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Water potential
The potential energy of water in a particular environment (the tendency of water to move to a new position)
ψs
solute potential
Always negative (more solute more neg)
Relative to solute potential in pure water
Ψp
pressure potential
tendency of water to move due to physical pressure on the water
Pos or neg
Neg pressure = tension (pulling)
What is the role of the cell wall in determining pressure potential in plants?
Cell wall exerts back pressure when water moves in
Causes inner pressure to build up as turgor pressure
Turgor pressure
Outward pressure of the protoplast against the cell wall
How turgor pressure functions to support the primary plant body
Allows plant cell to hold its shape, otherwise wilting happens
What is considered a turgid cell?
Cells that are exerting enough turgor pressure to induce wall pressure (cell wall pushing back with equal opposite force)
What is the role of the stomata?
Gas exchange – where transpiration occurs (water regulation) + CO2 taken in
Transpiration
Evaporation of water from surface of mesophyll cells and diffusion out of leaf via stomata
When are stomata open/closed
Open when guard cells have full turgor pressure on them, close when guard cells are flaccid
Why is water transport necessary in plants (3 reasons)?
To move water for photosynthesis, turgor pressure, and cellular processes
To move minerals carried from root to shoot
To cool the plant through evaporative cooling
How does water enter the root?
Water enters root through root hairs via osmosis
Soil ψw almost always higher than in roots
How can water move through the root to the xylem (3 options)?
Symplastic
Path through cytoplasm moving between cells via plasmodesmata
Symplast = continuous cytoplasm in plants
Transmembrane
Path through water channels in the plasma membrane (go through pores in plasma membrane)
Apoplastic
Path outside of plasma membrane through cell wall matrix
Apoplast
Continuous cell wall matrix
What is the role of the casparian strip in mediating water uptake in roots?
Blocks the apoplastic route
Water (and solutes) must pass through cytoplasm of endodermal cell to enter vascular tissue
Limits Na+, heavy metals, and pathogens
Casparian strip
band of suberin (waxy substance) on the endodermis
What is root pressure?
The positive upward force generated in plant roots by actively pumping nutrients in that then diffuses water in after it into the xylem
Cohesion-Tension Theory (passive process)
1. Water exits mesophyll cells via evaporation and diffuses out of leaves
2. Drop in water potential inside of mesophyll cell causes net water movement into those cells from surrounding cells
3. Cells bordering the xylem elements replace water with water from xylem
4. Cohesion (H2O molecules sticking together from H-bonds) and adhesion (sticking of H2O to cell walls) lifts water column in the xylem
5. Negative pressure (pulling) reaches to tip of roots, so water moves passively into xylem, then is pulled up the xylem to replace water lost by transpiration
How do plants prevent extensive water loss (3 ways)?
Reduce water loss through stomata
Sunken stomata
Trichomes
Help maintain high humidity at air/leaf interface to reduce water potential gradient
2. Reduce water loss via diffusion across epidermis
Thick cuticle
Multiple layers of epidermis
Reduce SA
3. Timing of stomata opening
CAM (Crassulacean Acid Metabolism)
Stomata are open to allow CO2 uptake at night
Pressure flow model (general description)
Sugars flow in from sources to sinks that are on the same side and region along a pressure gradient (high to low pressure)
How does pressure flow model work?
High pressure at the source
Sugars from source cells are actively transported to and loaded into companion cells and then into sieve tube members
Phloem loading
Water from adjacent xylem cells passively moves into sieve tube element due to more solute entering (turgor pressure increases)
Low pressure at the sink
Sugars are moved out of the sieve tube element and into sink cells
Phloem unloading
Water follows as solute concentration drops in phloem at sink
Turgor pressure falls
How does water flow in pressure flow model?
Water in sieve tube flows from high pressure at source to low pressure at sink
Carries sugar and other solutes with it
Source vs sink
Place where sugars enter system (where they’re either made or stored) vs where sugars exit phloem (where they’re used)
How do sources/sinks change seasonally?
In fall, sinks = roots, trunk, branches, tubers etc.
