UNIT%II

TRANSPORT%AND%TRANSLOCATION%OF%WATER%AND%SOLUTES

Topic 2
Water balance%of%plants

! The$soil)plant)atmosphere$continuum
! The$structure$and$properties$of$water.$Water$transport$processes
! Water$in$the$soil
! Water$absorption$by$roots
! Water$transport$through$the$xylem
! Water$movement$from$the$leaf$to$the$atmosphere
Lesson 2:(Water balance(of(plants The soil5plant5atmosphere continuum

Water limits both agricultural and(

Water is a(major factor(in(plant distribution natural(ecosystem productivity

Temperature

Sunlight
Water
Yellow,4orange,4red4and4pink4indicate4that4
water4is4a4key4climate4factor4limiting4plant4growth4

Plants have evolved several strategies to manage water

Some4desert4plants4tolerate4dry4 Most4tracheophytes cannot4

Many4bryophytes4can4tolerate4 Some4desert4plants4evade4
conditions4through4adaptations4such4 tolerate4desiccation4– they4die
desiccation4(i.e.,4drying4to4 drought.4They4survive4the4dry4
as4deep4roots,4C4 photosynthesis,4
equilibrium4with4the4atmosphere) season4as4seeds,4sprouting4and4 succulence4(water4storage)4and4tiny4
flowering4in4a4brief4period4of4rain or4absent4leaves
Lesson 2:(Water balance(of(plants The soil5plant5atmosphere continuum

Water moves by diffusion and(bulk flow

2.(Transport( 3.(Transport( ψw =*
in(the(xylem through(the(leaf( 7100*MPa 3.(Water*
and(into(the(air moves*from*
leaf*to*air

2.(Water*moves*
through*plant
Stoma

ψw =
70.3*MPa 1.(Water*
Endodermis moves*from*
soil*to*root

1.(Water( Vascular* Outer*root*layer

uptake(in(roots cylinder
Diffusion:(Random*movement*of*individual*molecules*(1,(3)

About 97%(of(the water absorbed(by roots is carried through

the plant and(is lost by transpiration through leaf surfaces

Bulk(flow:(water*in*streams,*tubes,*hoses,*water*column*
of*xylem*(1,(2)
Lesson 2:,Water balance,of,plants The structure and,properties of,water.,Water transport processes

The polarity of,water molecules give rise to hydrogen bond

The polarity of,water makes it an excellent solvent
The thermal properties of,water result from hydrogen bonding
The cohesive and,adhesive properties of,water are,due to hydrogen bonding
Cohesion,,adhesion and,surface tension give rise to capillarity
Water has,a,hight tensile strength
Surface tension =,cohesion
Water evaporates
liquid ! gaseous phase (vapour pressure gradient,,humidity +Tª)

3,principles govern water movement in,all living,systems

Mass flow: is the overall movement of,water from a,region of,high concentration to a,region of,low concentration.
The potential of,water to move from a,region of,high to low concentration is called water potential =,".,Pure water "=0
Anything dissolved in,water,,making it less pure,,lower " (All other " than pure water are,negative numbers!)
Species of,chemicals dissolved in,water =,solutes.,Measure of,total,solutes "#
Pressure driven bulk flow drives,long distance water transport
Diffusion: movement of,any molecular,species from a,region of,high to low concentration until the concentrations are,the
same.,Diffusion is the movement of,molecules by random thermal agitation.,Diffusion is rapid over short,distances but
extremely slow over long distances
Osmosis:,diffusion through a,selectively permeable,membrane.,Osmosis,is driven by a,water potential gradient
Lesson 2:(Water balance(of(plants The structure and(properties of(water.(Water transport processes

The chemical potential of(water represents the free(energy status(of(water

Three major factors contribute to cell water potential:(concentration,(pressure and(gravity
!" ,(the solute potential (osmotic potential)(represents solutes dilution of(water and(the reduction of(the free(energy of(water
!p ,(is the hydrostatic pressure of(the solution.(Positive(pressure (turgor pressure)(raises the water potential;(negative pressure,(reduces(it
Lesson 2:(Water balance(of(plants The structure and(properties of(water.(Water transport processes

Plant cells tipically have negative water potentials

Water enters the cell

along a(water potential

Water can(also leave the

cell in(response(to a(
water potential gradient
Lesson 2:(Water balance(of(plants The structure and(properties of(water.(Water transport processes

Water can(also leave the cell in(response(to a(water potential gradient

Water can(also leave the cell in(response(to a(water potential gradient

water enters the cell along a(water potential
Lesson 2:(Water balance(of(plants Water transport in(the plant

