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Plant Transport Plants Plant: terrestrial (mostly), multicellular, photoautotrophic, eukaryote, true tissues and organs Plant Structure Tissue Basic Tissue Types: pg 717 - give rise to specialized cells o Dermal - outer coat o Vascular – transport tubes o Ground – between Dermal and Vascular Basic Tissue Layout Dermal Ground Vascular Dermal tissue Ground tissue Vascular tissue Dermal Tissue Epidermis Function: protection: secretes the cuticle, forms prickles and root hairs Thorns, Spines and Prickles Based on where they originate Thorns – modified stems Spines – modified leaves Prickles – modified epidermal cells Thorn Spine Prickle Rose “thorns” are prickles “A rose between two prickles.” Vascular Tissue Xylem: Water conducting – unidirectional (up) Dead at maturity – pg 719 Phloem: Sugar conduction – bidirectional Sieve Tube Members: alive and functional – lack many organelles Companion Cells: connected to Sieve Tube Members by plasmodesmata – supports the STM with its organelle function Xylem Phloem WATER-CONDUCTING CELLS OF THE XYLEM Vessel Tracheids SUGAR-CONDUCTING CELLS OF THE PHLOEM Sieve-tube members: longitudinal view 100 m Pits Companion cell Sieve-tube member Sieve plate Tracheids and vessels Vessel element Vessel elements with partially perforated end walls Nucleus 30 m 15 m Tracheids Cytoplasm Figure. 35.9 Companion cell Ground Tissue Occupies the space between the vascular tissue and the dermal tissue Functions: Storage – roots and stems Support – stems Photosynthesis – leaves and some stems Types of Ground Tissue 1.Parenchyma: undifferentiated, thin cell walls (still flexible) – used for metabolism and photosynthesis Ex: Pallisade and Spongy Mesophyll of leaf Potato, Fruit pulp 2. Collenchyma: unevenly thickened cell walls – support young parts of plants – no lignin, but stronger than parenchyma Ex: “Strings” in celery 3. Sclerenchyma: highly thickened cell walls – lignified – support mature tissue – hard and dead Two types: Fibers and Sclerids Ex: Walnut Shell, Stone Cells in Pears PARENCHYMA CELLS COLLENCHYMA CELLS 80 m Cortical parenchyma cells SCLERENCHYMA CELLS 5 m Sclereid cells in pear 25 m Cell wall Parenchyma cells 60 m Collenchyma cells Fiber cells Plant Parts: Roots, Stems and Leaves Roots: Functions: - absorb water, nutrients and minerals - anchor plant in soil - store food and water - support the plant (a) Prop roots (d) Buttress roots (b) Storage roots (c) “Strangling” aerial roots (e) Pneumatophores Increasing Absorption - Root hairs – extensions of the epidermis Branching roots – lateral roots Mycorrhizae Root Structure Outside In Epidermis (D) Cortex (G) – storage and nutrient transfer Endodermis (G) – separates ground and vascular tissue – important for water transfer Pericycle (V) – forms the lateral roots Stele (Xylem and Phloem) (V) Epidermis Cortex Vascular cylinder Endodermis Pericycle Core of parenchyma cells Xylem 100 m Phloem 100 m (a) Transverse section of a typical root. In the roots of typical gymnosperms and eudicots, as well as some monocots, the stele is a vascular cylinder consisting of a lobed core of xylem with phloem between the lobes. (b) Endodermis Key Dermal Pericycle Ground Vascular Xylem Phloem Transverse section of a root with parenchyma in the center. The stele of many monocot roots is a vascular cylinder with a core of parenchyma surrounded by a ring of alternating xylem and phloem. Eudicot Root – Cross Section From: http://www.inclinehs.org/smb/Sungirls/images/dicot_stem.JPG Monocot Root Cross Section From: http://www.inclinehs.org/smb/Sungirls/images/monocot_stem.JPG Monocot Root Vascular Cylinder Monocot Stele From: http://www.botany.hawaii.edu/faculty/webb/BOT201/Angiosperm/MagnoliophytaLab99/SmilaxRotM aturePhloemXylem300Lab.jpg Growth of Lateral Roots 100 m Emerging lateral root Cortex Vascular cylinder 1 2 Epidermis Lateral root 3 4 Eudicot & Monocot Roots - External Eudicot – tap root Monocot – fibrous roots Stems Function: - support leaves and flowers - photosynthesis (non-woody plants – herbaceous) - storage: food (tubers – potato) and water (cactus) Stem Structure Nodes: points where leaves are/were attached Internodes: area of growth between the nodes Bud: Developing leaves Terminal/Apical Bud: end of a branch Lateral/Axillary Bud: lateral growth – between leaf petiole (“stem” of leaf) and main stem Bud Scale Scars: Sites of old bud scales (protective layers around the buds) - # of bud scale scars indicates the age of the stem Leaf Scars: Sites where leaves were attached to the stem Lenticles: “bumps” of cork lined pores that allow for oxygen exchange in the stem Terminal bud Bud scale Axillary buds Leaf scar This year’s growth (one year old) Node Stem Internode One-year-old side branch formed from axillary bud near shoot apex Leaf scar Last year’s growth (two years old) Growth of two years ago (three years old) Scars left by terminal bud scales of previous winters Leaf scar Stem: Internal Anatomy Epidermis Ground Tissue Pith Vascular Bundles Contain Xylem and Phloem May contain: Vascular Cambium, Cork Cambium, Sclerenchyma Monocot Stem Structure Ground tissue Epidermis Vascular bundles 1 mm (b) A monocot stem. A monocot stem (maize) with vascular bundles scattered throughout the ground tissue. In such an arrangement, ground tissue is not partitioned into pith and cortex. (LM of transverse section) Monocot Stem Vascular Bundles Xylem Phleom Monocot Stem Vascular Bundle From: http://iweb.tntech.edu/mcaprio/stem_dicot_400X_cs_E.jpg Eudicot Stem Structure Phloem Xylem Sclerenchyma (fiber cells) Ground tissue connecting pith to cortex Pith Key Cortex Epidermis Vascular bundle Dermal Ground Vascular 1 mm (a) A eudicot stem. A eudicot stem (sunflower), with vascular bundles forming a ring. Ground tissue toward the inside is called pith, and ground tissue toward the outside is called cortex. (LM of transverse section) Eudicot Stem Cross Section From: http://plantphys.info/plant_physiology/images/stemcs.jpg Eudicot Stem Vascular Bundle Sclerenchyma Phloem Xylem Vascular Cambium Leaves Functions: - photosynthesis - storage (succulent leaves, Aloe) - protection: spines, toxins, trichomes - reproduction: flowers (modified leaves) Leaves Functions: - photosynthesis - storage (succulent leaves, Aloe) - protection: spines, toxins, trichomes - reproduction: flowers (modified leaves) Leaves: External Structure - - Blade Petiole Stipule Axillary Bud Veins Stipule – growth at the base of petiole Leaves: Internal Structure - Cuticle - Upper Epidermis (Adaxil) - Mesophyll: - Palisade Layer - Spongy Layer - Air Spaces - Vascular Bundle - Bundle Sheath Cells - Xylem and Phloem -Lower Epidermis (Abaxil) - Stomata - Guard Cells - Cuticle Guard cells Key to labels Dermal Ground Stomatal pore Vascular Cuticle Epidermal cell Sclerenchyma fibers 50 µm (b) Surface view of a spiderwort (Tradescantia) leaf (LM) Stoma Upper epidermis Palisade mesophyll Bundlesheath cell Spongy mesophyll Lower epidermis Guard cells Cuticle Vein Xylem Phloem (a) Cutaway drawing of leaf tissues Guard cells Figure 35.17a–c Vein Air spaces Guard cells (c) Transverse section of a lilac 100 µm (Syringa) leaf (LM) Leaf Mesophyll Leaf Stomata Plant Transport Turgor loss in plants causes wilting Which can be reversed when the plant is watered Figure 36.7 Plant Transport of Solutes Proton Pumps: Active transport of H+ out of the cell Builds proton CYTOPLASM – – ATP – EXTRACELLULAR FLUID gradient + H+ + + H+ H+ H+ H+ H+ – – + + H+ Proton pump generates membrane potential and H+ gradient. H+ Functions: provides potential for the COTRANSPORT of materials across the membrane with the H+ – – – H+ + + + H+ H+ H+ H+ H+ H+ – – – H+ + + + Cell accumulates anions ( , for – NO example) by3 coupling their transport to the inward diffusion of H+ through a cotransporter. H+ H+ H+ H+ (b) Cotransport of anions Figure 36.4b H+ H+ S – – + + – + H+ H+ – – H+ – (c) Contransport of a neutral solute Figure 36.4c + + + Plant cells can also accumulate a neutral solute, such as sucrose H+ H+ H+ –+ H H+ S H+ H+ ( S ), by cotransporting H+ down the steep proton gradient. Water Flow from Cell to Cell Water moves between three major compartments of the plant cell. 1. 