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Resource Acquisition and Transport in Vascular Plants

Plant Adaptations

  • Adaptations aid in the acquisition of resources, including water, minerals, carbon dioxide, and light

  • Early nonvascular (no xylem & phloem) land plants lived in shallow water and had aerial shoots

    • Nothing could live on land until plants colonized it

    • Xylem: transports water and minerals from roots to shoots

    • Phloem: transports photosynthetic products from where they are made to where they are needed

Shoot Architecture and Light Capture

  • Trade-off between growing tall & branching

    • More energy invested in branching→ less energy available for height growth

  • Water availability signals leaf growth

  • Phyllotaxy: arrangement of leaves on a stem; a species-specific trait important for light capture → more leaves

    • Angle between leaves is 137.5 degrees

      • Minimizes shading of lower leaves

    • If there are too many top leaves, will shade bottom leaves

      • Competition among plants

  • Self-pruning: shedding of lower shaded leaves

    • Occurs when they respire more than they photosynthesize

      • Costs more than it benefits

    • Community: multiple different species living in the same area & interacting

  • Leaf-area index: ratio of total upper leaf surface of a plant divided by the surface area of land on which it grows

    • 7→ shading so self-pruning occurs

    • less leaves→ smaller leaf area index

Root Architecture and Acquisition of Water & Minerals

  • Roots are less competitive with other roots from the same plant than with roots from different plants

    • Nitrates necessary for growth

  • Mycorrhizae: mutualistic associations formed between roots and hyphae of soil fungi (roots/extensions of soil fungi)

    • ex/ increase surface area for absorbing water and minerals, especially phosphate (necessary for making DNA & phospholipids)

Transport

  • 2 major transport pathways

    • Apoplast: everything external to the plasma membrane

      • Includes cell walls, extracellular spaces, and interior of dead cells like vessel elements and tracheids (water-conducting cell in xylem)

    • Symplast: consist of cytosol of all living cells & plasmodesmata

      • everything inside plasma membrane

  • Apoplastic route: through cell walls and extracellular spaces

  • Symplastic route: where water and solutes cross a plasma membrane once and then travel through cytosol

  • Transmembrane route: water and solutes repeatedly cross membranes when they pass from cell to cell

    • Can also occur in cells with the same plasmodesmata (some nutrients may not want to travel through the cytosol)

  • Active transport: need ATP

  • Proton pumps establish membrane potential by pumping H+ and establishing a proton gradient (plants)

  • Membrane potential is established by pumping Na+ by sodium-potassium pumps (animals)

  • Plant cell membranes have ion channels that only allow certain ions to pass

  • Osmosis: the diffusion of water into our out of a cell that is affected by solute concentration and pressure

  • Water potential: quantity that includes effects of solute concentration and physical pressure

    • Determines direction of movement of water

    • Flows from regions of higher water potential to regions of lower water potential

    • Potential→ refers to water’s capacity to perform work

    • Unit of pressure is a megapascal

  • Diffusion: high to low concentration

  • Solute potential: osmotic potential; directly proportional to molarity

  • Pressure potential: physical pressure on a solution (can be positive or negative)

  • Turgor pressure: positive pressure exerted by plasma membrane against cell wall & cell wall against protoplast

    • Protoplast: living part of cell, includes plasma membrane

    • Turgor loss results in wilting, which can be reversed by watering the plant

  • Flaccid: limp/deflated cell

    • If flaccid cell placed in environment with higher solute concentration, cell will lose water and undergo plasmolysis

      • Plasmolysis: protoplast shrinks and pulls away from cell wall

    • If flaccid cell placed in solution with lower solute concentration, cell will gain water and become turgid (plump, lots of water)

  • Aquaporins: transport proteins in cell membrane that facilitate passage of water

    • Opening & closing of aquaporins affect rate of osmotic water movement across membrane

    • Hydrophilic head & hydrophobic tail

  • Bulk-flow: the movement of a fluid driven by a pressure gradient (long-distance transport)

    • Water and solutes move through tracheids and vessel elements (in xylem, tube-like components) of xylem & sieve-tube elements (tube-like element of phloem) of phloem

    • Enhanced by structural adaptations of xylem and phloem

      • Mature tracheids and vessel elements have no cytoplasm; inside of cells empty to move water and minerals (dead cells)

      • Sieve-tube elements have few organelles in their cytoplasm (alive)

    • Perforation plates connect vessel elements & porous sieve plates connect sieve-tube elements

Absorption of Water & Minerals by Root Cells

  • Transpiration: bulk flow up, using xylem; evaporation of water from a plant’s surface

  • Water and mineral absorption occurs near root tips, where there are root hairs and the epidermis is permeable to water

