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Chapter 36 Transport in Plants
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Pathways for Survival For vascular plants The evolutionary journey onto land involved the differentiation of the plant body into roots and shoots
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Vascular tissue Transports nutrients throughout a plant; such transport may occur over long distances Figure 36.1
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Transport occurs on 3 scales
Transport of water and solutes by individual cells, such as root hairs Short-distance transport of substances from cell to cell Long-distance transport within xylem and phloem
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Types of transport CO2 O2 Light H2O Sugar O2 H2O CO2 Figure 36.2
Sugars are produced by photosynthesis in the leaves. 5 Through stomata, leaves take in CO2 and expel O2. The CO2 provides carbon for photosynthesis. Some O2 produced by photosynthesis is used in cellular respiration. 4 CO2 O2 Light H2O Sugar Transpiration, the loss of water from leaves (mostly through stomata), creates a force within leaves that pulls xylem sap upward. 3 Sugars are transported as phloem sap to roots and other parts of the plant. 6 Water and minerals are transported upward from roots to shoots as xylem sap. 2 Roots exchange gases with the air spaces of soil, taking in O2 and discharging CO2. In cellular respiration, O2 supports the breakdown of sugars. 7 Roots absorb water and dissolved minerals from the soil. 1 O2 H2O CO2 Minerals Figure 36.2
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Central Role of Proton Pumps
Create a H+ (proton) gradient PE that can be harnessed to do work membrane potential CYTOPLASM EXTRACELLULAR FLUID ATP H+ Proton pump generates membrane potential and H+ gradient. – + Figure 36.3
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(b) Cotransport of anions
Coupled transport Figure 36.4b H+ NO3– NO3 – – + (b) Cotransport of anions of through a cotransporter. Cell accumulates anions ( , for example) by coupling their transport to the inward diffusion
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(c) Contransport of a neutral solute
“coattail” effect of cotransport Responsible for the uptake of sucrose by plant cells H+ S Plant cells can also accumulate a neutral solute, such as sucrose ( ), by cotransporting down the steep proton gradient. – + Figure 36.4c (c) Contransport of a neutral solute
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Effects of Differences in Water Potential
Plants must balance water uptake and loss Osmosis Determines uptake or loss Affected by solute conc. & pressure
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Water potential (y) Water Solute concentration and pressure
Direction of movement of water Water Flows f/ high y to low y
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Solute potential Pressure potential
Proportional to # of dissolved molecules Pressure potential
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Addition of solutes Reduces y Figure 36.5a (a) P = 0 S = 0.23
0.1 M solution H2O Pure waer P = 0 S = 0.23 = 0.23 MPa = 0 MPa (a) Figure 36.5a
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Application of pressure
Increases y H2O P = S = 0.23 = 0 MPa = 0 MPa (b) H2O P = S = 0.23 = MPa = 0 MPa (c) Figure 36.5b, c
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Negative pressure Decreases y Figure 36.5d (d) H2O P = 0.30 P = 0
= 0.23 MPa (d) P = 0.30 S = 0 = 0.30 MPa Figure 36.5d
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y Flaccid cells in higher solute concentration
Affects uptake and loss of water by plant cells Flaccid cells in higher solute concentration Cells lose water plasmolyzed Figure 36.6a 0.4 M sucrose solution: Initial flaccid cell: Plasmolyzed cell at osmotic equilibrium with its surroundings P = 0 S = 0.7 S = 0.9 = 0.9 MPa = 0.7 MPa
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If Flaccid cell solution w/ low conc
Cell gains water turgid Distilled water: Initial flaccid cell: Turgid cell at osmotic equilibrium with its surroundings P = 0 S = 0.7 S = 0 P = 0.7 Figure 36.6b = 0.7 MPa = 0 MPa = 0 MPa
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Turgor loss (wilting) Reversed when the plant is watered Figure 36.7
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Aquaporin Proteins Transport proteins in the cell membrane that allow the passage of water Do not affect y
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Plasma membrane Controls traffic of molecules into and out of the protoplast
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Vacuolar membrane Regulates transport between the cytosol and the vacuole Transport proteins in the plasma membrane regulate traffic of molecules between the cytosol and the cell wall. the vacuolar membrane regulate traffic of molecules between the cytosol and the vacuole. Plasmodesma Vacuolar membrane (tonoplast) Plasma membrane Cell wall Cytosol Vacuole Cell compartments. The cell wall, cytosol, and vacuole are the three main compartments of most mature plant cells. (a) Figure 36.8a
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Cell walls and cytosol continuous from cell to cell
Cytoplasmic continuum symplast Extracellular continuum apoplast
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Symplastic route Apoplastic route
