2Overview: Underground Plants Stone plants (Lithops) are adapted to life in the desertTwo succulent leaf tips are exposed above ground; the rest of the plant lives below ground
3Overview: Underground Plants (cont.) The success of plants depends on their ability to gather and conserve resources from their environmentThe transport of materials is central to the integrated functioning of the whole plant
4Plant EvolutionAlgal ancestors absorbed minerals and CO2 directly from waterEarly nonvascular land plants lived in shallow water and had aerial shootsNatural selection favored taller plants with flat appendages, multicellular branching roots, and efficient transportevolution of xylem & phloem made long-distance transport of water, minerals, and products of photosynthesis
5Why does over-watering kill a plant? Transport in plantsH2O & mineralstransport in xylemtranspirationevaporation, adhesion & cohesionnegative pressureSugarstransport in phloembulk flowCalvin cycle in leaves loads sucrose into phloempositive pressureGas exchangephotosynthesisCO2 in; O2 outstomatesrespirationO2 in; CO2 outroots exchange gases within air spaces in soilWhy does over-watering kill a plant?
6Ascent of xylem fluidTranspiration pull generated by leaf
7Apoplast & SymplastThe apoplast consists of everything external to the plasma membranecell walls, extracellular spaces, and the interior of vessel elements and tracheidsThe symplast consists of the cytosol of the living cells in a plant, as well as the plasmodesmata
9Water & mineral absorption Water absorption from soilosmosisaquaporinsMineral absorptionactive transportproton pumpsactive transport of H+aquaporinroot hairproton pumpsH2O
10Mineral absorption Proton pumps active transport of H+ ions out of cellchemiosmosisH+ gradientcreates membrane potentialdifference in chargedrives cation uptakecreates gradientcotransport of other solutes against their gradientThe most important active transport protein in the plasma membranes of plant cells is the proton pump , which uses energy from ATP to pump hydrogen ions (H+) out of the cell. This results in a proton gradient with a higher H+ concentration outside the cell than inside. Proton pumps provide energy for solute transport. By pumping H+ out of the cell, proton pumps produce an H+ gradient and a charge separation called a membrane potential. These two forms of potential energy can be used to drive the transport of solutes.Plant cells use energy stored in the proton gradient and membrane potential to drive the transport of many different solutes. For example, the membrane potential generated by proton pumps contributes to the uptake of K+ by root cells.In the mechanism called cotransport, a transport protein couples the downhill passage of one solute (H+) to the uphill passage of another (ex. NO3−). The “coattail” effect of cotransport is also responsible for the uptake of the sugar sucrose by plant cells. A membrane protein cotransports sucrose with the H+ that is moving down its gradient through the protein.The role of proton pumps in transport is an application of chemiosmosis.
11Figure 36.7CYTOPLASMEXTRACELLULAR FLUID(a) H+ and membrane potential+ATPH++Hydrogen ion+H+H+H+H+H++H+Proton pumpH+++(b) H+ and cotransport of neutral solutesSH+H++H+H++H+H+SH+SH+H+H+SS+SH+H+/sucrose cotransporter+Sucrose (neutral solute)++(c) H+ and cotransport of ionsH+H+NO3+NO3+H+H+H+H+NitrateH+NO3H+Figure 36.7 Solute transport across plant cell plasma membranes.NO3+NO3NO3+H+H+NO3 cotransporterH++H++(d) Ion channelsK+Potassium ion+K+K++K+K+K+K++Ion channel+
12Water flow through root Porous cell wallwater can flow through cell wall route & not enter cellsplant needs to force water into cellsCasparian stripThe endodermis, with its Casparian strip, ensures that no minerals can reach the vascular tissue of the root without crossing a selectively permeable plasma membrane. If minerals do not enter the symplast of cells in the epidermis or cortex, they must enter endodermal cells or be excluded from the vascular tissue. The endodermis also prevents solutes that have been accumulated in the xylem sap from leaking back into the soil solution. The structure of the endodermis and its strategic location in the root fit its function as sentry of the border between the cortex and the vascular cylinder, a function that contributes to the ability of roots to transport certain minerals preferentially from the soil into the xylem.
13Controlling the route of water in root Endodermiscell layer surrounding vascular cylinder of rootlined with impermeable Casparian stripforces fluid through selective cell membranefiltered & forced into xylem cells
16Mycorrhizae increase absorption Symbiotic relationship between fungi & plantsymbiotic fungi greatly increases surface area for absorption of water & mineralsincreases volume of soil reached by plantincreases transport to host plant
17Figure 36.5RootsFigure 36.5 A mycorrhiza, a mutualistic association of fungus and roots.Fungus
18MycorrhizaeThe hyphae of mycorrhizal fungi extend into soil, where their large surface area and efficient absorption enable them to obtain mineral nutrients, even if these are in short supply or are relatively immobile. Mycorrhizal fungi seem to be particularly important for absorption of phosphorus, a poorly mobile element, and a proportion of the phosphate that they absorb has been shown to be passed to the plant.