2. 3. Vacuole – surrounded by Tonoplast Cytosol – surrounded by the Cell Membrane Cell Wall – hydrophilic cellulose – absorbs water Cytosol Tonoplast Vacuole Cell Membrane Cell Wall Three compartments make up three major pathways of transport of water from cell to cell. 1. Apoplastic Route: movement of water and solutes through the cell walls 2. Symplastic Route: transfer of materials from cytosol to cytosol via plasmodesmata 3. Transmembrane Route: movement of water through the walls and cell membranes Key Symplast Apoplast Transmembrane route Apoplast The symplast is the continuum of cytosol connected by plasmodesmata. The apoplast is the continuum of cell walls and extracellular spaces. Symplast Symplastic route Apoplastic route (b) 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. Figure 36.8b Importance of Symplast and Apoplast - provides the route for lateral movement of water from the root epidermis to the vascular cylinder - Water Pathway: - Soil to root epidermis - - In the epidermis water can pass through the cell membrane, enter the symplastic route and travel to the xylem OR it can stay in the cell wall and follow the apoplastic route to the endodermis. Apoplastic Barrier: Endodermis Endodermal walls are infused with suberin (wax) that prevents the water from entering the vascular cylinder The water must enter the cell through the cell membrane and then into the xylem IMPORTANCE: This ensures that all the water and dissolved materials pass through at least one cell membrane before entering the xylem. Casparian strip Endodermal cell Pathway along apoplast Pathway through symplast 1 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. Casparian strip 2 Minerals and water that cross the plasma membranes of root hairs enter the symplast. 1 Plasma membrane Apoplastic route Vessels (xylem) 2 3 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. Symplastic route Root hair 4 Within the transverse and radial walls of each endodermal cell is the Figure 36.9 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. Epidermis Cortex Endodermis Vascular cylinder 5 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. Neither the apoplastic nor symplastic route is continuous to the xylem Apoplastic stops at the endodermis Symplastic stops at the xylem Since xylem cells are dead, the plasmodesmata from the symplastic route will not work so the water must exit the cells via the apoplastic route to go into the xylem walls Vertical Movement Water – Xylem – Pushing and Pulling Hydrostatic Pushing – Root Pressure Roots pump ions and solutes into the roots increasing the solute concentration Lowers the water potential resulting in an influx of water which builds pressure The pressure pushes water up the xylem Only good for short distances and may result in GUTTATION – forcible expulsion of water out of special structures called hydathodes (can be used as a salt gland for plants that live in high saline environments) Transpirational Pull Pulling water up the xylem Transpiration: regulation of the photosynthesis/transpiration compromise by the guard cells and stomata Proper gas exchange causes the loss of water from the air spaces in the spongy mesophyll The drier air space pulls water our of the mesophyll which gets the water from the xylem Water loss from the xylem pulls on the water molecules down the xylem 3 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. Y = –0.15 MPa Y = –10.00 MPa Cell wall Air-water interface Airspace Low rate of transpiration Cuticle High rate of transpiration Upper epidermis Cytoplasm Evaporation Mesophyll Airspace Airspace Cell wall Lower epidermis Evaporation Water film Cuticle CO2 O2 Xylem Water vapor 1 In transpiration, water vapor (shown as blue dots) diffuses from the moist air spaces of the leaf to the drier air outside via stomata. CO2 O2 Stoma Water vapor 