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Chapter 10

Transport in multicellular
         plants
Why do multicellular plants
need a transport system?

  like animals – need O2 and
  nutrients

  differ from animals – nature of
  the nutrients needed, gases
  required, and RATE
Particular requirements of plant
              cells

 CO2 - during daylight-
  photosynthesis
 6 CO2 + 6 H2O → C6H12O6 + 6 O2
Particular requirements of plant cells


 Oxygen (O2) – Photosynthesizing
 cells make enough O2 for their own
 needs, but at a slower rate than
 animals
Particular requirements of plant cells


 Organic nutrients (containing
 carbon) – some cells make their
 own but other parts must be
 supplied
Particular requirements of plant cells



 Inorganic ions & H2O → NH4+,
 NO3-, H20, get from roots
Transport of H20
 H20 is transported in xylem tissue
Transport of H20, the Process

 H20 uptake near root tips (root hairs)
 H20 enters xylem
 H20 moves up xylem
 H20 from xylem to leaf cells
 evaporation of H20 into leaf air spaces
 transpiration of H20 vapor through open
 stoma into the atmosphere
Overall
 H20 moves from high to low water
  potential (ψ)
 from soil to root
 from root to leaf
From Soil to Root Hair
 remember Chapter 4, page 61?
Root hairs
 extensions of epidermis
 H20 moves into root hair with the
 water potential (ψ) gradient
    Soil          Root cytoplasm
    Low solute    High solute (proteins, ions, sugars)
    High ψ        Low ψ


       H20
Root hairs
 large # of root hairs
 – large surface area
 - increase rate of osmosis
 - root hairs constantly being
 replaced
Mycorrhizas
 fungi which function like root hairs,
 form a mutualistic symbiotic
 relationship with the root
 - useful in poor soil that can’t
 grow root hairs
 - absorb nutrients, especially
 phosphate
Mycorrhizae
Root Hair to Xylem
H2O movement
   - H2O moves into the root hair through
  epidermal cells
 - H2O must move across cortical cells
  to the xylem tissue in the center of the
  root (Ψ gradient is responsible)
 H2O movement can take two
  pathways
 - which pathway depends on the type of
  plant and condition
Routes from cell to cell
 Moving water & solutes between cells
   transmembrane route
      repeated crossing of plasma membranes
      slowest route but offers more control
   symplast route
      move from cell to cell within cytosol
   apoplast route
      move through connected cell wall without crossing cell
       membrane
      fastest route but never enter cell
Apoplast
 H2O moves through the cortex by
  traveling along criss-cross fibers of
  cellulose fibers located in cortical
  cell walls
     - never enters the cells
Apoplast
Symplast
 H2O enters the cortical cells
 cytoplasm and vacuole
 - moves to the next cell by
 interconnecting plasmodesmata
 channels
 - enters the cells
Symplast
Water & mineral uptake by
            roots
 Mineral uptake by root hairs
   dilute solution in soil
   active transport pumps
      this concentrates solutes (~100x) in root cells

 Water uptake by root hairs
   flow from high H2O potential to low H2O
    potential
   creates root pressure
Route water takes through
               root
 Water uptake by root hairs
  a lot of flow can be through cell wall
   route
  apoplasty
H2O reaches the stele
 Once the H2O reaches the stele
 (center), apoplast pathway stops
 - endodermal cells have a thick
 waterproof waxy band of suberin
 in the cell walls – Casparian strip
 - can only pass into these cells
 by symplast or cytoplasm
 movment
Controlling the route of
        water in root
 Endodermis
  cell layer surrounding vascular cylinder of root
  lined with impervious Casparian strip
  forces fluid through
   selective cell membrane
   & into symplast
     filtered &
     forced into
     xylem vessels
                   Aaaaah…
              Structure-Function
                  yet again!
H2O reaches the stele

 in a young root the suberin
  deposits form bands in the cell
  walls called Caspian strips
 The symplast path remains open
H2O reaches the stele
 in older root, entire cells become
  suberised, closing the symplast
  path too
 Only passage cells are then
  permeable to water
Suberin deposits
 - as cell ages, some cells become
  completely blocked and can only
 use passage cells
 - allows the plant to control
  incoming nutrients
 - builds root pressure
 - H2O crosses the pericycle into
  the xylem
H2O reaches the stele
Xylem
 dual function (support & transport)
 Made up of four different types of cells
   parenchyma cells – standard plant cells, no
    chloroplast
   fibers – elongated cells with dead lignifies
    walls
   vessel elements
   tracheids
Xylem
Plant tissues
 Dermal
   “skin” of plant
   single layer of tightly
    packed cells that
    covers & protects plant
 Vascular
   transport materials
    between roots & shoots
   xylem & phloem
 Ground
   everything else:
    storage, photosynthetic
   bulk of plant tissue
Plant cell types in tissues
Plant cell types in                    Those would’ve