Transport of Water and Minerals into the Xylem

  • Endodermis: innermost layer of cells in the root cortex (external part of root)

    • surrounds vascular cylinder & is last checkpoint for selective passage of minerals from cortex to vascular tissue

  • Casparian strip: barrier at checkpoint of the endodermal wall, blocking apoplastic transfer of minerals from the cortex to the vascular cylinder

  • Endodermal cells discharge water and minerals from their protoplasts into their own cell walls

  • Xylem Sap: fluid in xylem containing water and dissolved minerals

    • transported from roots to leaves by bulk flow

Pushing Xylem Sap: Root Pressure

  • Water flows in from root cortex, generating root pressure (push of xylem sap)

    • Sometimes results in guttation

      • Guttation: exudation of water droplets on tips or edges of leaves

  • Positive root pressure is weak and is a minor mechanism of xylem bulk flow

Pulling Xylem Sap: The Cohesion-Tension Hypothesis

  • Cohesion-tension hypothesis: transpiration and water cohesion pull water from shoots to roots

    • Cohesion: water sticks to water

    • Adhesion: water sticked to other surfaces

      • prevents water from falling back down

    • Surface tension: water won’t spill over & bugs don’t sink in water

  • Xylem sap is normally under negative pressure/tension

  • Transpirational pull: positive + negative pressure

    • Water vapor in the air spaces of leaf diffuse down water potential gradient and exits via stomata

      • Air-water interface retreats into mesophyll cell walls

      • Creates negative pressure potential

    • Negative pressure potential lowers water potential

      • Water molecules pulled from more hydrated areas of lead by negative pressure potential created by air-water interface

    • Positive pressure → pushes up

    • Negative pressure → pulls up

  • Drought stress or freezing can cause a break in chain of water molecules through cavitation

    • Cavitation: formation of water vapor pocket; break in hydrogen bond

Xylem Sap Ascent by Bulk Flow

  • How does bulk flow differ from diffusion?

    • Driven by differences in pressure potential rather than solute potential

    • Occurs in hollow dead cells rather than membranes of living ells

    • Moves the entire solution instead of just water or solutes

    • Much faster

Mechanisms of Stomatal Opening and Closing

  • Changes in turgor pressure open & close stomata

    • Turgid→ guard cells bow outward and pore opens (expand & leave gap)

    • Flaccid→ guard cells become less bowed and pore closes (deflate & cover surface)

  • Generally open during day and close at night to minimize water loss

    • Opening triggered by light, CO2 depletion, and internal clock in guard cells

      • Circadian rhythms: 24 hr cycles; genes that control hormone release are exact

        • ex/ most heart attacks if at risk occur at 10am

    • Hormone Abscisic Acid (ABA) produced in response to water deficiency; causes closure of stomata

  • Sunny, warm, dry, and windy conditions cause evaporation & increase transpiration

    • More water molecules lost to atmosphere

      • Water sometimes leaks through cuticle

    • If uptake & transport can’t replace lost water, plant wilts

  • Transpiration→ evaporative cooling (lower temp of leaf)

    • Trade-off with losing water

    • Water has a high specific heat

Adaptations that Reduce Evaporative Water Loss

  • Xerophytes: plants adapted to dry conditions and climates

    • Crassulacean acid metabolism (CAM): stomatal gas exchange occurs at night

      • Reverse reaction

      • Change CO2 into a 4-carbon molecule and store it

Movement from Sugar Sources to Sugar Sinks

  • Translocation: opposite of transpiration; goes down using phloem

  • Phloem sap: aqueous solution high in sucrose

    • Travels from sugar source to sugar sink

    • Phloem made of sieve-tube elements

    • Moves through bulk flow driven by positive pressure called pressure flow

  • Sugar source: organ that is a net producer of sugar (ex/ mature leaves)

    • Pressure is high

  • Sugar sink: organ that is a net consumer or depository of sugar (ex/ roots, buds, fruits)

    • Sugar must be loaded into sieve-tube elements before being exported to sinks

    • Pressure is low

  • Companion cells: enhance solute movement between apoplast and symplast

  • Self-thinning: dropping of sugar sinks (ex/ flowers, seeds, fruits)

    • Occurs when there are more sugar sinks than sources can support

    • Usually don’t want to get rid of flowers because they attract pollinators

Misc Info

  • Plasmodesmata (plant can have multiple) open and close in response to turgor pressure, cytosolic Ca2+ levels or cytosilic pH

    • plant viruses can cause it to dilate, allowing viral RNA to pass between cells

      • plants try to stop spread of virus by closing plasmodesmata

  • Phloem is key for transport of macromolecules and viruses

    • Systemic communication through phloem allows the integration of plant functions

    • Electricity: movement of any charge (ion)