Water uptake is a(passive process

It#is#driven#by#differences#of#water#poten.al#between#the# #
atmosphere#(very#dry),#the#inner#plant#.ssues#(wet)#and#the#soil# It#is#controlled#ac.vely#in#the#shoot:#
(wet#3#very#wet)# 3  Leaf:#stomata,#cu.cle#
3  Stem:#xylem#development,#
cavita.on#sensi.vity,#embolism#
recovery#

It#is#controlled#ac.vely#in#the#roots:#
#
3  axial#(from#one#root#segment#to#the#
other)#
3  size#and#number#of#vessels#
#
3  radial#(from#the#xylem#vessels#to#the#
soil)#
3  diameter#of#the#root#
3  presence#of#aquaporins#
3  presence#of#endo3#/#exodermis#
Lesson 2:(Water balance(of(plants Water transport in(the plant

Water transport in(the plant:(from regions of(high water potential to regions of(low water potential

Plumbing system of(the plant =(pathway

Root ! stem ! leaf

Driving force for water movement in(plants is the

evaporation of(water from the leaves =(transpiration

Water is under tension.(Water evaporates

liquid ! gas(phase,(break(the hydrogen bond(
because the gradient is too high.(Water
molecules are(pulled out of(the leaves through
stomata

"p Negative ! tension

"p Positive(! pressure (turgor)

Water movement in(living(systems depends on

both solute concentration and(pressure

"w =("# +("p

Lesson 2:(Water balance(of(plants Water in(the soil

Water moves through the soil by bulk flow

The water content and(rate of(movement in(soils depends on soil type and(structure,(
which influence the pressure gradient in(the soil and(its hydraulic conductivity

Soils&are&made&up& Large&pore&space&(sandy&soil) Small&pore&space&(clayey&soil)

of&particles&of& Gravitational&pull&dominates Capillary&action&contributes&to&
various&sizes,& lateral&movement&
which&affect&how&
Sand water&moves&and&is&
retained&in&them

Silt
Clay
Lesson 2:(Water balance(of(plants Water absortion by roots

The intimate contact between root The type of(soil particles (Shape,(size)(
hairs and(soil particles greatly increases affects soil interactions with water
the surface area for water absortion
Field(capacity:(The&
level&above&which&
Increasing* water&drains&off
water*added*
to*soil
Field(capacity

Wilting(point:(The&
point&at&which&
most&plants&wilt&
Water uptake is mostly confined Wilting(point and&will&not&
recover&at&night
to regions near the root tips
Sand Sandy( Loam Silt( Clay( Clay
loam loam loam
2(mm .002(mm

Decreasing*particle*size

Other&factors&that&affect&water&uptake&from&soil&
include&organic&matter,&microbes,&and&salinity
Lesson 2:(Water balance(of(plants
balance(on plants Water absortion
Water in(the
by roots
soil

Water likely moves from the soil to stele through multiple paths

Apoplastic path((through(wall):
No#crossing#of#plasma#membrane

Symplastic path((through(plasmodesmata):(
Plasma#membrane#crossed#once wall

Transcellular path:
Plasma#membrane#crossed#many#times

Conductance of(cell walls and(membranes affects water uptake

The(exodermis and(endodermis(are(cell(layers(with(Casparian
bands(or(strips(that(restrict(their(conductance(of(water(and(solutes

Key(factors(that(may(affect(movement(
through(the(root(cortex: Xylem
• The#endodermis,-exodermis and#water#
impermeable#Casparian#strip
• Aquaporins (membrane:bound#water#
channels)
• Plasmodesmata-

Plasmodesmata are(small(plasma(membrane–lined
cytoplasmic(bridges(connecting(cells Aquaporins(are(regulated(water(
channels(in(cell(membranes
Lesson 2:(Water balance(of(plants
balance(on plants Water absortion
Water in(the
by roots
soil

Acuoporins facilitate the movement of(water across cell membranes

Channel'activity'is'modified'by'phosphorylation'and'pH
Lesson 2:(Water balance(of(plants Water absortion by roots

Water moves in(the root via the apoplastic

symplastic and(transmembrane pathways Water movement through
apoplast is obstructed by
the Caspary strip in(the
endodermis

Caspariany strip blocks(

apoplastic water flow.(
Water forced to cross a(
plasma(membrane is
effectivelly filtered

Solute accumulation in(the xylem can(generate “root pressure”

Lesson 2:(Water balance(of(plants
balance(on plants Water absortion
Water in(the
by roots
soil

Water flow into roots is affected by

root architecture and(conductance

The distribution of(root growth

is affected by water availability

Root$architecture$(length(of(
roots,(branching,(angle)(affects(
and(is(affected(by(water(
Lesson 2:(Water balance(of(plants Water transport through the xylem

Xylem conduits,(which can(be either single=celled tracheids or multicellular

vessels,(provide a(low resistence pathway for the transport of(water
Lesson 2:(Water balance(of(plants Water transport through the xylem
The cohesion:tension theory explains water transport in(the xylem

