Copyright 2005 Pearson Education Inc publishing as Benjamin
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Concept 36. 1: Physical forces drive the transport of materials in plants over a range of distances • Transport in vascular plants occurs on three scales: – Transport of water and solutes by individual cells, such as root hairs – Short-distance transport of substances from cell to cell at the levels of tissues and organs – Long-distance transport within xylem and phloem at the level of the whole plant Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -2_4 CO 2 Light H 2 O Sugar O 2 H 2 O Minerals CO 2
Selective Permeability of Membranes: A Review • The selective permeability of the plasma membrane controls movement of solutes into and out of the cell • Specific transport proteins enable plant cells to maintain an internal environment different from their surroundings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
The Central Role of Proton Pumps • Proton pumps in plant cells create a hydrogen ion gradient that is a form of potential energy that can be harnessed to do work • They contribute to a voltage known as a membrane potential Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -3 CYTOPLASM EXTRACELLULAR FLUID ATP Proton pump generates membrane potential and gradient.
• Plant cells use energy stored in the proton gradient and membrane potential to drive the transport of many different solutes Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -4 a CYTOPLASM EXTRACELLULAR FLUID Cations ( , for example) are driven into the cell by the membrane potential. Transport protein Membrane potential and cation uptake
• In the mechanism called cotransport, a transport protein couples the passage of one solute to the passage of another Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -4 b Cell accumulates anions ( , for example) by coupling their transport to; the inward diffusion of through a cotransporter. Cotransport of anions
• The “coattail” effect of cotransport is also responsible for the uptake of the sugar sucrose by plant cells Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -4 c Plant cells can also accumulate a neutral solute, such as sucrose ( ), by cotransporting down the steep proton gradient. Cotransport of a neutral solute
Effects of Differences in Water Potential • To survive, plants must balance water uptake and loss • Osmosis determines the net uptake or water loss by a cell is affected by solute concentration and pressure Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -8 a Cell wall Cytosol Vacuole Plasmodesma Key Symplast Apoplast Vacuolar membrane (tonoplast) Plasma membrane Cell compartments
• In most plant tissues, the cell walls and cytosol are continuous from cell to cell • The cytoplasmic continuum is called the symplast • The apoplast is the continuum of cell walls and extracellular spaces Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -8 b Key Symplast Apoplast Transmembrane route Apoplast Symplastic route Apoplastic route Transport routes between cells
Functions of the Symplast and Apoplast in Transport • Water and minerals can travel through a plant by three routes: – Transmembrane route: out of one cell, across a cell wall, and into another cell – Symplastic route: via the continuum of cytosol – Apoplastic route: via the cell walls and extracellular spaces Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Bulk Flow in Long-Distance Transport • In bulk flow, movement of fluid in the xylem and phloem is driven by pressure differences at opposite ends of the xylem vessels and sieve tubes Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Concept 36. 2: Roots absorb water and minerals from the soil • Water and mineral salts from the soil enter the plant through the epidermis of roots and ultimately flow to the shoot system Animation: Transport in Roots Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -9 Casparian strip Pathway along apoplast Endodermal cell Pathway through symplast Casparian strip Plasma membrane Apoplastic route Symplastic route Vessels (xylem) Root hair Epidermis Endodermis Vascular cylinder Cortex
The Roles of Root Hairs, Mycorrhizae, and Cortical Cells • Much of the absorption of water and minerals occurs near root tips, where the epidermis is permeable to water and root hairs are located • Root hairs account for much of the surface area of roots Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Most plants form mutually beneficial relationships with fungi, which facilitate absorption of water and minerals from the soil • Roots and fungi form mycorrhizae, symbiotic structures consisting of plant roots united with fungal hyphae Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -10 2. 5 mm
