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Lecture Notebook to accompany Sinauer Associates, Inc. MacMillan Copyright © 2014 Sinauer Associates, Inc. Cover photograph © Alex Mustard/naturepl.com. This document may not be modified or distributed (either electronically or on paper) without the permission of the publisher, with the following exception: Individual users may enter their own notes into this document and may print it for their own personal use. 00 35 Transport in Plants O2 (product of photosynthesis) CO2 (reactant of photosynthesis) H2O (transpiration) H2O, carbohydrates, etc. CO2 enters and O2 and H2O exit the leaves via the stomata (see Figure 35.8). H2O and dissolved minerals 35.1 The Pathways of Water and Solutes in a Plant (Page 727) LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 35.01 Date 07-13-12 To add your own notes to any page, use Adobe Reader’s Typewriter feature, accessible via the Typewriter bar at the top of the window. (Requires Adobe Reader 8 or later. Adobe Reader can be downloaded free of charge from the Adobe website: http://get.adobe.com/reader.) 2 © 2014 Sinauer Associates, Inc. Chapter 35 | Transport in Plants 3 (A) In this tube, the solute potentials on the two sides of the membrane differ, but the pressure potentials are the same. Pure water y = 0 MPa The right side of the tube has a lower water potential, so there is a net movement of water to the right. Membrane In this tube, a piston is used to increase the pressure potential of the right side. Solution yp = 0 MPa ys = –1.0 MPa Pure water y = 0 MPa y = –1.0 MPa The water potentials of the two sides are equal, so there is no net movement of water. Solution yp = +1.0 MPa ys = –1.0 MPa y = 0 MPa (B) The inside of the cell has a lower solute potential than the surrounding water. The cell has a pressure potential of zero. Pure water y = 0 MPa The cell has a lower water potential than the water outside, so there is net movement of water into the cell. The cell has a negative solute potential, but has a positive pressure potential. Flaccid cell yp = 0 MPa ys = –1.0 MPa The pressure potential of the cell balances its solute potential, so the cell’s water potential is zero. There is no net movement of water. Pure water y = 0 MPa y = –1.0 MPa Turgid cell yp = +1.0 MPa ys = –1.0 MPa y = 0 MPa 35.2 Water Potential, Solute Potential, and Pressure Potential (Page 728) The cells of this plant have low turgor pressure and the plant is wilted. LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 35.02 Date 07-13-12 The water potential of cells of this plant is zero because the negative solute potential is balanced by an equally positive pressure potential. The plant is upright because its cells are turgid. 35.3 A Wilted Plant (Page 728) © 2014 Sinauer Associates, Inc. Chapter 35 | Transport in Plants 1 A proton pump generates differences in H+ concentration and electric charge across the membrane. Outside of cell Plasma membrane H+ 4 2 The difference in electric charge causes cations such as K+ to enter the cell. H+ H+ H+ H+ K+ Proton pump ATP Inside of cell H+ ADP + fusion of H+ to the transport (against an electrochemical gradient) of anions such as Cl– into the cell. H+ H+ H+ 3 Symport couples the dif- H+ Cl– H+ Symport protein Potassium channel K+ K+ Pi K+ + K + K H+ Cl– Cl– Cl– 35.4 The Proton Pump in Transport of K+ and Cl– (Page 729) LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 35.04 Date 07-13-12 Cell membrane Cytoplasm Plasmodesmata Cell wall Water and solutes can move in the symplast by crossing a cell membrane and passing through plasmodesmata. Water and solutes can move through the apoplast without passing through cell membrane. 35.5 Apoplast and Symplast (Page 729) © 2014 Sinauer Associates, Inc. Chapter 35 | Transport in Plants 5 The Casparian strip prevents water and solutes in the apoplast from passing between the endodermal cells into the stele. Epidermis Root hair Endodermis Cortex Cortex Endodermis Pericycle Pericycle Xylem Stele Casparian strip Soil solution Apoplast Symplast Plasmodesmata Water and solutes travel through the symplast or apoplast until they reach the endodermis. At the Casparian strip, water and solutes in the apoplast must enter the symplast to cross the endodermis. Inside the stele, solutes are actively transported into the apoplast and water follows passively, forming the xylem sap. 35.6 Pathways to the Root Xylem (Page 730) LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 35.06 Date 09-10-12 © 2014 Sinauer Associates, Inc. Chapter 35 | Transport in Plants 6 3 Tension pulls water from the veins into the apoplast of the mesophyll cells... Leaf Vein 4 ...then pulls the water column through the veins in the leaves... 