       tissues                         been great names
                                         for my kids!
 Parenchyma
   “typical” plant cells = least specialized
   photosynthetic cells, storage cells
   tissue of leaves, stem, fruit, storage roots
 Collenchyma
   unevenly thickened primary walls = support
 Sclerenchyma
   very thick, “woody” secondary walls =
    support
   rigid cells that can’t elongate
   dead at functional maturity
Plant cell types in tissues
Parenchyma
Parenchyma cells are relatively unspecialized, thin,
flexible & carry out many metabolic functions
   all types of cells develop from parenchyma
Xylem - vessel elements
 make up the xylem vessels
 began as normal cell, but lignin in
  cells
    walls impermeable to water
        - as it builds up, cells die,
  empty lumen left
       - no lignin at plasmodesmata
  – kept open = pits
Xylem - tracheids
 - non living lignified walls
        - not open ended, not tubes
        - H2O moves through pits
         - primitive plants, ferns &
    conifers
Xylem
dead cells →
water-conducting
cells of xylem
Xylem
Xylem
H2O movement in the
       xylem

 H2O crossed cortex, epidermis,
  pericycle into xylem vessels
    - moves up vessels to the
  leaves
    - in root - xylem in center
    - in stem – near the outer
  region
How does H20 get from the
    xylem to the leaf?

 - H20 evaporates from cell walls of
 mesophyll
 - H20 replaced from xylem
Ascent of xylem “sap”
Transpiration pull generated by leaf
Rise of water in a tree by bulk
             flow
 Transpiration pull
    adhesion & cohesion
       H bonding
    brings water &
     minerals to shoot
 Water potential
    high in soil →
     low in leaves
 Root pressure push
    due to flow of H2O
     from soil to root cells
    upward push of
     xylem sap
From leaf to
       atmosphere –
       transpiration
 - inside leaf mesophyll – air spaces
 that are saturated with H20 vapor
 - H20 evaporates fro mesophyll cell
 walls
 - internal leafs spaces are in direct
 contact with the atmosphere through
 the stomata
 - diffuse down the gradient to the
 atmosphere
transpiration

 loss of H20
     - low humidity – transpiration
  HIGH, large gradient
     - high humidity – transpiration
  LOW
     - high temperature/wind –
  transpiration rate HIGH
Control of transpiration
 Stomata function
   always a compromise between
    photosynthesis & transpiration
     leaf may transpire more than its weight in
      water in a day…this loss must be balanced
      with plant’s need for CO2 for photosynthesis
        a corn plant transpires 125 L of water in a
        growing season
Regulation of stomata
 Microfibril mechanism
    guard cells attached at tips
    microfibrils in cell walls
        elongate causing cells to
         arch open = open stomata
        shorten = close when water
         is lost
 Ion mechanism
    uptake of K+ ions by guard
     cells
        proton pumps
        water enters by osmosis
        guard cells become turgid
    loss of K+ ions by guard cells
       water leaves by osmosis
       guard cells become flaccid
 Other cues
   light trigger
       blue-light receptor in plasma membrane of guard cells
        triggers ATP-powered proton pumps causing K+ uptake
           stomates open

   depletion of CO2
       CO2 is depleted during photosynthesis (Calvin cycle)