      • Triggers cells to act/behave differently

T

Resource Acquisition and Transport in Vascular Plants

Plant Adaptations

  • Adaptations aid in the acquisition of resources, including water, minerals, carbon dioxide, and light

  • Early nonvascular (no xylem & phloem) land plants lived in shallow water and had aerial shoots

    • Nothing could live on land until plants colonized it

    • Xylem: transports water and minerals from roots to shoots

    • Phloem: transports photosynthetic products from where they are made to where they are needed

Shoot Architecture and Light Capture

  • Trade-off between growing tall & branching

    • More energy invested in branching→ less energy available for height growth

  • Water availability signals leaf growth

  • Phyllotaxy: arrangement of leaves on a stem; a species-specific trait important for light capture → more leaves

    • Angle between leaves is 137.5 degrees

      • Minimizes shading of lower leaves

    • If there are too many top leaves, will shade bottom leaves

      • Competition among plants

  • Self-pruning: shedding of lower shaded leaves

    • Occurs when they respire more than they photosynthesize

      • Costs more than it benefits

    • Community: multiple different species living in the same area & interacting

  • Leaf-area index: ratio of total upper leaf surface of a plant divided by the surface area of land on which it grows

    • 7→ shading so self-pruning occurs

    • less leaves→ smaller leaf area index

Root Architecture and Acquisition of Water & Minerals

  • Roots are less competitive with other roots from the same plant than with roots from different plants

    • Nitrates necessary for growth

  • Mycorrhizae: mutualistic associations formed between roots and hyphae of soil fungi (roots/extensions of soil fungi)

    • ex/ increase surface area for absorbing water and minerals, especially phosphate (necessary for making DNA & phospholipids)

Transport

  • 2 major transport pathways

    • Apoplast: everything external to the plasma membrane

      • Includes cell walls, extracellular spaces, and interior of dead cells like vessel elements and tracheids (water-conducting cell in xylem)

    • Symplast: consist of cytosol of all living cells & plasmodesmata

      • everything inside plasma membrane

  • Apoplastic route: through cell walls and extracellular spaces

  • Symplastic route: where water and solutes cross a plasma membrane once and then travel through cytosol

  • Transmembrane route: water and solutes repeatedly cross membranes when they pass from cell to cell

    • Can also occur in cells with the same plasmodesmata (some nutrients may not want to travel through the cytosol)

  • Active transport: need ATP

  • Proton pumps establish membrane potential by pumping H+ and establishing a proton gradient (plants)

  • Membrane potential is established by pumping Na+ by sodium-potassium pumps (animals)

  • Plant cell membranes have ion channels that only allow certain ions to pass

  • Osmosis: the diffusion of water into our out of a cell that is affected by solute concentration and pressure

  • Water potential: quantity that includes effects of solute concentration and physical pressure

    • Determines direction of movement of water

    • Flows from regions of higher water potential to regions of lower water potential

    • Potential→ refers to water’s capacity to perform work

    • Unit of pressure is a megapascal

  • Diffusion: high to low concentration

  • Solute potential: osmotic potential; directly proportional to molarity

  • Pressure potential: physical pressure on a solution (can be positive or negative)

  • Turgor pressure: positive pressure exerted by plasma membrane against cell wall & cell wall against protoplast

    • Protoplast: living part of cell, includes plasma membrane

    • Turgor loss results in wilting, which can be reversed by watering the plant

  • Flaccid: limp/deflated cell

    • If flaccid cell placed in environment with higher solute concentration, cell will lose water and undergo plasmolysis

      • Plasmolysis: protoplast shrinks and pulls away from cell wall

    • If flaccid cell placed in solution with lower solute concentration, cell will gain water and become turgid (plump, lots of water)

  • Aquaporins: transport proteins in cell membrane that facilitate passage of water

    • Opening & closing of aquaporins affect rate of osmotic water movement across membrane

    • Hydrophilic head & hydrophobic tail

  • Bulk-flow: the movement of a fluid driven by a pressure gradient (long-distance transport)

    • Water and solutes move through tracheids and vessel elements (in xylem, tube-like components) of xylem & sieve-tube elements (tube-like element of phloem) of phloem

    • Enhanced by structural adaptations of xylem and phloem

      • Mature tracheids and vessel elements have no cytoplasm; inside of cells empty to move water and minerals (dead cells)

      • Sieve-tube elements have few organelles in their cytoplasm (alive)

    • Perforation plates connect vessel elements & porous sieve plates connect sieve-tube elements

Absorption of Water & Minerals by Root Cells

  • Transpiration: bulk flow up, using xylem; evaporation of water from a plant’s surface

  • Water and mineral absorption occurs near root tips, where there are root hairs and the epidermis is permeable to water