Key Symplast Apoplast The symplast is the continuum of cytosol connected by plasmodesmata. The apoplast is the continuum of cell walls and extracellular spaces. Transmembrane route Symplastic route Apoplastic route Transport routes between cells. At the tissue level, there are three passages: the transmembrane, symplastic, and apoplastic routes. Substances may transfer from one route to another. (b) Figure 36.8b
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Bulk Flow in Long-Distance Transport
Movement of fluid in the xylem and phloem is driven by pressure differences
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Water and mineral salts f/ soil epidermis of roots shoot system
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Lateral transport Figure 36.9 Casparian strip Endodermal cell
1 2 3 Uptake of soil solution by the hydrophilic walls of root hairs provides access to the apoplast. Water and minerals can then soak into the cortex along this matrix of walls. Minerals and water that cross the plasma membranes of root hairs enter the symplast. As soil solution moves along the apoplast, some water and minerals are transported into the protoplasts of cells of the epidermis and cortex and then move inward via the symplast. Within the transverse and radial walls of each endodermal cell is the Casparian strip, a belt of waxy material (purple band) that blocks the passage of water and dissolved minerals. Only minerals already in the symplast or entering that pathway by crossing the plasma membrane of an endodermal cell can detour around the Casparian strip and pass into the vascular cylinder. Endodermal cells and also parenchyma cells within the vascular cylinder discharge water and minerals into their walls (apoplast). The xylem vessels transport the water and minerals upward into the shoot system. Casparian strip Pathway along apoplast Pathway through symplast Plasma membrane Apoplastic route Symplastic Root hair Epidermis Cortex Endodermis Vascular cylinder Vessels (xylem) Endodermal cell 4 5
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Root hairs account for much of the surface area of roots
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Mycorrhizae Mutually beneficial relationships (symbiosis) with fungal hyphae, facilitate the absorption of water and minerals 2.5 mm Figure 36.10
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The Endodermis: A Selective Sentry
Innermost layer of cells in the root cortex Surrounds the vascular cylinder and functions as the last checkpoint for the selective passage of minerals from the cortex into the vascular tissue
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Waxy Casparian strip of the endodermal wall blocks apoplastic transfer
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Water and minerals ascend from roots to shoots through the xylem
Plants lose an enormous amount of water through transpiration, loss of water f/ leaves
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Xylem sap Rises to heights of more than 100 m in the tallest plants
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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
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Root pressure results in guttation, (exudation) of water f/ tips of leaf
Figure 36.11
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Pulling Xylem Sap: The Transpiration-Cohesion-Tension Mechanism
Water is pulled upward by negative pressure in the xylem Based on cohesion of water molecules which is based on hydrogen bonding
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Water vapor in the airspaces of a leaf
Transpirational Pull Water vapor in the airspaces of a leaf Diffuses down its water potential gradient and exits the leaf via stomata
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Transpiration produces negative pressure (tension) in the leaf
Pulls on water in the xylem, pulling water into the leaf Evaporation causes the air-water interface to retreat farther into the cell wall and become more curved as the rate of transpiration increases. As the interface becomes more curved, the water film’s pressure becomes more negative. This negative pressure, or tension, pulls water from the xylem, where the pressure is greater. Cuticle Upper epidermis Mesophyll Lower Water vapor CO2 O2 Xylem Stoma Evaporation At first, the water vapor lost by transpiration is replaced by evaporation from the water film that coats mesophyll cells. In transpiration, water vapor (shown as blue dots) diffuses from the moist air spaces of the leaf to the drier air outside via stomata. Airspace Cytoplasm Cell wall Vacuole Water film Low rate of transpiration High rate of Air-water interface Y = –0.15 MPa Y = –10.00 MPa 3 1 2 Figure 36.12 Air- space
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Cohesion and Adhesion in the Ascent of Xylem Sap
Transpirational pull on xylem sap Transmitted f/ leaves to roots and even into the soil solution Facilitated by cohesion and adhesion
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Water potential gradient
Ascent of xylem sap Xylem sap Outside air Y = –100.0 MPa Leaf Y (air spaces) = –7.0 MPa Leaf Y (cell walls) = –1.0 MPa Trunk xylem Y = – 0.8 MPa Water potential gradient Root xylem Y = – 0.6 MPa Soil Y = – 0.3 MPa Mesophyll cells Stoma Water molecule Atmosphere Transpiration Adhesion Cell wall Cohesion, by hydrogen bonding Root hair Soil particle Cohesion and adhesion in the xylem Water uptake from soil Figure 36.13
