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BIOLOGY 125 LECTURE Plant Physiology - Week 10

LECTURE 10.1: ★ Soil structure determined by sand, silt, & clay


WATER ABSORPTION AND TRANSPORT ★ Soil structure affects:
○ Porosity - interconnected channels between
★ Water moves from soil → plant body → air
irregularly shaped soil particles
○ Water moves from high water potential to low
(Pores are 40-60% of soil)
water potential
■ Large pores
○ Water potential is less in the atmosphere than
■ Capillary pores
soil
○ Water Retention
○ Explains upward movement of water
■ Water not readily held in larger pores
○ Considered as soil-plant-atmosphere
(10-60 µm)
continuum
■ If soil is freshly watered, water will
percollate in pores and fill it up
■ Water holding capacity - soil saturated
with water
■ Field capacity - water that remains after
free drainage is complete
● Held in capillary pores
● Plants be watered better at field
capacity

Soil-plant-water continuum

Water transport in Soil


★ Soil
○ Complex medium with solid, liquid, and gas
■ Solid - inorganic particles & organic
materials
○ Aeration
■ Liquid - water solution
■ Air supply in soil
■ Gas - air / gas pockets
■ Sandy soils have larger aeration
■ Algae, bacteria, fungi, earthworms, etc
● Poor water retention,, excellent
aeration
■ Clay soils have less aeration
● Excellent water retention, poor
aeration
■ Loam - 40% sand, 40% silt, 20% clay
● Good soil

Available and Non-available Water in Soil


★ Gravitational water drains out of soil
○ Leaves behind moisture held at soil surfaces
○ Fills small channels of soil
○ These moisture are turned into capillary water
★ Capillary water - principal water source of plants
Soil and Water ○ Attractive forces are not powerful
○ Can be readily absorbed by plants
BIOLOGY 125 LECTURE Plant Physiology - Week 10
★ Hygroscopic water
○ Thin films on soil surfaces
○ Cannot be absorbed by plant

Hygroscopic - can’t be absorbed; Capillary - principal


source; Gravitational - drains away
Formation of Meniscus → Surface tension
★ Permanent Wilting point
○ Proportion of water in soil when plant wilts & ★ Greater attraction with water molecules at air
will not recover unless water is added to soil interface creates pressure on liquid
○ Occurs before all capillary water is absorbed ★ Water is pulled towards roots
■ Some capillary water are held too strongly ○ Soil has higher hydrostatic pressure
for plants to absorb ○ Near the root has lower hydrostatic pressure

Water Potential of Soil ★ More water removed → more acute menisci


★ Two components ○ Makes greater tensions
○ Osmotic potential
■ Negligible
■ Solute concentrations of soil are low
■ Osmotic concentration of soil remain
same
○ Pressure potential (Hydrostatic pressure)
■ Soil water potential is principally
determined by this
★ Uptake of water occurs due to water potential
gradient ○ Develops negative pressure
○ From higher potential to lower
○ Driven principally by pressure gradient
★ Water moves through soil due to pressure ○ T - surface tension of water (7.28*10-8 MPa/m)
difference ○ r - radius of curvature of air-water interface
■ Pressure potential can be negative due to
★ Water-filled pore spaces in soil are interconnected radius of curvature of air-water interface
★ Water is absorbed by roots from center of largest being very small in drying soil
spaces between particles
○ Attractive forces not very powerful and water Water Transport through Roots
is absorbed ★ Most absorption takes place in root hair zone
★ Soil dries out → water recedes to smaller packets ★ Absorption takes place when:
○ Causes surface of soil solution to develop ○ Water potential of root cell sap is more
concave menisci negative than soil solution
○ Brings surface tension to solution ■ If solute concentration of cell is increased
★ Meniscus - formed due to adhesion of water to ■ If turgor pressure is decreased
sides & cohesion to water molecules below ★ Suberized tissues may take water via lenticels
BIOLOGY 125 LECTURE Plant Physiology - Week 10
Lateral Transport Routes ★ Nutrients in form of ions
★ Apoplast Pathway ○ Ions enter soil via epidermis of roots
○ Water moves only through cell wall ○ Root cortex → endodermis → stele
○ No crossing through membrane ★ Ions move via apoplastic route & symplastic
○ Continuous system of cell walls & intercellular route
spaces ★ Casparian band prevents further apoplastic
★ Transmembrane Pathway diffusion through endodermis to stele
○ Enter cell on one side & exit on other ○ Only possible route for ions through
○ Water crosses at least two membranes each endodermis is to enter symplast via carrier /
cell in this path channel mediated transport in cell
○ Transport through tonoplast membrane
★ Symplast Pathway ○ Charged particles - cannot diffuse freely
○ Travel from one cell to another via inside cell
plasmodesmata ○ Transport proteins - in outer tangential wall of
endodermal cell / epidermal or cortical cell
★ Symplastic connection facilitate passive
movement towards stele
○ Unloaded into xylem vessel elements for long
distance transport to other organs

