion through protein pore C) Facilitated diffusion D) Active transport E) Bulk transport 1) Exocytosis 2) Endocytosis Selective Active
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Nutrient uptake Then must cross plasma membrane Gases, small uncharged & non-polar molecules diffuse
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Nutrient uptake Then must cross plasma membrane Gases, small uncharged & non-polar molecules diffuse down their ∆ [ ] Important for CO2, auxin & NH3 transport
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Nutrient uptake Then must cross plasma membrane Gases, small uncharged & non-polar molecules diffuse down their ∆ [ ] Polar chems must go through proteins!
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Selective Transport 1) Channels integral membrane proteins with pore that specific ions diffuse through
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Selective Transport 1) Channels integral membrane proteins with pore that specific ions diffuse through depends on size & charge
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Channels integral membrane proteins with pore that specific ions diffuse through depends on size & charge O in selectivity filter bind ion (replace H2O)
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Channels integral membrane proteins with pore that specific ions diffuse through depends on size & charge O in selectivity filter bind ion (replace H2O) only right one fits
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Channels O in selectivity filter bind ion (replace H2O) only right one fits driving force? electrochemical D
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Channels driving force : electrochemical D “non-saturable”
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Channels driving force : electrochemical D “non-saturable” regulate by opening & closing
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Channels regulate by opening & closing ligand-gated channels open/close when bind specific chemicals
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Channels ligand-gated channels open/close when bind specific chemicals Stress-activated channels open/close in response to mechanical stimulation
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Channels Stress-activated channels open/close in response to mechanical stimulation voltage-gated channels open/close in response to changes in electrical potential
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Channels Old model: S4 slides up/down Paddle model: S4 rotates
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Channels Old model: S4 slides up/down Paddle model: S4 rotates 3 states Closed Open Inactivated
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Selective Transport 1) Channels 2) Facilitated Diffusion (carriers) Carrier binds molecule
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Selective Transport Facilitated Diffusion (carriers) Carrier binds molecule carries it through membrane & releases it inside
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Selective Transport Facilitated Diffusion (carriers) Carrier binds molecule carries it through membrane & releases it inside driving force = ∆ [ ]
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Selective Transport Facilitated Diffusion (carriers) Carrier binds molecule carries it through membrane & releases it inside driving force = ∆ [ ] Important for sugar transport
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Selective Transport Facilitated Diffusion (carriers) Characteristics 1) saturable 2) specific 3) passive: transports down ∆ []
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Selective Transport 1) Channels 2) Facilitated Diffusion (carriers) Passive transport should equalize [ ] Nothing in a plant cell is at equilibrium!
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Selective Transport Passive transport should equalize [ ] Nothing in a plant cell is at equilibrium! Solution: use energy to transport specific ions against their ∆ [ ]
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Active Transport Integral membrane proteins use energy to transport specific ions against their ∆ [ ] allow cells to concentrate some chemicals, exclude others
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Active Transport Characteristics 1) saturable ions/s molecules/s
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Active Transport Characteristics 1) saturable 2) specific
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Active Transport Characteristics 1) saturable 2) specific 3) active: transport up ∆ [ ] (or ∆ Em)
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4 classes of Active transport ATPase proteins
1) P-type ATPases (P = “phosphorylation”) Na/K pump Ca pump in ER & PM H+ pump in PM pumps H+ out of cell
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4 classes of Active transport ATPase proteins
1) P-type ATPases (P = “phosphorylation”) 2) V-type ATPases (V = “vacuole”) H+ pump in vacuoles
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4 classes of Active transport ATPase proteins
1) P-type ATPases (P = “phosphorylation”) 2) V-type ATPases (V=“vacuole”) 3) F-type ATPases (F = “factor”) a.k.a. ATP synthases mitochondrial ATP synthase chloroplast ATP synthase
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4 classes of Active transport ATPase proteins
1) P-type ATPases (P = “phosphorylation”) 2) V-type ATPases (V = “vacuole”) 3) F-type ATPases (F = “factor”) 4) ABC ATPases (ABC = “ATP Binding Cassette”) multidrug resistance proteins
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4 classes of Active transport ATPase proteins
1) P-type ATPases (P = “phosphorylation”) 2) V-type ATPases (V = “vacuole”) 3) F-type ATPases (F = “factor”) 4) ABC ATPases (ABC = “ATP Binding Cassette”) multidrug resistance proteins pump hydrophobic drugs out of cells very broad specificity
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Secondary active transport
Uses ∆ [ ] created by active transport to pump something else across a membrane against its ∆ [ ]
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Secondary active transport
Uses ∆ [ ] created by active transport to pump something else across a membrane against its ∆ [ ] Symport: both substances pumped same way
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Secondary active transport
Uses ∆ [ ] created by active transport to pump something else across a membrane against its ∆ [ ] Symport: both substances pumped same way Antiport: substances pumped opposite ways
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Secondary active transport
Uses ∆ [ ] created by active transport to pump something else across a membrane against its ∆ [ ] Symport: both substances pumped same way Antiport: substances pumped opposite ways
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Nutrient uptake Gases enter/exit by diffusion down their ∆ [ ]
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Nutrient uptake Gases enter/exit by diffusion down their ∆ [ ] Ions vary dramatically!