!p Negative # tension
!p Positive(# pressure (turgor)

With transpiration:
water is pulled from the xylem into
the cell walls of(leaf mesophyll
before evaporating into the leaf´s air(
spaces.(The xylem is under tension
with a(negative !p

With low transpiration:

the continued transport of(solutes
into the xylem fluid(leads(to a(
decrease in(!" and(!w ,(providing
the force for water absortion and(a(
positive(!p,(which yields a(positive(
hydrostatic pressure in(the xylem
Lesson 2:(Water balance(of(plants Water transport through the xylem

Cavitation breaks the water column and(

prevents the transport of(water under tension

Some(pits(are(more(embolismBresistant(than(others

Plants minimize the consequences of(xylem cavitation

Winter – Freezing)induced-embolisms
Early(spring(–Root)pressure-refilling
Late(spring(– New-xylem-formation
Lesson 2:(Water balance(of(plants Water transport through the xylem

Partial rootzone drying is a(

method to reduce(water use

Well$watered* Exposing*part*of*the*root*system*to*dry*
plant*with*open* conditions*reduces*stomatal*aperture*
stomata*and*high* and*water*use*without*inducing*
transpiration*rate detrimental*drought*stress*effects
Lesson 2:(Water balance(of(plants Water transport through the xylem

Leaf%morphology%by%height

Ψw =(Ψp +(Ψπ +(Ψg

Gravitational(potential(lowers(
Ψw(by(@0.01(MPa per(m(height(((
Lesson 2:(Water balance(of(plants Water movement from the leaf to the atmosphere

Guttation
Lesson 2:(Water balance(of(plants Water movement from the leaf to the atmosphere

The driving force for the water loss is the difference in(water vapour concentration

Water loss is also regulated by the pathway resistences (leaf stomatal resistance and(boundary layer resistance)

Stomatal control(couples leaf transpiration to leaf photosynthesis

Lesson 2:(Water balance(of(plants Water movement from the leaf to the atmosphere

The cell walls of(guard cells have specialized features

Lesson 2:(Water balance(of(plants Water movement from the leaf to the atmosphere
Lesson 2:(Water balance(of(plants Water movement from the leaf to the atmosphere

Stomatal control(couples leaf transpiration to leaf photosynthesis

The transpiration ratio(measures the relationship between water loss and(carbon gain
Lesson 2:(Water balance(of(plants Water movement from the leaf to the atmosphere

Stomatatal opening Stomatatal closing

Plasmatic Cell wall

membrane
Citoplasm K+ Apoplast
d( e Citoplasm
(an arg
Blue(light pH ch
ic d ive(

e
ha (
e c and
rg
A sit Ψπ decrease
po
tiv H(
ga c(p (more(negative)
ne asi
B

Malate K+
Phototropin

H+((pump ATP
H+ K+
H+ Malate
Vacuole ClA ClA ClA
ADP(+(P Ψπ increase
(less negative)
Vacuole
K+
K+ K+ Water
K Channel
+
ClA ADP(+(P
ClA
H+ H+
ClA H+ ATP Glucids
H+
Water PEP Water
CO2
Malate PEPc
OAAA
Water
MalateA
Lesson 2:(Water balance(of(plants Water movement from the leaf to the atmosphere

Abscisic acid (ABA)(closes stomata in(response(to water stress

Lesson 2:%Water balance%of%plants Water movement from the leaf to the atmosphere

ABA%controls%stomatal aperture%by%changing%the%volume%of%guard%cels

When%stomata%are%open,%plants%lose%water%
through%transpiration.%ABA%induced%by%
drought%causes%the%guard%cells%to%close%and%
prevents%their%reopening,%conserving%water%

ABAAinduced%stomatal closure%is%extremely%rapid%and%involves%changes%in%ion%channel%activities

ABA%triggers%an%increase%in%cytosolic calcium%(Ca2+),%which%
activates%anion%channels%(AA)%allowing%ClA to%leave%the%cell.%
ABA%activates%channels%that%move%potassium%out%of%the%cell%
Cl% (K+out)%and%inhibits%%channels%that%move%potassium%into%the%
cell%(K+in).%The%net%result%is%a%large%movement%of%ions%out%of%
AA channel the%cell.

As%ions%leave%the%cell,%so%does%water%(by%osmosis),%causing%
K+in%channel
the%cells%to%lose%volume%and%close%over%the%pore

K+
H2O
Lesson 2:(Water balance(of(plants Water movement from the leaf to the atmosphere

ABA(closes stomata in(response(to water stress

ABA7induced(stomatal closure(is(extremely(rapid(and(involves(changes(in(ion(channel(activities
Lesson 2:(Water balance(of(plants Water movement from the leaf to the atmosphere
Lesson 2:(Water balance(on plants