★ Release of ion into xylem require active transport


○ Transfer from symplast to apoplast
○ Carrier mediated process
★ Ion concentration in apoplast of stele is much
higher than in surrounding cortex
○ Ions are accumulated in xylem against
concentration gradient
Path of water movement through root; Symplastic ○ Hence, active transport
ends in xylem & tracheids; Apoplastic ends in phloem ■ Causes water to diffuse to ion
in stele concentrated xylem & stele
■ Water transport by bulk flow
What drives lateral transport of water in root? ○ Casparian bands function to prevent ion loss
★ Osmotic potential gradient from stele
○ Scale relative to outside the stele

Ion Uptake by roots


★ Liquid part of soil is not composed of pure water
○ Contains dissolved mineral nutrients
BIOLOGY 125 LECTURE Plant Physiology - Week 10
Transport of Water Transport of Water In Leaf
★ Water moves from leaves to atmosphere due to
vapor concentration difference
○ From substomatal chamber
○ Steep water vapor gradient

★ Root pressure - pressure developed in tracheary


elements of xylem
○ Dye to metabolic activity of roots
○ Active absorption of salt
○ Not major force in rise of water in plants
■ Not enough to push water to heights
★ Water vapor in cellular spaces > Water vapor in
reached by most trees
atmosphere
■ Absent from conifers
★ Tension → Negative pressure in leaf (spongy layer)
■ Exudation rates is slower than
○ Depleting water in cell drives water to gather
transpiration rates
closer and adhere to cell wall
○ Significant factor in water movement when
○ Forms meniscus
transpiration is poor however (e.g. midnight)
★ Water potential - driving force in water absorption
★ Guttation - exudation of water droplets
★ More concave meniscus, more negative pressure
○ Water absorption > water loss
of water film → greater pulling force
○ Water from root rises into stem → exits
★ Water lost from stomata → replaced by water from
through vein endings (hydathodes)
leaf xylem

Transport of Water through Stems


★ Column of water through xylem cells sticks
★ Cohesion-Tension theory
together due to cohesion
○ Water is not pushed from below but pulled
○ Creates tension within xylem
from above
○ Causes decrease of diameter of tree during
○ Evaporation of water from lead creates
warm day (high transpiration)
negative pressure
○ Similar to sipping through straw
○ Water from below seeks to fill it
○ Driving force for upward movement of water in
Translocation of Water
stem xylem
★ Pure water has tensile strength that can withstand
3MPa required to pull uninterrupted water column
to top of tall trees
○ Cohesion tension theory
★ Cavitation – formation of pockets in xylem
tracheary elements; breaks water column
○ Prevents water transport
○ Cause dehydration and death of leaves
BIOLOGY 125 LECTURE Plant Physiology - Week 10
★ Formation of new growth rings
○ New xylem functions before old xylem ceases
due to gas bubbles

Air pockets caused by cavitation

★ Cavitation can be prevented by


○ Lodging of torus in pit
○ Blocks gas bubbles

Long-distance transport of water from roots to leaves


occurs by bulk flow

LECTURE 10.2:
WATER LOSS

★ Epidermis contain pores


○ Stomatal complex – controls gas exchange &
water loss
○ Managed by guard cells → supported by
subsidiary cells
★ Detour around tracheary elements
○ Blocked cavitated elements ★ Plant must spread leaves
○ Water travels to other tracheary elements ○ Obtain CO2 and sunlight
○ CO2 diffuses in → O2 diffuses out
★ 90% if water loss is through stomata
★ Guard cells control stomatal size
○ Balances plant water conservation

Stomatal Movement
★ Is a direct response to changes in osmotic
potential of guard cell
○ Water moves in → turgid guard cells (open)
○ Water moves out → flaccid guard cells (close)
★ Anatomy of Stoma
○ Cellulose microfibrils making up wall of guard
cells are radially arranged

★ When transpiration is low - pressure potential in


plant increases → water vapor and gases dissolve
back to xylem solutions
○ Some plants develop positive pressures (root
pressure) → shrinks gas bubbles
BIOLOGY 125 LECTURE Plant Physiology - Week 10
○ Guard cell adjacent to wall is thicker than ★ Water movement towards guard cells → increased
outer wall turgidity → stomatal movement
★ Exit of K+ & Cl- in guard cells → Stomatal closing
→ Osmotic loss of water