   circadian rhythm = internal “clock”
       automatic 24-hour cycle
stomata
 stomata open/close – control rate of
  transpiration
 - CO2 must come in, so H20 will leaves
  (HIGH transpiration)
     - bright day, CO2 more needed for
  photosynthesis
     - but then more water loss through
  transpiration
     - plant will have to compromise
Hydrostatic pressure
 pressure exerted by liquid
  - H20 removed from the xylem
 reduces pressure
 - creates a high to low gradient
 - (e.g. suck on a straw, reduces
 pressure at the top, liquid rises)
 - H20 flows up the xylem
Hydrostatic pressure
 - H20 moves by mass flow (moves as
 one body of liquid)
 -H20 molecules attracted to each
 other (H bonds) – cohesion
 -H20 molecules attracted to lignin (H
 bonds) – adhesion
 - air bubbles stop movement (break
 water column) but pits allow H20 to
 move around
Root pressure
 transpiration reduces hydrostatic
 pressure at top of xylem
 - plants also increases root
 pressure to increase pressure
 differences
 - active secretion of solutes,
 mineral ions, into the water of the
 xylem vessels in the root
Root pressure
 cells surrounding the xylem use active
 transport to pump solutes across their
 membranes and into the xylem
 - the presence of solutes lowers the
 water potential drawing water from the
 surrounding root cells
 - increases pressure at the base of
 the xylem vessels
 - root pressure is not essential
Comparing rates of
      transpiration
 - hard to measure H2O vapor
 leaving the plant
 - since most H2O lost by a plant
 was taken in by the roots
 - a measurement of transpiration
 could be approximated by
 measuring the amount of water
 uptake
potometer
 apparatus used to measure uptake
    - must be airtight
    - plant stem in water with rest of
  plant above the airtight seal
      - cut the stem with a slanting cut
      - as water evaporates it is drawn
  into the xylem from the capillary tubing
  and can be measured
     - an experiment can expose the
  plant to varying conditions to compare
  the rate of transpiration
Xerohytes
 plants that live in places where water
  is scarce
 - adaptations include
     -dune grass - eaves can roll up
  exposing a tough waterproof cuticle
  to the outside air while the stomata
  remain open in the enclosed middle of
  the roll
Xerohytes
 hairs help trap a layer of moist air close to
  the leaf surface
     - cactus – flattened photosynthetic
  stem, stores water, leaves are reduced to
  spines which reduces surface area for
  transpiration
    - waxy coating cuts down on water loss
All Cacti are xerophytes
Transverse Section Through Leaf of Xerophytic Plant
Left and right Epidermis of the cactus Rhipsalis
dissimilis.
Left: View of the epidermis surface. The crater-shaped
depressions with a guard cell each at their base can be
seen.
Right: X-section through the epidermis & underlying
tissues. The guard cells are countersunk, the cuticle is
thickened. These are classic xerophyte adaptations.
Marram grass possesses:
rolled leaves, leaf hairs and
sunken stomata. These
adaptations make it resistant
to dry conditions and of
course sand-dunes which
drain very quickly retain very
little water.
BYB3 June 2001 Question 8 part c
BYB3 June 2001 Question 8 part c
           ANSWERS
Translocation
 - term used to describe the
 transport of soluble organic
 substances (assimilates) – such as
 sugars
     - assimilates are transported in sieve tube
     elements in the phloem tissue
Phloem tissue
 types of cells
   sieve (tube) elements
   companion cells
   parenchyma
   fibers
Sieve elements
 sieve tube – made up of many sieve
  elements joined end to end vertically
  to form a continuous column
 a living cell with a cellulose cell wall, a
  plasma membrane, cytoplasm
  containing ER and mitochondria
Sieve elements
 amount of cytoplasm is very small,
  only providing a thin layer inside the
  cell wall,
 no nucleus or ribosomes
 sieve plate – end walls made of
  perforated sieve plate
 pores allow free movement of liquids
Companion cells
 each sieve elements has at least on
 companion cell lying close beside it
 same organelles of other plant cells,
 including small vacuole and nucleus
Companion cells
 contain more mitochondria and
  ribosomes, are metabolically very
  active
 numerous plasmodesmata pass
  through their cell walls making direct
  contact between the cytoplasm of the
  companion cell and sieve element
The contents of phloem
        sieve tubes
 liquid called phloem sap or sap
 not easy to collect phloem sap
 when phloem tissue is cut, the sieve
 elements respond by rapidly blocking
 the sieve pores in a process called
 clotting
phloem sieve tubes
 pores become blocked first by plugs
 of phloem protein strands and then
 the carbohydrate callose
 callose – molecular structure similar
 to cellulose, long chains of glucose
 units linked by β 1-3 glycosidic bonds
Phloem: food-conducting
           cells
 sieve tube elements & companion cells
Aaaaah…

                Phloem
                                        Structure-Function
                                              again!

 Living cells at functional maturity
    lack nucleus, ribosomes & vacuole
      more room: specialized for
      liquid food (sucrose) transport
 Cells
   sieve tubes
      end walls, sieve plates, have pores to
      facilitate flow of fluid between cells
   companion cells
      nucleated cells connected to the sieve-
       tube
      help sieve tubes
sieve plate
        Phloem


sieve
tubes
Phloem: food-conducting cells
 sieve tube elements & companion cells
Vascular tissue in herbaceous
  dicot     stems      monocot
trees & shrubs       grasses & lilies
Root structure: monocot
Root structure: dicot




phloem    xylem
How are aphids used to
       sample sap?
 - aphids have a tubular mouthpart
  – stylets
 - stylet is inserted through the
  surface of the plant’s stem/leaf
  into the phloem
 - phoem sap flows through the
  stylet
Why doesn’t the stylet clot?
 - the diameter of the stylet is so
  small it does not allow the sap to
  flow out rapidly enough to switch
  on the plant’s phloem clotting
  mechanism
 stimulus-response
How translocation occurs
 - like water movement in the xylem,
  sap moves by mass flow
 - mass flow in xylem resulted from
  pressure difference created by the
  water potential gradient between the
  soil and air, this required no energy
  from the plant
 - to create the pressure differences
  needed for mass flow in the phloem, the
  plant has to use energy, an active
  process
How translocation occurs
 - active loading – sucrose is actively
  loaded into the sieve elements , usually
  where photosynthesis takes place
 - as sucrose is loaded, this decreases
  the water potential in the sap
 - through osmosis water enter the sap,
  moving down the water potential
  gradient
 - in the root, (or other area along the
  way), sucrose may be removed, again
  water follows by osmosis
How translocation occurs
 - this creates a pressure difference:
  hydrostatic pressure is high in the part
  of the sieve tube in the leaf and lower in
  the part of the root
 - water flows from high to low taking
  dissolved solutes with it

