● Vascular plants evolved because of the need of better resource transportation (water to

larger leaves the evaporated more, larger shoots needed better anchorage, larger plants
needed better long-distance transport)
● More stomata = more CO2 and more evaporation
○ adaptatons of palnts strike balance between gas exchange and water loss

Shoot architecture and Light Capture:

● Branching and growing tall affect shoot architecture
○ Plants must balance these to get the most light
● Leaf size and Structure other factor of diversity
○ larger leaves in tropical humid enviros
○ smaller found in dry or cold
■ H2O is scarce and evaporative loss more problematic
● phyllotaxy: arrangement of leaves
○ architectural feature important in light capture
○ determined by shoot apical meristems
○ specific to each species
■ one leaf per node
● alternate- most ANGIOSPERMS
○ leaves in ascending spiral around stem each leaf at
○ 137.5 degrees
■ minimizes shading of lower leaves
● spiral
■ 2 leafs per node
● opposite
■ More
● whorled
○ When too many layers of vegetation, lower leaves respire more than they
photosynthesize
■ nonproductive leaves/branches do programmed cell death
● self-pruning
○ How to reduce self shading?
○ Factors affecting light captrue
■ leaf area index: measurement of this, ratio of total upper leaf surface/ SA
of land on which plant/crop grows
● 7 is common for mature crops
● little benefit of an index >7
○ more causes self-pruning to occur
■ Leaf orientation
● horizontal
● vertical (ex grasses)
 ○ some places, especially grasslands, light too intense for
horizontal leaves
○ light penetrates more deeply to lower leaves

Root Architecture And Acquisition of Water and MInerals

● Roots respond to where soil is mos nutrient rich
○ will penetrate straight through pockets of low nitrate instead of branching within
○ Roots will branch extensively in pockets rich with nitrate
■ Will synthesize more proteins involved in nitrate transport and assimilation
● Roots have reduced competition with roots of the same plant (even cuttings in certain
plants retain ability to recognize itself)

Different mechanisms transport substance over different

distances
● 2 major pathways of transport
○ apoplast
■ everything external to plasma membranes
● cell walls
● extracellular space
● interior of dead cells
○ vessel elements
○ tracheids
○ symplast
■ All cytosol in living cells
■ plasmodesmata
○ 3 routes of transport
■ apoplastic route
■ symplastic route
● only need to cross plasma membrane once
■ Transmembrane
● keep moving through plasma membranes
■ NOT mutually exclusive, some substance may use more than one route
to varying degrees

Short distance transportation

● SOLUTES-across plasma membranes
○ Plants have plasma membranes with same general types of transport proteins as
other cell
○ Difference to animal cells
 ■ Plants use hydrogen iones (H+) unlike sodium ionws (Na+) in animal cels
in basic transport processes
■ membrane potential established though H+ punbping w/ proton pumps
NOT Na+ with sodium-potassium pumps
■ H+ is most often cotransported unlike Na+
● runs absorption of neutral solutes ex sucrose
● cotransport facilitates movement of ions
○ ex NO3- by root cells
■ Also have ion channels like anima cells
● produce electrical signals like action potential in animal
○ In plants, these are 1000 times slower and us Ca2+
activated anion channels not Na+ ion channels in animal
cells
● WATER-across plasma membranes
○ osmosis: how water is absorbed or lost, with the diffusion of FREE water
■ water potential: control direction of flow
● high to low
■ measured in tridents (psi)
● unites: megapascals (1 MPa = 10 MPa)
■ psi of pure water at sea level room temperature = 0MPa
● psi of plant cell = 0.5 MPa (twice air pressure in inflated car tire)
■ water potential = psi solute potential + psi pressure potential
● psi solute potential ALWAYS negative (pure water defined to be 0)
■ Pressure potential is physical pressure
● + or -
● in cells, the protoplast: (living part of cell including plasma
membrane) presses against cell wall creating turgor pressure
○ like air in a tire
○ helps maintain stiffness of plant tissues
○ driving force of cell elongation
● Water in xylem cells often - pressure potential <-2 MPa
○ Dynamic equilibrium is reached when inside outside water pressure is equal
■ know turbid, flaccid/wilted
○ Aquaporins-can open and close affect rate at which water moves across
membrane
■ close more with increases of cytosolic Ca2+ or decreases in cytosolic pH

Long distance tranport: The Role of Bulk Flow

● Diffusion is too slow over long-distance transport
● bulk flow: the movement of liquid in response to pressure gradient
○ always higher to lower pressure
○ INDEPENDENT of solute concentration
 ● occurs within tracheid sand vessel elements of xylem and sieve-tube elements of the
phloem
○ these are either dead holow cells or have very little cloggin organelles
○ assited by perforation plates
○ porous sieve plates