In spring, sources are sinks from fall
Roots, trunk, branches, tubers send sugars from starch to buds and new shoots
Buds and new shoots = sinks
Phloem loading
Done actively, with cotransporters (secondary transport)
Movement of sucrose coupled with movement of H+ down gradient moving in same direction (symporter)
Phloem unloading
Passively or actively depending on where its being unloaded
Passively
Sucrose moves along concentration gradient out of sieve-tube member to companion cell then developing leaf cell where it’ll be used for photosynthesis
Actively
Sucrose moves along gradient out of sieve-tube member to companion cell then root cell where it is stored in vacuole
High concentration in vacuole, so active transport
Why plants need to uptake mineral nutrients (1 reasons, 4 examples)
To synthesize molecules necessary for growth and reproduction
Amino acids
Nucleic acids
Enzymes
Chlorophylls
Examples of macronutrients
N, P, and K
Micronutrients
Nutrients plants need in relatively small quantities that act as cofactors for specific enzymes (not part of enzyme structure, but required for enzyme to work)
Micronutrients examples
E.g. Cl, Fe, Mn, Zn, etc.
Examples of typical limiting nutrients
Often, N, P, and K
Mobile nutrients
Elements that can move inside the plant tissue to areas of new growth when supplies are low
Mobile nutrient symptoms
Old leaves exhibit nutrient deficiency of these
Plant moves limited nutrients to younger parts
Mobile nutrient examples
N, P, K, and Mg
Immobile nutrient symptoms
Young leaves exhibit nutrient deficiency of immobile nutrients
Nutrients are stuck in old leaves
Immobile nutrient examples
Fe, Ca
Immobile nutrients
Elements that CANNOT move inside the plant tissue to areas of new growth when supplies are low
How does the plant uptake nutrients (describe process)?
SIMPLY: 1) pump H+ out of cell, use secondary transport for + and - ions 2) move inner to reach endo
1. Proton pumps establish electrochemical gradient by moving protons out of cell
Builds up voltage across membrane (membrane potential)
Cations move along gradient, anions move into cell against gradient by coupling movement with protons
2. Ions move from epidermal cells through root cortex to reach endodermis
Endodermis has casparian strip, so all materials must pass thru endodermal cell
What is passive exclusion?
Ions are excluded if no transporter is present
The number of transporters affects how many ions can reach the xylem
Nitrogen fixation
The conversion of N2 to ammonium, nitrate, or nitrite by bacteria and archaea
Why is nitrogen fixation essential?
Eukaryotes can’t use N2, so they need form they can use
How does nitrogen fixation in plants work?
Plants of legume family enter symbiotic relationship with rhizobia (nitrogen fixing bacteria) in root nodules
What is the root nodule?
Bumps that form on the roots of legumes that host rhizobia
What is the role of leghemoglobin?
Binds O2 and keeps O2 concentrations low inside nodules
Protects oxygen-sensitive enzyme nitrogenase needed for fixation
Provides source of O2 for cellular respiration
George Washington Carver’s contributions
Proposed planting crops like peanuts in rotation with cotton grown in the south
allows for a restoration of nutrients to depleted soil
Communicated these strategies with poor farmers
helped increase productivity and reduce poverty in the South
The three sisters
Agricultural system used by native americans involving squash, maize, and beans
How does the three sisters system work?
Planted squash, maize, and beans together
Corn supports beans (trellis for beans to grow on)
Beans fix nitrogen
Squash acts as weed control and shade to the soil
What is the purpose of crop rotation?
To allow for the restoration of nutrients in depleted soil
Rotate with legumes to restore nitrogen in soil
What is a parasitic plant?
Plant that lives in close physical contact with plants from another species and lowers the fitness of those plants by obtaining water or nutrients from them
Haustoria
Structures that penetrate host plant’s vascular tissue to extract water and nutrients
Are parasitic plants heterotrophic?
Can be heterotrophic but most are photosynthetic still
Just use host for water and nutrients
What is an epiphyte?
Plants that grow on top of other plants but are not parasitic (commensal usually) and absorb water and nutrients from collected rainwater, dust, and particles that collect on them
Carnivorous plants
Trap and kill insects, digest them, and absorb nutrients
Found in bogs or habitats with limited nitrogen
Make their own carbs (photosynthetic) but need nitrogen supplement
The general mechanisms by which signals are perceived and transduced.
