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Transport in plants occurs across a network of vessels and over long distances 1 Lecture 6 Outline (Ch. 36 & 37) I. Plant Transport Overview II. Driving Forces A. B. C. D. Water potential Transpiration & Bulk Flow in Xylem Stomata Control Positive Pressure & Bulk Flow in Phloem III. Mineral Acquisition IV. Essential Nutrients V. Relationships with other organisms VI. Preparation for next lecture 2 Transport in Plants Physical forces drive the transport of materials in plants over a range of distances Transport occurs on three scales 1. Within a cell – cellular level 2. Short-distance cell to cell – tissue level 3. Long-distance in xylem & phloem - whole plant level Transport occurs by 3 mechanisms: A. Osmosis & Diffusion B. Active Transport C. Bulk Flow 3 Transport in Plants – Water Potential Roots xylem stomata 4 To survive Water Potential – Plants must balance water uptake and loss • What is Osmosis? What is diffusion? • Water potential : predicts water movement due to solute concentration & pressure – designated as psi (ψ) Water molecules are attracted to: • Each other (cohesion) • Solid surfaces (adhesion) 5 Water Potential • Free water flows from regions of high water potential to regions of low water potential Ψ changes with: • Adding solutes • Adding pressure Water potential = Potential energy of water = Energy per volume of water in megapascals (MPa) ψTotal = ψsolute + ψpressure 6 Water Potential (a) • Solutes added decreases ψ 0.1 M solution (water less likely to cross membrane) Pure water (in an open area, no pressure, so ψp = 0) H2O = 0 MPa P = 0 S = 0.23 = 0.23 MPa 7 Water Potential • Application of physical pressure increases ψ (water more likely to cross membrane) (b) (c) H2O P = 0.23 S = 0.23 = 0 MPa = 0 MPa = 0 MPa H2O P = 0.30 S = 0.23 = 0.07 MPa 8 Water Potential Water Potential ψ = ψs + ψp Which direction will water move? ψcell = – 0.7 MPa + 0.5 MPa = – 0.2 MPa ψsolution = –0.3 MPa (solution has no pressure potential) 9 Water Potential • Water potential – Affects uptake and loss of water by plant cells • If a flaccid cell is placed in an environment with a higher solute concentration – The cell will lose water and become plasmolyzed 0.4 M sucrose solution: Plasmolyzed cell at osmotic equilibrium with its surroundings P = 0 S = 0.9 = 0.9 MPa P = 0 S = 0.9 = 0.9 MPa Initial flaccid cell: P = 0 S = 0.7 = 0.7 MPa 10 Water Potential Uses of turgor pressure: • • • Inexpensive cell growth Hydrostatic skeleton Phloem transport 11 Water Route Most plant tissues - cell walls and cytosol are continuous cell to cell (via?) - cytoplasmic continuum called the symplast apoplast = continuum of cell walls plus extracellular spaces 12 Water Route How do water and minerals get from the soil to vascular tissue? Symporters (cotransporters) contribute to the gradient that determines the directional flow of water. Here, pumps in H+ and mineral ions Water enters plants via the roots. H2O Soil Soil Cytosol Symporter H+ Mineral ions Water 13 14 Water Potential Minerals & ions pumped into root cells, then moved past endodermis What happens to ψ between soil and endodermis? Where is osmosis occurring? 14 Water Potential Once water & minerals cross the endodermis, they are transported through the xylem to upper parts of the plant. 16 Xylem Water exits plant through stomata. Smooth surface Rippled surface H2O Water moves up plant through xylem. Water film that coats mesophyll cell walls evaporates. Adhesion to xylem cells Cohesion between water molecules 16 16 17 Transpiration = loss of water from the shoot system to the surrounding environment. Bulk Flow = movement of fluid due to pressure gradient • Transpiration drives bulk flow of xylem sap. • Water is PULLED up a plant. • Ring/spiral wall thickening protects against vessel collapse 17 Xylem Ascent by Bulk Flow • The movement of xylem sap is against gravity – maintained by the transpiration-cohesion-tension • Stomata help regulate the rate of transpiration • Leaves generally have broad surface areas • These characteristics – Increase photosynthesis – Increase water loss through stomata 20 µm 18 19 Xylem What happens if rate of transpiration nears zero? i.e. – at night, water pressure builds up in the roots • Guttation 19 H+ pumped out Stomata Control K+ flow in H2O flow in Why? stomata open Why? K+ channels, aquaporins and radially oriented cellulose fibers play important roles. Cues for opening stomata: Light Depleted CO2 Internal cell “clocks” 20 21 Phloem • • • Direction is source to sink Near source to near sink Phloem under positive pressure Phloem sap composition: Phloem tissue • • • • • Sugar (mainly sucrose) amino acids hormones minerals enzymes Are tubers and bulbs sources or sinks? Aphid 21 Phloem Pressure Flow Hypothesis Vessel (xylem) Sieve tube Source cell (phloem) (leaf) H2O Sucrose H2O 1 Where are sugars made? 2 Pressure flow Water potential increased, turgor pressure increased, sap PUSHED through phloem Sugars removed (actively) at sink water potential decreased, water leaves phloem 2 3 Transpiration stream Sugars actively transported into companion cells plasmodesmata to sieve tube elements Via H+/sucrose Water follows (WHY?!) cotransporters 1 4 Sink cell (storage root) 3 H2O 4 Sucrose 22 Overview: A Nutritional Network • Every organism – Continually exchanges energy and materials with its environment • The branching root and shoot system provides high SA:V to collect resources – Plants’ resources are diffuse (scattered, at low concentration) What are these diffuse resources? 