 source – area of the plant where
  sucrose is loaded
 sink - area of the plant where sucrose
  is taken out
Loading sucrose into phloem
 photosynthesis produces trioses
  which are converted into sucrose
 sucrose, in solution, moves from
  the mesophyll cells to the phloem
 - moves by the symplast pathway
  (cell-to-cell via the plamodesmata)
  or the apoplast pathway traveling
  along the cell walls
Loading sucrose into phloem
 - sucrose is loaded into companion cells
  by active transport
 - ATP is used to pump H+ ions outside
  the companion cell
 - H+ reenter the cell traveling down
  their concentration gradient traveling
  with sucrose (against its through carrier
  proteins (co-transporter proteins) in
  the plasma membrane
Unloading sucrose from
        phloem
 little is known about this process
 - believed sucrose moves from the
  phloem though diffusion down a
  concentration gradient
 - this gradient is maintained through
  cells converting sucrose to something
  else to maintain the gradient
 - invertase – enzyme that hydrolyzes
  sucrose to glucose and fructose
Evidence for the mechanism
    of phloem transport
 - the role of sieve pores and phloem proteins
    were topics of considerable arguments in the
    70’s and 80’s
   - now known phloem proteins are not present
    in living active phloem tissue
   - evidence that sap moves by mass flow is
    considerable
   - rate of phloem transport is 10,000 X faster
    than diffusion
   - actual rates of transport measured match
    closely with those calculated from measured
    pressure differences at source and sink
Evidence for the mechanism
    of phloem transport
 - evidence of ‘active’ loading of sucrose into
  sieve elements in sources such as leaves
 - - phloem sap always has a relatively high pH
  about 8, expected if H+ ions are being
  transported out of the cells
 - - electrical potential of -150mV, consistent
  with an excess of H+ ions outside the cell
  compared to inside
 - - ATP is present in phloem sieve elements in
  large amounts, expected it is required for
  active transport
Comparison of sieve elements
    and xylem vessels
  Similarities
  - liquid moves in mass flow along
   a pressure gradient
  - liquid moves in tubes formed
   from cells stacked end to end
Comparison of sieve
elements and xylem vessels
 Differences
 - water transported through dead xylem
  vessels
 - translocation through live phloem sieve tubes
  requires active loading of sucrose at sources
  thus requiring living cells
 - xylem has lignified walls, phloem does not
 - lignified, dead xylem vessels are entirely
  empty providing a tube through which water
  can flow unimpeded
Comparison of sieve elements
    and xylem vessels
  - lignified, dead xylem vessels have
   strong walls for plant support
  - end walls of elements disappear where
   phloem sieve elements form sieve
   plates which prevent the phloem sieve
   elements from collapsing
  - sieve plates allow the phloem to seal
   itself if damaged, i.e. part of a blade of
   grass is eaten by a herbivore
Comparison of sieve
elements and xylem vessels
 - phloem sap has a high turgor
  pressure because of its high solute
  content, it would leak out quickly if the
  sieve elements did not seal
 - phloem sap contains valuable
  substances such as sucrose, which
  clotting protects from loss
 - clotting may also prevent the entry of
  microorganisms which might feed on
  the nutritious sap or cause disease