Transpiration

Absorption of Water and Minerals by Root Cells

● Cells near tips of roots particularly important, since most absorption occurs there
○ Epidermal cells permeable to water here
○ Many differentiated into root hairs
■ Rot hairs absorb soil solution (water and dissolved mineral ions not bound
tightly to soil particles)
■ Solution is drawn into hydrophilic walls of epidermal cells
● Pass apoplasticly along the cortex
○ This flow increases SA for absorption than surface of the
epidermis alone
○ Active transport allows roots to accumulate 100x more
minerals that soil with a low mineral concentration

Transport of Water and Minerals into Xylem

● Water and minerals can’t be transported to rest of plant until they inter xylem
○ Endodermis, last layer of cells in root cortex function as las checkpoint for what
gets into vascular cylinder
■ Minerals from symplast were already screened by plasma membrane
they had to cross to enter symplast
■ Apoplastic minerals encounter Casparian strip: suberin waxy belt
impervious to water and solutes in wall of each endodermal cell
■ Water an minerals passively move though selectively permeable plasma
membrane of endodermal cell
● Filters out unneeded and toxic substances
● Prevent solutes in xylem from leaking back out
■ Endodermal cells and live cells in vascular cylinder discharge minerals
from their protoplasts into their own cell walls
● Both diffusion and active transport are involved in this tranger
(solutes form symplast to apoplast)
■ Water and minerals enter tracheids and vessel elements, transproted to
shoot system through builk flow
Bulk Flow Transport via Xylem
● Xylem sap transported by bulk flow to veins n each leaf
● Much faster than diffusion or active transport
○ Peak speed 15-45 m/hr for trees with wide vessel elements
● Involves transpiration
○ Single maize plant transpires 60 L a growing season
● Is xylem sap PUSHED or PULLED?
○ Pushed; Root Pressure
■ Night, with almost no transpiration
● Root cels keep pumping mineral ions into xylem
● Casparian strip prevents leakage
● Accumulation of solutes lowers water potential
○ Water flows in from root cortex, generating root pressure
■ This is a PUSH
■ SOmetimes adds more water than is transpired,
● Causes guttation, root pressure forces
excess water out of leaves
■ Root pressure is mostly minor mechanism, pushes a few meters at most
■ Too weak
■ Root pressure can’t keep up with transpiration after sunrise
○ PULLED: Cohesion-tension hypothesis
■ Functions even if the leaf it reaches is dead for weeks
■ Xylem sap normaully under negative pressure (tension)
■ Stomata on leaf’s surface leads to lots of internal air space (exposes
mesophyll cells to CO2 for photosynthesis)
● Air her is saturated with water vapor because it border moist walls
of cells
● Most days, air outside leaf is drier
○ Outside air has LOWER water potential
○ Water vapor diffuses out the stomata
■ This is transpiration
■ Transpiration produces negative pressure potential that pulls water up
through xylem
■ As water leaves, the water curves more between microfibrils on cell wall,
and surface tension pulls water up, this force is transferred via cohesion
property
● Cohesion attracts water molecules together
○ This gives xylem tensile strength equivalent to steel wire of
same diameter
● Adhesion attracts water to cellulose in xylem walls
● Negative pressure causes diameter of tree trunks to shrink
○ Thick secondary walls of tracheids and bessel elements
prevent them from collapsing
 ■ Transpiration can only pull through UNBROKEN chian of water
moelculues
● Cavitation breaks the change
● More common in wide wessel elements no tracheids
○ Occurs during drought stress or when sap freezes
○ Air bubbles can expand and block water channels
■ Rapid expansion of air bubbles produces clicking
noises
● Chains of H2O can detour between adjacent cells throug pits
● Root pressure lets small plants refill blocked cells
● Speculation: cavitation can even be repaired by sap under
negative pressure
● Only youngest, outermost secondary xylem layers transport water
○ Oldest provides support

Stomata regulation of rate of transpiration

- Leaves generally have large SA
- Aid in photosynthesis
- Spongy mesophyll is very irregular shaped,
- Leafs internal surface may be 10-30 times external SA
- Causes high amount of transpiration
- Plants need for water primarily consequence of need for gas exchange
- Stomata help plant balance water consumption and gas exchange
- Stomata
- Stomatal density is though genetic (ex desert plants) but still environmental
(development plasticity)
- High light and low CO2 leads to increased density in many species
- 95% H2O loss is through these
- Though pores are only 1-2% of external leaf SA
- Waxy cuticle prevents elsewhere
- Guard cells operate on turgidity, water from neighboring cells
- In most angiosperms, cell walls have uneven thickness, and cellulose
microfibrils are oriented specifically
- Guards cells bow outward when turgid.
- Stomata open when guard cells actively accumulate K+ form neighboring cells
- Flow of K+ across plasma membrane of guard cell is power by generation
of membrane potential through proton pumps
- Most K+ and H2O stored in vacuole
- Vacuole membrane has role in regulating guard cell
- Stomata closes when loss of K+ to neighbor cell causes osmotic loss of water
- Vacuole membrane and aquaporins have role in regulation
- Stimuli for Stomatal Opening and Closing
- Generally, stomata open during day, not night
- Only open when photosynthesis can occur
- At least 3 cues contribute to stomatal opening at dawn
- Light
- Causes guard cells to accumulate K+ to become turgid
- Triggered by blue-light receptors in plasma membrane of
guards cells
- Stimulates proton pumps, which promote K+ absorption
- CO2 Depletion
- As photosynthesis decreases CO2, stomata progressively
opens if even H2O is provided
- Internal “clock” in guard cells
- Occurs even if in the dark
-
-
- Environmental stress (drought, high temp, wind, water deficiency)
- Cause stomata to close
- Abscisic acid (ABA): a hormone produced in roots and leaves ins
response to H2O deficiency signals guard cells to close stomata
- Decrease H2O loss, ALSO decrease in photosynthesis
- Also, since turgor is necessary for elongation, growth ceases
- Guard cells control on a MOMENT-TO-MOMENT basis
- Stomata change every second
Effects of transpiration on Wilting and leaf temp
● Extensive wilting can damage leaves
● Transpiration result in evaporative cooling
○ Can lower leaf temp as much as 10C
○ Prevents leaf from reaching denaturing temperatures