23 Mineral Acquisition What’s in dirt?! Mineral Acquisition • After heavy rainfall, water drains away from the larger spaces in soil – But smaller spaces retain water – attraction to surfaces, clay and other particles • The film of loosely bound water available to plants Soil particle surrounded by film of water Root hair Water available to plant Air space 25 Mineral Acquisition Soil particle Cation Exchange • Makes cations available for uptake. K+ – –– – Cu2+ K+ – – Mg2+ – + K – – Ca2+ H+ CO2 H+ Root hair H2O Steps: 1. Roots acidify soil solution via respired CO2 and H+/ATPase pumps 2. H+ attracted to soil particle (-) which “releases” cations 3. Roots absorb cations 26 27 Essential Nutrients and Deficiencies • Plants require certain chemicals to thrive • Plants derive most organic mass from the CO2 of air – Also depend on soil nutrients like water and minerals Essential elements: Required for a plant to complete its life cycle 27 Essential Nutrients and Deficiencies • Photosynthesis = major source of plant nutrition • Overall need – Macronutrients – used in larger amounts • Nine = C, O, H, N, K, Ca, Mg, P, and S – Micronutrients – used in minute amounts • Seven = Cl, Fe, Mn, Zn, B, Cu, and Mo Healthy Deficiency of any one can have severe effects on plant growth Phosphate-deficient Potassium-deficient Nitrogen-deficient 28 29 Relationship with other organisms • • • • Mycorrhizae Root nodulation Parasitic plants Carnivorous plants 29 Relationship with other organisms • Symbiotic associations with mycorrhizal fungi are found in about 90% of vascular plants – Substantially expand the surface area available for nutrient uptake – Enhance uptake of phosphorus and micronutrients The fungus gets: sugars from plant Agriculturally, farmers and foresters …Often inoculate seeds with spores of mycorrhizae to promote mycorrhizal relationships. 30 Nitrogen, Soil Bacteria and Nitrogen Availability • Plants need ammonia (NH3) or nitrate (NO3–) for: Proteins, nucleic acids, chlorophyll… • Nitrogen-fixing soil bacteria convert atmospheric N2 to nitrogenous minerals that plants can absorb N2 N2 Atmosphere Soil N2 Nitrogen-fixing bacteria Denitrifying bacteria H+ (From soil) Soil + NH4 NH3 (ammonia) – + NH4 (ammonium) Organic material (humus) Nitrate and nitrogenous organic compounds exported in xylem to shoot system Nitrifying bacteria NO3 (nitrate) Ammonifying bacteria Root Symbiotic relationships form between nitrogen-fixing bacteria and certain plants - Mainly legume family (e.g. peas, beans) 31 • Nodules: Swellings of plant cells “infected” by Rhizobium bacteria Nodules Bacteroids within vesicle 5 m Roots (a) Pea plant root (b) Bacteroids in a soybean root nodule. In this TEM, a cell from a root nodule of soybean is filled with bacteroids in vesicles. The cells on the left are uninfected. • Inside the nodule – Rhizobium bacteria assume a form called bacteroids, which are contained within vesicles formed by the root cell 32 Epiphytes, Parasitic, and Carnivorous Plants EPIPHYTES Anchored on another plant, self-nourished PARASITIC PLANTS Absorb sugar/minerals from host plant Staghorn fern, an epiphyte Pitcher plants cavity filled with digestive fluid Venus flytrap Mistletoe, a photosynthetic parasite To gain extra nitrogen Things To Do After Lecture 6… Reading and Preparation: 1. Re-read today’s lecture, highlight all vocabulary you do not understand, and look up terms. 2. Ch. 36 Self-Quiz: #2, 3, 4, 6, 7, 8, 9 (correct answers in back of book) Ch. 37 Self-Quiz: #1, 2, 8, 9, 10 (correct answers in back of book) 3. Read chapters 36 & 37, focus on material covered in lecture (terms, concepts, and figures!) 4. Skim next lecture. “HOMEWORK” (NOT COLLECTED – but things to think about for studying): 1. Explain the two components of water potential – which of these is due to osmosis? 2. Diagram the movement of water in a plant via xylem versus sugar movement through phloem. List similarities and differences. 3. Discuss how mycorrhizae and Rhizobium are different and the benefits each provide to plants. 4. Think about what types of environments might be more likely to have carnivorous plants – what do plants gain by digesting insects?