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Nutrient uptake Gases enter/exit by diffusion down their ∆ [ ] Ions vary dramatically! H+ is actively pumped out of cell by P-type H+ -ATPase
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Nutrient uptake Gases enter/exit by diffusion down their ∆ [ ] Ions vary dramatically! H+ is actively pumped out of cell by P-type H+ -ATPase and into vacuole by V-type!
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Nutrient uptake H+ is actively pumped out of cell by P-type H+ -ATPase and into vacuole by V-type! Main way plants make membrane potential (∆Em)!
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Nutrient uptake H+ is actively pumped out of cell by P-type H+ -ATPase and into vacuole by V-type! Main way plants make membrane potential (∆Em)! K+ diffuses through channels down ∆Em
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Nutrient uptake K+ diffuses through channels down ∆Em Also taken up by transporters
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Nutrient uptake K+ diffuses through channels down ∆Em Also taken up by transporters some also transport Na+
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Nutrient uptake K+ diffuses through channels down ∆Em Also taken up by transporters some also transport Na+ why Na+ slows K+ uptake?
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Nutrient uptake K+ diffuses through channels down ∆Em Also taken up by transporters some also transport Na+ why Na+ slows K+ uptake? Na+ is also expelled by H+ antiport
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Nutrient uptake K+ diffuses through channels down ∆Em Also taken up by transporters some also transport Na+ why Na+ slows K+ uptake? Na+ is also expelled by H+ antiport Enters through channels
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Nutrient uptake Na+ is also expelled by H+ antiport Enters through channels Ca2+ is expelled by P-type ATPases in PM
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Nutrient uptake Na+ is also expelled by H+ antiport Enters through channels Ca2+ is expelled by P-type ATPases in PM & pumped into vacuole
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Nutrient uptake Ca2+ is expelled by P-type ATPases in PM pumped into vacuole & ER by H+ antiport & P-type enters cytosol via gated channels
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Nutrient uptake PO43-, SO42-, Cl- & NO3- enter by H+ symport
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Nutrient uptake PO43-, SO42-, Cl- & NO3- enter by H+ symport also have anion transporters of ABC type
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Nutrient uptake PO43-, SO42-, Cl- & NO3- enter by H+ symport also have anion transporters of ABC type and anion channels
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Nutrient uptake PO43-, SO42-, Cl- & NO3- enter by H+ symport also have anion transporters of ABC type and anion channels Plants take up N many ways
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Nutrient uptake Plants take up N many ways: NO3- & NH4+ are main forms
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Nutrient uptake Plants take up N many other ways NO3- also by channels NH3 by diffusion NH4+ by carriers
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Nutrient uptake Plants take up N many other ways NO3- by channels NH3 by diffusion NH4+ by carriers NH4+ by channels
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Nutrient uptake Plants take up N many other ways 3 families of H+ symporters take up amino acids
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Nutrient uptake Plants take up N many other ways 3 families of H+ symporters take up amino acids Also have many peptide transporters some take up di- & tri- peptides by H+ symport
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Nutrient uptake Plants take up N many other ways 3 families of H+ symporters take up amino acids Also have many peptide transporters some take up di- & tri- peptides by H+ symport others take up tetra- & penta-peptides by H+ symport
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Nutrient uptake Plants take up N many other ways 3 families of H+ symporters take up amino acids Also have many peptide transporters some take up di- & tri- peptides by H+ symport others take up tetra- & penta-peptides by H+ symport Also have ABC transporters that import peptides
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Nutrient uptake Plants take up N many other ways 3 families of H+ symporters take up amino acids Also have many peptide transporters some take up di- & tri- peptides by H+ symport others take up tetra- & penta-peptides by H+ symport Also have ABC transporters that import peptides N is vital! NO3- & NH4+ are main forms
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Nutrient uptake Metals are taken up by ZIP proteins & by ABC transporters same protein may import Fe, Zn & Mn!
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Nutrient uptake Much is coupled to pH gradient
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Nutrient transport in roots
Move from soil to endodermis in apoplast
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Nutrient transport in roots
Move from soil to endodermis in apoplast Move from endodermis to xylem in symplast
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Nutrient transport in roots
Move from endodermis to xylem in symplast Transported into xylem by H+ antiporters
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Nutrient transport in roots
Move from endodermis to xylem in symplast Transported into xylem by H+ antiporters, channels
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Nutrient transport in roots
Transported into xylem by H+ antiporters, channels,pumps
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Nutrient transport in roots
Transported into xylem by H+ antiporters, channels,pumps Lowers xylem water potential -> root pressure
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Transport to shoot Nutrients move up plant in xylem sap
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Nutrient transport in leaves
Xylem sap moves through apoplast Leaf cells take up what they want
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