19Transport of sugars in phloem Loading of sucrose into phloemflow through cells via plasmodesmataproton pumpscotransport of sucrose into cells down proton gradient
20Pressure flow in phloem Mass flow hypothesis“source to sink” flowdirection of transport in phloem is dependent on plant’s needsphloem loadingactive transport of sucrose into phloemincreased sucrose concentration decreases H2O potentialwater flows in from xylem cellsincrease in pressure due to increase in H2O causes flowcan flow 1m/hrIn contrast to the unidirectional transport of xylem sap from roots to leaves, the direction that phloem sap travels is variable. However, sieve tubes always carry sugars from a sugar source to a sugar sink. A sugar source is a plant organ that is a net producer of sugar, by photosynthesis or by breakdown of starch. Mature leaves are the primary sugar sources. A sugar sink is an organ that is a net consumer or storer of sugar. Growing roots, buds, stems, and fruits are sugar sinks. A storage organ, such as a tuber or a bulb, may be a source or a sink, depending on the season. When stockpiling carbohydrates in the summer, it is a sugar sink. After breaking dormancy in the spring, it is a source as its starch is broken down to sugar, which is carried to the growing tips of the plant.A sugar sink usually receives sugar from the nearest sources. Upper leaves on a branch may send sugar to the growing shoot tip, whereas lower leaves export sugar to roots. A growing fruit may monopolize sugar sources around it. For each sieve tube, the direction of transport depends on the locations of the source and sink connected by that tube. Therefore, neighboring tubes may carry sap in opposite directions. Direction of flow may also vary by season or developmental stage of the plant.
21Experimentation Testing pressure flow hypothesis using aphids to measure sap flow & sugar concentration along plant stemPressure Flow: The Mechanism of Translocation in AngiospermsPhloem sap flows from source to sink at rates as great as 1 m/hr, much too fast to be accounted for by either diffusion or cytoplasmic streaming. In studying angiosperms, researchers have concluded that sap moves through a sieve tube by bulk flow driven by positive pressure (thus the synonym pressure flow.The building of pressure at the source end and reduction of that pressure at the sink end cause water to flow from source to sink, carrying the sugar along. Xylem recycles the water from sink to source.The pressure flow hypothesis explains why phloem sap always flows from source to sink.
23Figure 36.8Solutes have a negative effect on by binding water molecules.Positive pressure has a positive effect on by pushing water.Solutes and positive pressure have opposing effects on water movement.Negative pressure (tension) has a negative effect on by pulling water.Pure water at equilibriumPure water at equilibriumPure water at equilibriumPure water at equilibriumH2OH2OH2OH2OAdding solutes to the right arm makes lower there, resulting in net movement of water to the right arm:Applying positive pressure to the right arm makes higher there, resulting in net movement of water to the left arm:In this example, the effect of adding solutes is offset by positive pressure, resulting in no net movement of water:Applying negative pressure to the right arm makes lower there, resulting in net movement of water to the right arm:Figure 36.8 Effects of solutes and pressure on water potential () and water movement.Positive pressurePositive pressureNegative pressurePure waterSolutesSolutesMembraneH2OH2OH2OH2O
24Outside air 100.0 MPa Leaf (air spaces) 7.0 MPa Figure 36.13Xylem sapOutside air Mesophyll cells 100.0 MPaStomaLeaf (air spaces)Water molecule 7.0 MPaAtmosphereTranspirationLeaf (cell walls)Adhesion by hydrogen bonding 1.0 MPaXylem cellsCell wallWater potential gradientTrunk xylem Cohesion by hydrogen bonding 0.8 MPaCohesion and adhesion in the xylemFigure Ascent of xylem sap.Water moleculeRoot hairTrunk xylem 0.6 MPaSoil particleSoil WaterWater uptake from soil 0.3 MPa
25water moves into guard cells water moves out of guard cells Control of StomatesEpidermal cellGuard cellChloroplastsNucleusUptake of K+ ions by guard cellsproton pumpswater enters by osmosisguard cells become turgidLoss of K+ ions by guard cellswater leaves by osmosisguard cells become flaccidK+K+H2OH2OH2OH2OK+K+K+K+H2OH2OH2OH2OK+K+Thickened innercell wall (rigid)H2OH2OH2OH2OK+K+K+K+Stoma openStoma closedwater moves into guard cellswater moves out of guard cells
26Control of transpiration Balancing stomate functionalways a compromise between photosynthesis & transpirationleaf may transpire more than its weight in water in a day…this loss must be balanced with plant’s need for CO2 for photosynthesis