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Chapter 10

  • 1. Chapter 10 Transport in multicellular plants
  • 2. Why do multicellular plants need a transport system?  like animals – need O2 and nutrients  differ from animals – nature of the nutrients needed, gases required, and RATE
  • 3. Particular requirements of plant cells  CO2 - during daylight- photosynthesis  6 CO2 + 6 H2O → C6H12O6 + 6 O2
  • 4. Particular requirements of plant cells  Oxygen (O2) – Photosynthesizing cells make enough O2 for their own needs, but at a slower rate than animals
  • 5. Particular requirements of plant cells  Organic nutrients (containing carbon) – some cells make their own but other parts must be supplied
  • 6. Particular requirements of plant cells  Inorganic ions & H2O → NH4+, NO3-, H20, get from roots
  • 7. Transport of H20  H20 is transported in xylem tissue
  • 8. Transport of H20, the Process  H20 uptake near root tips (root hairs)  H20 enters xylem  H20 moves up xylem  H20 from xylem to leaf cells  evaporation of H20 into leaf air spaces  transpiration of H20 vapor through open stoma into the atmosphere
  • 9. Overall  H20 moves from high to low water potential (ψ)  from soil to root  from root to leaf
  • 10. From Soil to Root Hair  remember Chapter 4, page 61?
  • 11. Root hairs  extensions of epidermis  H20 moves into root hair with the water potential (ψ) gradient Soil Root cytoplasm Low solute High solute (proteins, ions, sugars) High ψ Low ψ H20
  • 12. Root hairs  large # of root hairs  – large surface area  - increase rate of osmosis  - root hairs constantly being replaced
  • 13. Mycorrhizas  fungi which function like root hairs,  form a mutualistic symbiotic relationship with the root  - useful in poor soil that can’t grow root hairs  - absorb nutrients, especially phosphate
  • 15. Root Hair to Xylem
  • 16. H2O movement  - H2O moves into the root hair through epidermal cells  - H2O must move across cortical cells to the xylem tissue in the center of the root (Ψ gradient is responsible)  H2O movement can take two pathways  - which pathway depends on the type of plant and condition
  • 17. Routes from cell to cell  Moving water & solutes between cells  transmembrane route  repeated crossing of plasma membranes  slowest route but offers more control  symplast route  move from cell to cell within cytosol  apoplast route  move through connected cell wall without crossing cell membrane  fastest route but never enter cell
  • 18. Apoplast  H2O moves through the cortex by traveling along criss-cross fibers of cellulose fibers located in cortical cell walls  - never enters the cells
  • 20. Symplast  H2O enters the cortical cells cytoplasm and vacuole  - moves to the next cell by interconnecting plasmodesmata channels  - enters the cells
  • 22.
  • 23. Water & mineral uptake by roots  Mineral uptake by root hairs  dilute solution in soil  active transport pumps  this concentrates solutes (~100x) in root cells  Water uptake by root hairs  flow from high H2O potential to low H2O potential  creates root pressure
  • 24. Route water takes through root  Water uptake by root hairs  a lot of flow can be through cell wall route  apoplasty
  • 25. H2O reaches the stele  Once the H2O reaches the stele (center), apoplast pathway stops  - endodermal cells have a thick waterproof waxy band of suberin in the cell walls – Casparian strip  - can only pass into these cells by symplast or cytoplasm movment
  • 26. Controlling the route of water in root  Endodermis  cell layer surrounding vascular cylinder of root  lined with impervious Casparian strip  forces fluid through selective cell membrane & into symplast  filtered & forced into xylem vessels Aaaaah… Structure-Function yet again!
  • 27. H2O reaches the stele  in a young root the suberin deposits form bands in the cell walls called Caspian strips  The symplast path remains open
  • 28. H2O reaches the stele  in older root, entire cells become suberised, closing the symplast path too  Only passage cells are then permeable to water
  • 29. Suberin deposits  - as cell ages, some cells become completely blocked and can only  use passage cells  - allows the plant to control incoming nutrients  - builds root pressure  - H2O crosses the pericycle into the xylem
  • 31. Xylem  dual function (support & transport)  Made up of four different types of cells  parenchyma cells – standard plant cells, no chloroplast  fibers – elongated cells with dead lignifies walls  vessel elements  tracheids
  • 32. Xylem
  • 33. Plant tissues  Dermal  “skin” of plant  single layer of tightly packed cells that covers & protects plant  Vascular  transport materials between roots & shoots  xylem & phloem  Ground  everything else: storage, photosynthetic  bulk of plant tissue
  • 34. Plant cell types in tissues
  • 35. Plant cell types in Those would’ve tissues been great names for my kids!  Parenchyma  “typical” plant cells = least specialized  photosynthetic cells, storage cells  tissue of leaves, stem, fruit, storage roots  Collenchyma  unevenly thickened primary walls = support  Sclerenchyma  very thick, “woody” secondary walls = support  rigid cells that can’t elongate  dead at functional maturity
  • 36. Plant cell types in tissues
  • 37. Parenchyma Parenchyma cells are relatively unspecialized, thin, flexible & carry out many metabolic functions all types of cells develop from parenchyma
  • 38. Xylem - vessel elements  make up the xylem vessels  began as normal cell, but lignin in cells  walls impermeable to water  - as it builds up, cells die, empty lumen left  - no lignin at plasmodesmata – kept open = pits
  • 39. Xylem - tracheids  - non living lignified walls  - not open ended, not tubes  - H2O moves through pits  - primitive plants, ferns & conifers
  • 41. Xylem
  • 42. Xylem
  • 43. H2O movement in the xylem  H2O crossed cortex, epidermis, pericycle into xylem vessels  - moves up vessels to the leaves  - in root - xylem in center  - in stem – near the outer region
  • 44. How does H20 get from the xylem to the leaf?  - H20 evaporates from cell walls of mesophyll  - H20 replaced from xylem