Adaptation that reduce water loss

- H2O is most important not because it’s used in photosynthesis, rather it’s required to
keep stomata open to take in CO2
- Xerophytes: plants adapted to arid environments
- Some complete short life cycles during rainy reasons
- Fleshy stems to store water
- Cacti have reduced leaves
- Crassulacean acid metabolism (CAM)
- Specialized form of photosynthesis in succulents of family Crassulaceae
and few others
- Take CO2 in at night, stomata close during day

Sugar transport: sources to sinks via phloem

● Transport of products of photosynthesis known as translocation
● In angiosperms, sieve-tube elements make up paths for translocation
○ Sieve plates are in between, forming long sieve tubes
● Phloem sap:
○ Largest solute is sugar, usually sucrose
■ Concentration can reach 30% by weight
■ Sap is syrupy and thick
○ Can also contain amino acids, hormones, minerals
● Phloem is not unidirectional
● Phloem moves from sugar source to sugar sinks
○ Growing roots, buds, stems, fruits, even leaves and sugar sinks
○ Mature illuminated leaf is sugar source
○ Storage organ, ex tuber or bulb, source or sink depending on season
■ Sink during carb stockpiling in summer
■ After breaking dormancy in spring, it’s a sugar source
● Starch broken down to sugar and carried to growing shoot tips
○ Sinks usually get sugar from NEAREST source
● Directionality:
○ Direction depends on location of sources and sinks
○ Neighboring sieve tubes may carry sap in opposite directions
● Sugars must be loaded into sieve-tube elements before exporting
 ○ Some move from mesophyll to s-t via symplast
○ Some move both symplastic and apoplastic
■ Ex maize
○ Some companion cells have many ingrowths to aid solute transfer
○ Many plants, sugar requires active transport to get into s-t and companion cells
■ Use proton pumps and H+/sucrose cotransport
● Sucrose is unloaded
○ Sugar concentration is always lower in the sink, so sugar molecules simple
diffuse, and water floors with osmosis

Bulk flow by positive pressure: translocation in angiosperms

- Phloem sap speed is max 1 m/hr
- Phloem flows through bulk flow driven by positive pressure, known as pressure flow
- Build pressure at source and reduce pressure at sink to cause flow
- Pressure-flow hypothesis explain why phloem sap flows source to sink
- However study using electron microscope suggest non-flowering vascular plants
may be too small.obstructed to allow pressure flow
- Sometimes too many sinks
- Plants may self-thin, removing sinks
- Ex removing young fruits so trees grow bigger fewer fruits

Symplast is cool (active)

- It’s dynamic and changes according is environment stressors
- Examples
- Change in plasmodesmatal number and pore size
- RECENT studies change idea that plasmodesmata are unchanging
pore-like structure
- Highly dynamic
- Open quickly to turgor pressure
- Cytosolic Ca2+ levels
- Cytosolic pH
- Some plasmodesmata form during cytokinesis but some form
much LATER
- Some disappear during differentiation (loss of function is
common during cell differentiation)
- Pore size can also change,
- Normal pores 2.5 nm
- Plant viruses use viral movement proteins to caus dilation
- 10 nm or more
 - High cytosolic interconnectedness exists only in certain groups and cells
and tissues
- *symplastic domains*
- Info molecules (proteins and RNAs) coordinate
development between each domain
- Disruption of this communication affect development
greatly
- Phloem: information line
- PHloem is “super-highway” for transport of macromolecules ad viruses!
- Entire-system transportation
- System communication is through phloem
- Plasmodesmata are unique to gap junctions in ability to traffic proteins
and RNA
- Electrical Signaling in PHloem
- Rapid-long distance electrical signliang through phloem is another
dynamic feature
- Plants with rapid leaf movments, ex Venus flytrap
- In other plants, migt have nerve-function
- Quick electrica communicaotn between far-away organs