• Water can cross the cortex via the symplast or apoplast • The waxy Casparian strip of the endodermal wall blocks apoplastic transfer of minerals from the cortex to the vascular cylinder Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Concept 36. 3: Water and minerals ascend from roots to shoots through the xylem • Plants lose an enormous amount of water through transpiration, the loss of water vapor from leaves and other aerial parts of the plant • The transpired water must be replaced by water transported up from the roots Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Factors Affecting the Ascent of Xylem Sap • Xylem sap rises to heights of more than 100 m in the tallest plants • Is the sap pushed upward from the roots, or is it pulled upward by the leaves? Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Pushing Xylem Sap: Root Pressure • At night, when transpiration is very low, root cells continue pumping mineral ions into the xylem of the vascular cylinder, lowering the water potential • Water flows in from the root cortex, generating root pressure Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Root pressure sometimes results in guttation, the exudation of water droplets on tips of grass blades or the leaf margins of some small, herbaceous eudicots Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Pulling Xylem Sap: The Transpiration-Cohesion Tension Mechanism • Water is pulled upward by negative pressure in the xylem Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Transpirational Pull • Water vapor in the airspaces of a leaf diffuses down its water potential gradient and exits the leaf via stomata • Transpiration produces negative pressure (tension) in the leaf, which exerts a pulling force on water in the xylem, pulling water into the leaf Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -12 Y = – 0. 15 MPa Y = – 10. 00 MPa Cell wall Air-water interface Airspace Low rate of transpiration Cuticle Upper epidermis High rate of transpiration Cytoplasm Evaporation Mesophyll Airspace Air space Cell wall Evaporation Water film Lower epidermis Cuticle CO 2 Xylem CO 2 Stoma Vacuole
Cohesion and Adhesion in the Ascent of Xylem Sap • The transpirational pull on xylem sap is transmitted all the way from the leaves to the root tips and even into the soil solution • Transpirational pull is facilitated by cohesion and adhesion Animation: Transpiration Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -13 Xylem sap Outside air = – 100. 0 MPa Mesophyll cells Stoma Leaf (air spaces) = – 7. 0 MPa Water molecule Transpiration Atmosphere Trunk xylem = – 0. 8 Mpa Water potential gradient Leaf (cell walls) = – 1. 0 MPa Xylem cells Adhesion Cell wall Cohesion, Cohesion and by adhesion in hydrogen the xylem bonding Water molecule Root hair Root xylem = – 0. 6 MPa Soil = – 0. 3 MPa Soil particle Water uptake from soil Water
Xylem Sap Ascent by Bulk Flow: A Review • The movement of xylem sap against gravity is maintained by the transpiration-cohesion-tension mechanism Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Concept 36. 4: Stomata help regulate the rate of transpiration • Leaves generally have broad surface areas and high surface-to-volume ratios • These characteristics increase photosynthesis and increase water loss through stomata Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -14 20 µm
Effects of Transpiration on Wilting and Leaf Temperature • Plants lose a large amount of water by transpiration • If the lost water is not replaced by absorption through the roots, the plant will lose water and wilt Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Transpiration also results in evaporative cooling, which can lower the temperature of a leaf and prevent denaturation of various enzymes involved in photosynthesis and other metabolic processes Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Stomata: Major Pathways for Water Loss • About 90% of the water a plant loses escapes through stomata • Each stoma is flanked by a pair of guard cells, which control the diameter of the stoma by changing shape Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -15 a Cells turgid/Stoma open Cells flaccid/Stoma closed Radially oriented cellulose microfibrils Cell wall Vacuole Guard cell Changes in guard cell shape and stomatal opening and closing (surface view)
• Changes in turgor pressure that open and close stomata result primarily from the reversible uptake and loss of potassium ions by the guard cells Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -15 b Cells turgid/Stoma open H 2 O Cells flaccid/Stoma closed H 2 O K+ H 2 O H 2 O Role of potassium in stomatal opening and closing H 2 O
Xerophyte Adaptations That Reduce Transpiration • Xerophytes are plants adapted to arid climates • They have leaf modifications that reduce the rate of transpiration • Their stomata are concentrated on the lower leaf surface, often in depressions that provide shelter from dry wind Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -16 Cuticle Upper epidermal tissue Lower epidermal Trichomes Stomata tissue (“hairs”) 100 µm
Concept 36. 5: Organic nutrients are translocated through the phloem • Translocation is the transport of organic nutrients in a plant Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Movement from Sugar Sources to Sugar Sinks • Phloem sap is an aqueous solution that is mostly sucrose • It travels from a sugar source to a sugar sink • A sugar source is an organ that is a net producer of sugar, such as mature leaves • A sugar sink is an organ that is a net consumer or storer of sugar, such as a tuber or bulb Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -17 Key Apoplast Symplast Companion (transfer) cell Mesophyll cell Cell walls (apoplast) Sieve-tube member High H+ concentration Cotransporter Proton pump Plasma membrane Plasmodesmata Sucrose Mesophyll cell Bundlesheath cell Phloem parenchyma cell Low H+ concentration
• In many plants, phloem loading requires active transport • Proton pumping and cotransport of sucrose and H+ enable the cells to accumulate sucrose Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
LE 36 -18 Sieve tube (phloem) Vessel (xylem) H 2 O Source cell (leaf) Sucrose Pres sure flow Transpiration stream H 2 O Sink cell (storage root) Sucrose H 2 O
- Slides: 50