  • 45. Ascent of xylem “sap” Transpiration pull generated by leaf
  • 46. Rise of water in a tree by bulk flow  Transpiration pull  adhesion & cohesion  H bonding  brings water & minerals to shoot  Water potential  high in soil → low in leaves  Root pressure push  due to flow of H2O from soil to root cells  upward push of xylem sap
  • 47.
  • 48.
  • 49. From leaf to atmosphere – transpiration  - inside leaf mesophyll – air spaces that are saturated with H20 vapor  - H20 evaporates fro mesophyll cell walls  - internal leafs spaces are in direct contact with the atmosphere through the stomata  - diffuse down the gradient to the atmosphere
  • 50. transpiration  loss of H20  - low humidity – transpiration HIGH, large gradient  - high humidity – transpiration LOW  - high temperature/wind – transpiration rate HIGH
  • 51. Control of transpiration  Stomata function  always a compromise between photosynthesis & transpiration  leaf may transpire more than its weight in water in a day…this loss must be balanced with plant’s need for CO2 for photosynthesis  a corn plant transpires 125 L of water in a growing season
  • 52. Regulation of stomata  Microfibril mechanism  guard cells attached at tips  microfibrils in cell walls  elongate causing cells to arch open = open stomata  shorten = close when water is lost  Ion mechanism  uptake of K+ ions by guard cells  proton pumps  water enters by osmosis  guard cells become turgid  loss of K+ ions by guard cells  water leaves by osmosis  guard cells become flaccid
  • 53.  Other cues  light trigger  blue-light receptor in plasma membrane of guard cells triggers ATP-powered proton pumps causing K+ uptake  stomates open  depletion of CO2  CO2 is depleted during photosynthesis (Calvin cycle)  circadian rhythm = internal “clock”  automatic 24-hour cycle
  • 54. stomata  stomata open/close – control rate of transpiration  - CO2 must come in, so H20 will leaves (HIGH transpiration)  - bright day, CO2 more needed for photosynthesis  - but then more water loss through transpiration  - plant will have to compromise
  • 55. Hydrostatic pressure  pressure exerted by liquid  - H20 removed from the xylem reduces pressure  - creates a high to low gradient  - (e.g. suck on a straw, reduces pressure at the top, liquid rises)  - H20 flows up the xylem
  • 56. Hydrostatic pressure  - H20 moves by mass flow (moves as one body of liquid)  -H20 molecules attracted to each other (H bonds) – cohesion  -H20 molecules attracted to lignin (H bonds) – adhesion  - air bubbles stop movement (break water column) but pits allow H20 to move around
  • 57. Root pressure  transpiration reduces hydrostatic pressure at top of xylem  - plants also increases root pressure to increase pressure differences  - active secretion of solutes, mineral ions, into the water of the xylem vessels in the root
  • 58. Root pressure  cells surrounding the xylem use active transport to pump solutes across their membranes and into the xylem  - the presence of solutes lowers the water potential drawing water from the surrounding root cells  - increases pressure at the base of the xylem vessels  - root pressure is not essential
  • 59. Comparing rates of transpiration  - hard to measure H2O vapor leaving the plant  - since most H2O lost by a plant was taken in by the roots  - a measurement of transpiration could be approximated by measuring the amount of water uptake
  • 60. potometer  apparatus used to measure uptake  - must be airtight  - plant stem in water with rest of plant above the airtight seal  - cut the stem with a slanting cut  - as water evaporates it is drawn into the xylem from the capillary tubing and can be measured  - an experiment can expose the plant to varying conditions to compare the rate of transpiration
  • 61.
  • 62. Xerohytes  plants that live in places where water is scarce  - adaptations include  -dune grass - eaves can roll up exposing a tough waterproof cuticle to the outside air while the stomata remain open in the enclosed middle of the roll
  • 63. Xerohytes  hairs help trap a layer of moist air close to the leaf surface  - cactus – flattened photosynthetic stem, stores water, leaves are reduced to spines which reduces surface area for transpiration  - waxy coating cuts down on water loss
  • 64. All Cacti are xerophytes
  • 65. Transverse Section Through Leaf of Xerophytic Plant
  • 66. Left and right Epidermis of the cactus Rhipsalis dissimilis. Left: View of the epidermis surface. The crater-shaped depressions with a guard cell each at their base can be seen. Right: X-section through the epidermis & underlying tissues. The guard cells are countersunk, the cuticle is thickened. These are classic xerophyte adaptations.
  • 67. Marram grass possesses: rolled leaves, leaf hairs and sunken stomata. These adaptations make it resistant to dry conditions and of course sand-dunes which drain very quickly retain very little water.
  • 68.
  • 69.
  • 70.
  • 71. BYB3 June 2001 Question 8 part c
  • 72. BYB3 June 2001 Question 8 part c ANSWERS
  • 73. Translocation  - term used to describe the transport of soluble organic substances (assimilates) – such as sugars  - assimilates are transported in sieve tube elements in the phloem tissue
  • 74. Phloem tissue  types of cells  sieve (tube) elements  companion cells  parenchyma  fibers
  • 75. Sieve elements  sieve tube – made up of many sieve elements joined end to end vertically to form a continuous column  a living cell with a cellulose cell wall, a plasma membrane, cytoplasm containing ER and mitochondria
  • 76. Sieve elements  amount of cytoplasm is very small, only providing a thin layer inside the cell wall,  no nucleus or ribosomes  sieve plate – end walls made of perforated sieve plate  pores allow free movement of liquids
  • 77. Companion cells  each sieve elements has at least on companion cell lying close beside it  same organelles of other plant cells, including small vacuole and nucleus