2 Water evaporates from mesophyll cell walls. Mesophyll cell Stem H2O 5 ...and then upward in the xylem of the root and stem. 1 During transpiration water vapor diffuses out of the stomata. Xylem H H H O O H H O H H H O Root 6 Water molecules form a cohesive water column from the roots to the leaves. H2O 7 Water moves H2O into the xylem by osmosis. 8 Water enters the root from Xylem the soil by osmosis. 35.7 The Transpiration–Cohesion–Tension Mechanism (Page 731) LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 35.06 Date 07-13-12 © 2014 Sinauer Associates, Inc. Chapter 35 | Transport in Plants 7 RESEARCHTOOLS 35.8 Measuring the Pressure of Xylem Sap with a Pressure Chamber Without pressure 1 By applying just enough pressure… With pressure 2 …so that xylem sap is pushed back to the cut surface of a plant sample,… 3 …a scientist can determine the tension on the sap in the living plant. Gas pressure Pressure gauge Pressure release valve (Page 732) Life10e_35.07 Sinauer 7-11-12 © 2014 Sinauer Associates, Inc. Chapter 35 | Transport in Plants 8 (A) Guard cells 10 µm Stoma (B) Cl– K+ 1 In the light, guard cells actively pump protons out, thus facilitating the entry of K+ and Cl–. H+ 2 Higher internal K+ and Guard cells Cl– concentrations give guard cells a more negative water potential, causing them to take up water, increase in pressure, and stretch, opening the stoma. H2O Stoma Cellulose microfibrils K+ H2O Cl– 35.9 Stomata (Page 733) 3 In the absence of light, K+ and Cl– diffuse passively out of the guard cells, and water follows by osmosis. The guard cells shrink and the stoma closes. Remove a ring of bark to girdle the tree. Organic solutes accumulate in the phloem above the girdle, causing swelling. Time LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 35.08 Date 07-13-12 Bark Wood In-Text Art (Page 734) © 2014 Sinauer Associates, Inc. Chapter 35 | Transport in Plants 9 Pores of sieve plate Sieve plate Mature sieve tube elements do not have nuclei and have lost most of their organelles. Sieve tube element Phloem sap The companion cell is a fully functional cell with a nucleus. Companion cell Pores Sieve plate Dr. R. Kessel & Dr. G. Shih/Visuals Unlimited. 35.10 Sieve Tubes (Page 735) Sieve tube element LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 35.09 Date 07-13-12 The aphid’s stylet has successfully penetrated the sieve tube. Sap droplet Longistigma caryae (aphid) Stylet In-Text Art (Page 735) © 2014 Sinauer Associates, Inc. Chapter 35 | Transport in Plants 10 Phloem sieve tube Xylem 1 Transpiration pulls water up xylem vessels. Source cell 2 Source cells load sucrose into phloem sieve tubes, reducing their water potential… H2O H2O Sucrose 3 …so water is taken up from xylem vessels by osmosis, raising the pressure potential in the sieve tubes. H2O Sink cell 4 Internal pressure differences drive the sap along the sieve tube to sink cells. Sucrose H2O 5 Sucrose is unloaded into sink cells, increasing the water potential in the sieve tube… 6 …and water moves back to xylem vessels. 35.11 The Pressure Flow Model (Page 736) LIFE The Science of Biology 10E Sadava Sinauer Associates Morales Studio Figure 35.10 Date 07-13-12 TablE35.1 Mechanisms of Sap Flow in Plant Vascular Tissues Xylem Phloem Driving force for bulk flow Transpiration from leaves Active transport of sucrose at source and sink Site of bulk flow Nonliving vessel Living sieve tube elements and tracheids elements Pressure Negative (pull from top; Positive (push from potential in sap tension) source; pressure) (Page 736) © 2014 Sinauer Associates, Inc. Chapter 35 | Transport in Plants 11 INVESTIGATINGLIFE 35.12 Manipulating Sucrose Transport from the Phloem HYPOTHESIS Reducing the sucrose concentration in a sink organ will increase the transport and unloading of sucrose from the phloem. Method Plants were transformed with a gene for invertase, an enzyme that hydrolyzes sucrose. The potato plant has underground tubers, modified stems that store starch. Results Wild-type plants Transgenic plants Phloem Tuber sink cell Sucrose The wild-type plants had a high level of sucrose in developing tubers. The tubers were normal in size and number. The genetically modified plants had a low level of sucrose in developing tubers and produced fewer but much larger potatoes. CONCLUSION Increasing sink strength increases sucrose transport into developing tissues. Go to BioPortal for discussion and relevant links for all INVESTIGATINGLIFE figures. (Page 737) 11 © 2014 Sinauer Associates, Inc.