Step 1: Sensory cells receive external signal
External stimulus is detected by sensory cell, typically by signal receptor proteins
Signal receptor proteins change shape, converting signal to intracellular one
External signal examples
Light, gravity, pressure, day-length, temperature
Step 2: The signal moves through the plant body to target cells
Chemical signals move in the form of hormones (usually)
Step 3: Target cells receive the signal and change their activity in response
Cells must have the appropriate receptor for a response to occur
Most receptor molecules are located in cell membrane
This initiates signal transduction
Involves phosphorylation cascades
Involves secondary messengers
The general characteristics of hormones (4)
Active in small quantities
Can have more than one action
Can act in combo or with non hormonal factors (Ca+)
Can inhibit or stimulate action or growth
What is a hormone?
Organic molecule synthesized by plant in one place that acts in another to initiate physiological response
Phototropic response (stimulus, perception, and signal)
Stimulus
Blue light wavelength
Perception
Perceived at the tip of coleoptile in receptor PHOTOTROPIN 1 (PHOT1)
Growth stimulus (auxin) is produced in tip and transmitted to the growth zone
Signal
Diffuses downward and causes elongation of cells
Called auxin meaning “to grow”
Later identified as Indoleacetic acid (IAA)
Cholodny-Went hypothesis
Phototropic response results from the asymmetrical distribution of auxin
Auxin moves to shaded side in response to light
More auxin transported down shaded side of coleoptile than down side exposed to light
Auxin concentrated on shaded side causes shaded cells to elongate to greater degree
Plant bends toward light
How does the acid growth hypothesis explain cell expansion?
1. IAA (auxin) stimulates proton pumps in membrane
2. Proton pumps secrete proton into cell wall and decreases pH to 4.5
3. Stimulates pH dependent enzyme (expansin) that breaks bonds in cellulose of plant wall
4. Breaking of bonds causes wall to loosen, and turgor pressure causes cell to expand
How do stomata open in response to blue light?
1. Blue light strikes photoreceptor (phototropin)
2. Proton pumps are stimulated
Protons pumped out of guard cells, and an electrochemical gradient builds up
3. K+ and Cl- enter down electrochemical gradient through K+ channels and H+ / Cl- cotransporter
4. Water enters by osmosis
5. Cells swell and pore opens
How does phytochrome act as a molecular switch?
Exhibits photoreversibility
Reversible change in shape in response to light
Switches between Pr and Pfr form depending on what light it absorbs
Pr vs Pfr
Pr (phytochrome red)
Absorbs red light
Inactive form
Pfr (phytochrome far-red)
Absorbs far-red light
Active form that triggers biological response
What responses is phytochrome involved in?
Seed germination
Pr absorbs red light, changes form to Pfr, which activates biological response
Flowering
Phytochrome triggers flowering based on daylength
Controls switch from the vegetative to reproductive meristem
Photoperiodism
Etiolation
Conversion to Pfr switches developmental pattern
Photomorphogenesis
Photomorphogenesis
light-mediated development and growth pattern of plants and other organisms separate from photosynthesis (i.e. germination & flowering) AFTER etiolation
3 categories of plants when it comes to flowering
3 categories
Long day plants
Flower when nights are shorter than certain length
Short-day plants
Flower when nights are longer than certain length
Day-neutral plants
Flower without regard to photo period
Photoperiodism
any response by an organism that is based on photoperiod (relative lengths of day and night)
What is etiolation?
Response by seedling that results in abnormal elongation of stem and no chloroplast development
Significance of etiolation/why its adaptive
Produces maximum growth in length allowed by food reserved to attempt to reach light source in the absence of light
What is gravitropism?
Directional growth of a plant in response to gravity
Auxin’s role in gravitropism
In vertical root, auxin distributed normally
When root tip moved to horizontal position, auxin is redistributed by gravity-sensing cells so that more goes to bottom side
Root bends because higher auxin concentration inhibits cell growth on lower side but stimulates growth on upper side, leading to bending
Statolith hypothesis
Gravity is perceived in the root cap
Root cap cells contain amyloplasts (plastids full of starch)
Amyloplasts accumulate on the bottom of the cell due to gravity (denser than rest of cell parts)
Acts as a statolith - structure that senses gravity
Puts pressure on the cell membrane
How do plants respond to wind and touch?