  • 78. Companion cells  contain more mitochondria and ribosomes, are metabolically very active  numerous plasmodesmata pass through their cell walls making direct contact between the cytoplasm of the companion cell and sieve element
  • 79. The contents of phloem sieve tubes  liquid called phloem sap or sap  not easy to collect phloem sap  when phloem tissue is cut, the sieve elements respond by rapidly blocking the sieve pores in a process called clotting
  • 80. phloem sieve tubes  pores become blocked first by plugs of phloem protein strands and then the carbohydrate callose  callose – molecular structure similar to cellulose, long chains of glucose units linked by β 1-3 glycosidic bonds
  • 81. Phloem: food-conducting cells  sieve tube elements & companion cells
  • 82.
  • 83. Aaaaah… Phloem Structure-Function again!  Living cells at functional maturity  lack nucleus, ribosomes & vacuole  more room: specialized for liquid food (sucrose) transport  Cells  sieve tubes  end walls, sieve plates, have pores to facilitate flow of fluid between cells  companion cells  nucleated cells connected to the sieve- tube  help sieve tubes
  • 84. sieve plate Phloem sieve tubes
  • 85. Phloem: food-conducting cells  sieve tube elements & companion cells
  • 86. Vascular tissue in herbaceous dicot stems monocot trees & shrubs grasses & lilies
  • 89. How are aphids used to sample sap?  - aphids have a tubular mouthpart – stylets  - stylet is inserted through the surface of the plant’s stem/leaf into the phloem  - phoem sap flows through the stylet
  • 90. Why doesn’t the stylet clot?  - the diameter of the stylet is so small it does not allow the sap to flow out rapidly enough to switch on the plant’s phloem clotting mechanism  stimulus-response
  • 91. How translocation occurs  - like water movement in the xylem, sap moves by mass flow  - mass flow in xylem resulted from pressure difference created by the water potential gradient between the soil and air, this required no energy from the plant  - to create the pressure differences needed for mass flow in the phloem, the plant has to use energy, an active process
  • 92. How translocation occurs  - active loading – sucrose is actively loaded into the sieve elements , usually where photosynthesis takes place  - as sucrose is loaded, this decreases the water potential in the sap  - through osmosis water enter the sap, moving down the water potential gradient  - in the root, (or other area along the way), sucrose may be removed, again water follows by osmosis
  • 93. How translocation occurs  - this creates a pressure difference: hydrostatic pressure is high in the part of the sieve tube in the leaf and lower in the part of the root  - water flows from high to low taking dissolved solutes with it  source – area of the plant where sucrose is loaded  sink - area of the plant where sucrose is taken out
  • 94. Loading sucrose into phloem  photosynthesis produces trioses which are converted into sucrose  sucrose, in solution, moves from the mesophyll cells to the phloem  - moves by the symplast pathway (cell-to-cell via the plamodesmata) or the apoplast pathway traveling along the cell walls
  • 95. Loading sucrose into phloem  - sucrose is loaded into companion cells by active transport  - ATP is used to pump H+ ions outside the companion cell  - H+ reenter the cell traveling down their concentration gradient traveling with sucrose (against its through carrier proteins (co-transporter proteins) in the plasma membrane
  • 96. Unloading sucrose from phloem  little is known about this process  - believed sucrose moves from the phloem though diffusion down a concentration gradient  - this gradient is maintained through cells converting sucrose to something else to maintain the gradient  - invertase – enzyme that hydrolyzes sucrose to glucose and fructose
  • 97. Evidence for the mechanism of phloem transport  - the role of sieve pores and phloem proteins were topics of considerable arguments in the 70’s and 80’s  - now known phloem proteins are not present in living active phloem tissue  - evidence that sap moves by mass flow is considerable  - rate of phloem transport is 10,000 X faster than diffusion  - actual rates of transport measured match closely with those calculated from measured pressure differences at source and sink
  • 98. Evidence for the mechanism of phloem transport  - evidence of ‘active’ loading of sucrose into sieve elements in sources such as leaves  - - phloem sap always has a relatively high pH about 8, expected if H+ ions are being transported out of the cells  - - electrical potential of -150mV, consistent with an excess of H+ ions outside the cell compared to inside  - - ATP is present in phloem sieve elements in large amounts, expected it is required for active transport
  • 99. Comparison of sieve elements and xylem vessels  Similarities  - liquid moves in mass flow along a pressure gradient  - liquid moves in tubes formed from cells stacked end to end
  • 100. Comparison of sieve elements and xylem vessels  Differences  - water transported through dead xylem vessels  - translocation through live phloem sieve tubes requires active loading of sucrose at sources thus requiring living cells  - xylem has lignified walls, phloem does not  - lignified, dead xylem vessels are entirely empty providing a tube through which water can flow unimpeded
  • 101. Comparison of sieve elements and xylem vessels  - lignified, dead xylem vessels have strong walls for plant support  - end walls of elements disappear where phloem sieve elements form sieve plates which prevent the phloem sieve elements from collapsing  - sieve plates allow the phloem to seal itself if damaged, i.e. part of a blade of grass is eaten by a herbivore
  • 102. Comparison of sieve elements and xylem vessels  - phloem sap has a high turgor pressure because of its high solute content, it would leak out quickly if the sieve elements did not seal  - phloem sap contains valuable substances such as sucrose, which clotting protects from loss  - clotting may also prevent the entry of microorganisms which might feed on the nutritious sap or cause disease