Transport of Water and Minerals into the Xylem

  • Endodermis: innermost layer of cells in the root cortex (external part of root)

    • surrounds vascular cylinder & is last checkpoint for selective passage of minerals from cortex to vascular tissue

  • Casparian strip: barrier at checkpoint of the endodermal wall, blocking apoplastic transfer of minerals from the cortex to the vascular cylinder

  • Endodermal cells discharge water and minerals from their protoplasts into their own cell walls

  • Xylem Sap: fluid in xylem containing water and dissolved minerals

    • transported from roots to leaves by bulk flow

Pushing Xylem Sap: Root Pressure

  • Water flows in from root cortex, generating root pressure (push of xylem sap)

    • Sometimes results in guttation

      • Guttation: exudation of water droplets on tips or edges of leaves

  • Positive root pressure is weak and is a minor mechanism of xylem bulk flow

Pulling Xylem Sap: The Cohesion-Tension Hypothesis

  • Cohesion-tension hypothesis: transpiration and water cohesion pull water from shoots to roots

    • Cohesion: water sticks to water

    • Adhesion: water sticked to other surfaces

      • prevents water from falling back down

    • Surface tension: water won’t spill over & bugs don’t sink in water

  • Xylem sap is normally under negative pressure/tension

  • Transpirational pull: positive + negative pressure

    • Water vapor in the air spaces of leaf diffuse down water potential gradient and exits via stomata

      • Air-water interface retreats into mesophyll cell walls

      • Creates negative pressure potential

    • Negative pressure potential lowers water potential

      • Water molecules pulled from more hydrated areas of lead by negative pressure potential created by air-water interface

    • Positive pressure → pushes up

    • Negative pressure → pulls up

  • Drought stress or freezing can cause a break in chain of water molecules through cavitation

    • Cavitation: formation of water vapor pocket; break in hydrogen bond

Xylem Sap Ascent by Bulk Flow

  • How does bulk flow differ from diffusion?

    • Driven by differences in pressure potential rather than solute potential

    • Occurs in hollow dead cells rather than membranes of living ells

    • Moves the entire solution instead of just water or solutes

    • Much faster

Mechanisms of Stomatal Opening and Closing

  • Changes in turgor pressure open & close stomata

    • Turgid→ guard cells bow outward and pore opens (expand & leave gap)

    • Flaccid→ guard cells become less bowed and pore closes (deflate & cover surface)

  • Generally open during day and close at night to minimize water loss

    • Opening triggered by light, CO2 depletion, and internal clock in guard cells

      • Circadian rhythms: 24 hr cycles; genes that control hormone release are exact

        • ex/ most heart attacks if at risk occur at 10am

    • Hormone Abscisic Acid (ABA) produced in response to water deficiency; causes closure of stomata

  • Sunny, warm, dry, and windy conditions cause evaporation & increase transpiration

    • More water molecules lost to atmosphere

      • Water sometimes leaks through cuticle

    • If uptake & transport can’t replace lost water, plant wilts

  • Transpiration→ evaporative cooling (lower temp of leaf)

    • Trade-off with losing water

    • Water has a high specific heat

Adaptations that Reduce Evaporative Water Loss

  • Xerophytes: plants adapted to dry conditions and climates

    • Crassulacean acid metabolism (CAM): stomatal gas exchange occurs at night

      • Reverse reaction

      • Change CO2 into a 4-carbon molecule and store it

Movement from Sugar Sources to Sugar Sinks

  • Translocation: opposite of transpiration; goes down using phloem

  • Phloem sap: aqueous solution high in sucrose

    • Travels from sugar source to sugar sink

    • Phloem made of sieve-tube elements

    • Moves through bulk flow driven by positive pressure called pressure flow

  • Sugar source: organ that is a net producer of sugar (ex/ mature leaves)

    • Pressure is high

  • Sugar sink: organ that is a net consumer or depository of sugar (ex/ roots, buds, fruits)

    • Sugar must be loaded into sieve-tube elements before being exported to sinks

    • Pressure is low

  • Companion cells: enhance solute movement between apoplast and symplast

  • Self-thinning: dropping of sugar sinks (ex/ flowers, seeds, fruits)

    • Occurs when there are more sugar sinks than sources can support

    • Usually don’t want to get rid of flowers because they attract pollinators

Misc Info

  • Plasmodesmata (plant can have multiple) open and close in response to turgor pressure, cytosolic Ca2+ levels or cytosilic pH

    • plant viruses can cause it to dilate, allowing viral RNA to pass between cells

      • plants try to stop spread of virus by closing plasmodesmata

  • Phloem is key for transport of macromolecules and viruses

    • Systemic communication through phloem allows the integration of plant functions

    • Electricity: movement of any charge (ion)

      • Triggers cells to act/behave differently