Thigmomorphogenesis, Thigmotropism, and thigmonasty
Thigmomorphogenesis
Change in development usually due to wind
Whole plant phenotype changes
Thigmotropism
Bending due to touch that is directional
E.g. tendrils on herbaceous vines
Interesting observations about thigmotropism
1. Response can be rapid
5-10 minutes for tendril to enclose
2. Response is long lasting
Brief contact with tendril can induce response that lasts for days
3. Tendrils touched in dark can retain “memory”
Response only occurs later in light, but doesn’t require continued stimulus to occur
Thigmonasty
Very rapid
Involves the generation of an electrical signal by building up membrane potential
Involves rapid changes in turgor pressure
E.g. venus flytrap closing
What is a nastic movement?
Movements that don’t involve growth
Reversible
Nondirectional
What responses is Auxin involved in?
Maintaining apical dominance (inhibits axillary buds)
Why if you cut shoot tip, you cut off production of auxin, so with no auxin to inhibit axillary buds, they start growing
Stimulates formation of adventitious roots
Leaf abscission (inhibits abscission zone)
When leaf damaged, less auxin produced, abscission zone activated, and leaf falls off
cell elongation (e.g. phototropism)
What response is ethylene involved in?
Fruit ripening
Stem elongation
What response is ABA (abscisic acid) involved in?
Seed dormancy
Viviporous (giving live birth) mutants lack ABA gene and therefore never go dormant
Closing stomata in response to drought
How do stomata close?
1. Water stress (condition where plant loses water faster than can take up) stimulates synthesis of the hormone ABA in roots
2. ABA moves up the plant via xylem
3. Pumping by proton pumps in guard cells stops pushing H+ out
4. Changes in membrane potential open Cl- channels
Cl- exits along electrochemical gradient
5. Changes in membrane potential also trigger opening of K+ channels
K+ exits along the electrochemical gradient
6. Water follows by osmosis, guard cells shrink, and stoma closes
Mechanical defense strategies in plants
Cuticle
Trichomes, thorns, and spines
Chemical defense strategies in plants
Secondary metabolites
Usually bi products of metabolic pathways
Peppermint, basil, and sage oil
Pitch (tar like substance from pines and firs)
Latex (milkweeds)
Tannins (cloves, cinnamon, oak)
Opium, caffeine, cocaine, nicotine, THC
Inducible defense
Defensive compounds are EXPENSIVE, so only produce defensive compounds when under attack
Salicylic acid/methyl salicylate (SA)
Involved in pathogen attack response in triggering SAR that is activated by the HR
Systemin
Stimulates production of jasmonic acid
Jasmonic acid (JA)
Stimulates production of gene products that include proteinase inhibitors
How do plants respond to pathogen attack?
HR (hypersensitive response) and SAR (systemic acquired resistance)
HR (Hypersensitive response)
Pathogen enters through wound or stoma
If pathogens recognized by protein receptors, initiates signaling cascade resulting in HR
Stomata closure
Production of toxins
Reinforcement of neighboring cell walls
Rapid cell death of infected cells
SAR (Systemic Acquired Resistance)
Triggered by SA that was triggered by HR
Slower acting, long-term defensive events
Results in changes in gene transcription and increases defense in whole plant
Trigger hormone: Salicylic Acid (SA) derivative Methyl Salicylate (MeSA)
How do plants respond to herbivore attack?
Insect damage → systemin → JA → gene transcription → proteinase inhibitors
Damaged cells produce hormone Systemin that travels thru plant and stimulates Jasmonic Acid production
JA stimulates the production of new gene products including proteinase inhibitors
Block digestive enzymes of herbivores and makes them sick
Herbivores avoid plant that produces proteinase inhibitors
Which responses/processes involve a proton pump?
Nutrient uptake
Stomata opening/closing
Acid-growth hypothesis