Notas do Editor

  1. The 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.
  2. Functions of the Symplast and Apoplast in Transport How do water and solutes move from one location to another within plant tissues and organs? For example, what mechanisms transport water and minerals from the root hairs to the vascular cylinder of the root? Such short–distance transport is sometimes called lateral transport because its usual direction is along the radial axis of plant organs, rather than up and down along the length of the plant. Three routes are available for this transport. By the first route, substances move out of one cell, across the cell wall, and into the neighboring cell, which may then pass the substances along to the next cell in the pathway by the same mechanism. This transmembrane route requires repeated crossings of plasma membranes as the solutes exit one cell and enter the next. The second route, via the symplast, the continuum of cytosol within a plant tissue, requires only one crossing of a plasma membrane. After entering one cell, solutes and water can then move from cell to cell via plasmodesmata. The third route for short–distance transport within a plant tissue or organ is along the apoplast, the pathway consisting of cell walls and extracellular spaces. Without entering a protoplast, water and solutes can move from one location to another within a root or other organ along the byways provided by the continuum of cell walls.
  3. The 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.
  4. The 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.
  5. The transpiration–cohesion–tension mechanism that transports xylem sap against gravity is an excellent example of how physical principles apply to biological processes. In the long–distance transport of water from roots to leaves by bulk flow, the movement of fluid is driven by a water potential difference at opposite ends of a conduit. In a plant, the conduits are vessels or chains of tracheids. The water potential difference is generated at the leaf end by transpirational pull, which lowers the water potential (increases tension) at the “upstream” end of the xylem. On a smaller scale, water potential gradients drive the osmotic movement of water from cell to cell within root and leaf tissue. Differences in both solute concentration and turgor pressure contribute to this short–distance transport. In contrast, bulk flow depends only on pressure. Another contrast with osmosis, which moves only water, is that bulk flow moves the whole solution, water plus minerals and any other solutes dissolved in the water. The plant expends no energy to lift xylem sap by bulk flow. Instead, the absorption of sunlight drives transpiration by causing water to evaporate from the moist walls of mesophyll cells and by lowering the water potential in the air spaces within a leaf. Thus, the ascent of xylem sap is ultimately solar powered.
  6. Leaves generally have broad surface areas and high surface area–to–volume ratios. The broad surface area is a morphological adaptation that enhances the absorption of light needed to drive photosynthesis. The high surface area–to–volume ratio aids in the uptake of carbon dioxide during photosynthesis as well as in the release of oxygen produced as a by–product of photosynthesis. Upon diffusing through the stomata, CO2 enters a honeycomb of air spaces formed by the spongy parenchyma cells. Because of the irregular shape of these cells, the internal surface area of the leaf may be 10 to 30 times greater than the external surface area we see when we look at the leaf. Although broad surface areas and high surface area–to–volume ratios increase photosynthesis, they also have the serious drawback of increasing water loss by way of the stomata. Thus, a plant’s tremendous requirement for water is part of the cost of making food by photosynthesis. By opening and closing the stomata, guard cells help balance the plant’s requirement to conserve water with its requirement for photosynthesis
  7. Guard cells arbitrate the photosynthesis–transpiration compromise on a moment–to–moment basis by integrating a variety of internal and external stimuli. Even the passage of a cloud or a transient shaft of sunlight through a forest canopy can affect the rate of transpiration.