○ Guard cell shape differs


■ Kidney-shaped - differently thin walls
■ Dumbbell shaped - handles separate;
sleeve between them widens when
opening; similar thin walls
Turgid guard cells (left); Flaccid guard cells (right)

Factors affecting Stomatal movement


★ CO2 concentration
○ When CO2 is used by photosynthesis, CO2
decreases from intercellular spaces
○ Low CO2 in leaf → Stomata opens
○ High CO2 in leaf → Stomata closes

○ Guard cells has chloroplasts ★ Light


○ Opens in presence of blue light
Control of Stomatal Movement ○ Phototropin - perceives blue light via blue light
★ Accumulation of K+ - universal process in opening receptors
stomata ○ Activation of receptors stimulate activity of
○ Accumulates in presence of blue light ATP-powered proton pumps
○ Driven by ATP powered H+ in plasma
membrane ★ Water deficit
■ Proton extrusion hyperpolarizes plasma ○ transpiration > water absorption
membrane ○ Guard cells lose water by direct evaporation
■ Creates pH gradient ○ Stomatal pores close (hydroactive closure)
■ Opens K+ channels ○ Abscisic acid (ABA) in mesophyll signals
★ Charge balanced is achieved by counterion guard cells to close
○ E.g. influx of Cl- ■ Primary regulator of stomatal apparatus in
○ Or by production of organic anion in cell water-stressed plants
■ ABA binding in receptors → intracellular
calcium → blocks K+ in channel & opens
Cl- out channels → membrane
depolarization
○ Proton pump is inhibited
■ Further depolarizes membrane
○ Membrane depolarization activated K+
channels
○ Higher water potential inside guard cell than
outside
■ Water diffuses outwardly
■ Stomates become flaccid & close
Movement of Ions in Guard cells
○ Slower transpiration but lower photosynthesis
BIOLOGY 125 LECTURE Plant Physiology - Week 10
★ Circadian Rhythm
○ Cycle of opening and closing of stomata
○ Approx. 24 hours
○ Some plants removed from natural light and
dark environment continues to exhibit this
cycle
■ Due to internal clock in guard cells

Transpiration
★ Loss of water in vapor form
Humidity
★ Three types
○ Stomatal transpiration
★ Temperature
○ Cuticular transpiration
○ Rise in temperature → high vapor pressure
○ Lenticular transpiration
due to higher motion of molecules → faster
★ Stomatal transpiration contributes greatest
transpiration
★ Two-stage process
○ Evaporation of water from stomata
○ Evaporation of water from moist cell walls into
substomatal air space
★ Wind
○ Diffusion of water vapor from stoma to
○ Quite complex
atmosphere
○ Air in immediate vicinity becomes saturated
★ Driving force of transpiration
with water vapor
○ Difference in water potential between
■ Lowers vapor concentration gradient →
substomatal air space & external atmosphere
Lowers transpiration
○ Vapor movement from higher conc. to lower
○ But, with presence of wind → disperses water
vapor outside leaf
Driving force of Transpiration
■ Does cooling effect on evaporating
★ Concentration of water molecules in vapor phase
surface
be expressed as
■ Lowers vapor pressure gradient &
○ Vapor density (g/m3)
transpiration
○ Vapor pressure (kPa)
■ Wind speed increases
★ Transpiration stream assists in delivery of
○ Before reaching bulk air, water vapor exiting
substances from root to leaves
leaf must diffuse
★ Transpiration → cooling effect
■ Via stomata and boundary layer
○ Lowers temp. of leaves by about 10 Co
■ Boundary layer - layer of unstirred air next
compared to surrounding air
to leaf surface; thickness determined by
○ Prevent enzyme-denaturing temperatures
wind speed
● Still air - thick boundary layer
External Factors affecting rate of Transpiration
● Strong wind - thin boundary layer
★ Light
■ Boundary layer thickness can be affected
○ Stomata opens in presence of light
by other anatomical structures
○ Hence, also increases transpiration
● In some xerophytes - stomata are in
crypts → shelter pores from dry wind
★ Humidity
● Trichomes can break air flow → keeps
○ Actual water content in atmosphere
humidity higher in crypt than in
○ Relative humidity - ratio of actual water
surrounding sphere
content of air to maximum amount of water
that can be held by air at a particular
temperature (RH x 100)
○ External atmosphere is usually unsaturated
○ Steeper gradient → faster transpiration
F. C. Cabuang

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