2 At first, the water vapor lost by transpiration is replaced by evaporation from the water film that coats mesophyll cells. Vacuole Transpirational pull results from the properties of cohesion and adhesion As one water molecule moves out of the xylem it tugs on the water molecule behind it because they are bound by cohesion forces of the hydrogen bonds between the molecules. Water does not move down the xylem because it is held in place by the adhesive forces between the water and the cellulose of the xylem walls. Xylem sap Outside air Y = –100.0 MPa Mesophyll cells Stoma Water molecule Leaf Y (air spaces) = –7.0 MPa Transpiration Atmosphere Leaf Y (cell walls) = –1.0 MPa Water potential gradient Trunk xylem Y = – 0.8 MPa Xylem cells Cohesion and adhesion in the xylem Adhesion Cell wall Cohesion, by hydrogen bonding Water molecule Root xylem Y = – 0.6 MPa Root hair Soil Y = – 0.3 MPa Soil particle Water uptake from soil Water Other Roles of Transpiration: Evaporative Cooling – helps keep leaves cooler during hot days Factors Affecting Transpiration: Temperature: Hotter = more Humidity: Higher = less Air flow (wind): Higher = more Hormone Signals (Abscisic Acid) – response to dry conditions: Release of hormone closes stomata Regulation of Transpiration: Guard Cells Regulate the size of stomatal openings for gas exchange – responsible for the photosynthesis/transpiration compromise Anatomy of Guard Cell: Eudicots: Kidney shaped Monocots: Dumbbell shaped Both: unevenly thickened cell walls (stomatal side is thicker) 20 µm Figure 36.14 (a) 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. Cells turgid/Stoma open Cells flaccid/Stoma closed Radially oriented cellulose microfibrils Cell wall Vacuole Guard cell Figure 36.15a Physiology Of the Guard Cell Potassium ions are pumped into the vacuole of the guard cell from surrounding cells Higher concentration of K+ reduces the water potential causing an influx of water More water causes the cell to swell Uneven thickness of the cell wall causes the cell to curve and open Loss of water causes the cell to become flaccid and close (b) 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. H2O K+ H2O H2O H2O H2O H2O H2O H2O Figure 36.15b H2O H2O Control of Guard Cells 1. Light stimulation gives energy for H+ pumps 2. 3. Results in the co-transport of K+ CO2 depletion in air space opens stomata Circadian rhythm: internal “clock” – plants kept in the dark still open their stomata when it should be day Stomatal Modifications Xerophytic Plants (dry) Cuticle Lower epidermal tissue Upper epidermal tissue Trichomes (“hairs”) Stomata 100 m Cavitation: Air bubble in the xylem – equivalent of an embolism in an artery – blocks the flow of water – plant reroutes through other xylem Translocation of Phloem Hydrostatic Push from Source to Sink Source: Location of Sugar Production Photosynthesis: Leaves (summer and fall) Starch Metabolism: Roots (spring) Sink: Location of Sugar Consumption or Storage Fall (Roots) Spring (buds for leaf and stem growth) Movement of Phloem Solution Sugar is produced Sugar is cotransported into the cell with H+ ions High H+ concentration H+ Proton pump Figure (b) 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 36.17b sucrose transport to the diffusion of H+ back into the cell. Cotransporter S Key ATP H+ Low H+ concentration H+ Sucrose S Apoplast Symplast Water potential in the cell is lowered Osmotic influx of water into the cell Builds pressure inside of the cell and pushes the solution through the cells to the sink. Vessel (xylem) Sieve tube (phloem) H2O Source cell (leaf) 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 This uptake of water generates a positive pressure that forces the sap to flow along the tube. 3 The pressure is relieved by the unloading of sugar and the consequent loss of water from the tube at the sink. 4 In the case of leaf-to-root translocation, xylem recycles water from sink to source. Sucrose 1 H2O 2 Pressure flow Transpiration stream 4 Sink cell (storage root) 3 Sucrose H2O