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Stomata help regulate the rate of transpiration
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Stomata regulate transpiration rate, also:
Photosynthetic rate Water loss rate 20 µm Figure 36.14
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Stomata About 90% of the water a plant loses through stomata
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Stomata flanked by guard cells
Control the diameter of the stoma by changing shape Cells flaccid/Stoma closed Cells turgid/Stoma open Radially oriented cellulose microfibrils Cell wall Vacuole Guard cell Changes in guard cell shape and stomatal opening and closing (surface view). Guard cells of a typical angiosperm are illustrated in their turgid (stoma open) and flaccid (stoma closed) states. The pair of guard cells buckle outward when turgid. Cellulose microfibrils in the walls resist stretching and compression in the direction parallel to the microfibrils. Thus, the radial orientation of the microfibrils causes the cells to increase in length more than width when turgor increases. The two guard cells are attached at their tips, so the increase in length causes buckling. (a) Figure 36.15a
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Chgs. in turgor pressure open and close stomata
From uptake and loss of K+ by the guard cells H2O K+ Role of potassium in stomatal opening and closing. The transport of K+ (potassium ions, symbolized here as red dots) across the plasma membrane and vacuolar membrane causes the turgor changes of guard cells. (b) Figure 36.15b
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Xerophytes Plants adapted to arid climates Leaf modifications
reduce transpiration
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Upper epidermal tissue
Stomata of xerophytes concentrated on the lower leaf surface, often located in depressions Lower epidermal tissue Trichomes (“hairs”) Cuticle Upper epidermal tissue Stomata 100 m Figure 36.16
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Translocation Transport of organic nutrients (e.g. sugar) in the plant
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Sugar Movement Phloem sap Aqueous sucrose soln.
Travels from a sugar source to a sugar sink
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Sugar source Sugar sink Sugar producer, e.g. leaves
Consumer or storer of sugar, e.g.tuber or bulb
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Sugar loaded into sieve-tube members
Figure 36.17a Mesophyll cell Cell walls (apoplast) Plasma membrane Plasmodesmata Companion (transfer) cell Sieve-tube member Phloem parenchyma cell Bundle- sheath cell Sucrose manufactured in mesophyll cells can travel via the symplast (blue arrows) to sieve-tube members. In some species, sucrose exits the symplast (red arrow) near sieve tubes and is actively accumulated from the apoplast by sieve-tube members and their companion cells. (a)
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Phloem loading requires active transport
Proton pumping and cotransport of sucrose and H+ A chemiosmotic mechanism is responsible for the active transport of sucrose into companion cells and sieve-tube members. Proton pumps generate an H+ gradient, which drives sucrose accumulation with the help of a cotransport protein that couples sucrose transport to the diffusion of H+ back into the cell. (b) High H+ concentration Cotransporter Proton pump ATP Key Sucrose Apoplast Symplast H+ Low H+ concentration S Figure 36.17b
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Pressure Flow Sap moves through a sieve tube by bulk flow driven by positive pressure Vessel (xylem) H2O Sieve tube phloem) Source cell (leaf) Sucrose Sink cell (storage root) 1 Loading of sugar (green dots) into the sieve tube at the source reduces water potential inside the sieve-tube members. This causes the tube to take up water by osmosis. 2 4 3 This uptake of water generates a positive pressure that forces the sap to flow along the tube. The pressure is relieved by the unloading of sugar and the consequent loss of water from the tube at the sink. In the case of leaf-to-root translocation, xylem recycles water from sink to source. Transpiration stream Pressure flow Figure 36.18
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Pressure flow explains why phloem sap always flows from source to sink
Aphid feeding Stylet in sieve-tube member Severed stylet exuding sap Sieve- Tube EXPERIMENT RESULTS CONCLUSION Sap droplet Stylet Sap droplet 25 m Sieve- tube member To test the pressure flow hypothesis,researchers used aphids that feed on phloem sap. An aphid probes with a hypodermic- like mouthpart called a stylet that penetrates a sieve-tube member. As sieve-tube pressure force-feeds aphids, they can be severed from their stylets, which serve as taps exuding sap for hours. Researchers measured the flow and sugar concentration of sap from stylets at different points between a source and sink. The closer the stylet was to a sugar source, the faster the sap flowed and the higher was its sugar concentration. The results of such experiments support the pressure flow hypothesis. Figure 36.19
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