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SIH1003

Stoma as an
interphase
with
environment

Aim..
Regulation of transpiration
Stomata: structure and
function
Sensing the environment

Soil-Plant-Atmosphere
Continuum

Normal plant water potential


-1 MPa (water content> 75%)

Atmospheric water potential


- 100 MPa
(T = 25oC and RH = 60%)

Water
potential
gradient
is
extremely
high (99
MPa) !!!

Due to the
different water
saturation levels
between
atmosphere and
substomatal
pores, water has
high tendency to
transpire into the
atmosphere. This
effectuates the
substomata to
close, except
when its
necessary to
open.

Water saturate

This interphase gradient is


STOMA.
interphase: the exchange of H2O and CO2
is due to different water concentration (plant
atmosphere).
-- highly temperature dependent
-- Evaporation will lead to leaf cooling
Stomatal interphase is complex: the rate
of efficiency is very low, 1/30 to 1/1000, due
substomatal pores. (higher leaf temperature,
the higher the rate)

Optimisation Theory
Transpiration Photosynthesis Compromise
500 kg soil water to produce 1 kg dry matter - transpiration is a
waste?????

Modus operandii - to regulate the rate of


water loss (i.e. transpiration)
To increase the efficiency of H2O and CO2
exchanges through stomatal pores.

Stomatal Conductance

Boundary layer and stomatal resistances control water loss


from leaf
RESISTANCE THEORY plant as one physical system that
exchanges water & CO2 (refer diagram 1).
Stomatal conductance; leaf stomatal opening
Important factors- diffusion conduction at leaf surface
Stomata function: control the rate of water loss and
recruitment allow influx of CO2 gaseous during photosynthesis
process.
Instrument : Porometer
Unit : cm s-1; gs = 1/rleaf = 1/ra + 1/rb
Conductances: calculated simply as (cm s-1) equivalent to
distance one molecule diffuses in one second, are finite and
easier to quantitate in practise

Heat

H2O

CO2

Boundry
layer
ra

ra

rs
Epidermis

Diagram 1

ra

rs
rc

rc

Mesophyll

rm

ra= Air boundary layer resistance


rs = Stomatal resistance
rc = Cuticular resistance
rm = Mesophyll resistance

ALL RESISTANCES IN

PHOTOSYNTHETIC PROCESS

Ecologically, we
can make some
generalisations
about maximal leaf
conductance
Largely, this will tie
in with the need to
restrict cavitation
and capacity for
the plant to
recharge water
status overnight

Stomata: structure
and function
Antagonism
between guard cell
and epidermal
turgor
Ultrastructural
modifications
alterations in the
shape & vol of
guard cells by
internal changes in
Figure 1:Theradialalignmentofthecellulosemicrofibrilsin
the hydrostatic
guardcellsandepidermalcellsof(A)akidney-shapedstoma
turgor pressure
and(B)agrasslikestoma(source:
Taiz L., Zeiger E., 2010)

Stomata: structure and function


Guardcellsarefoundinleavesofallvascularplants.
In grasses, guard cells have a characteristic dumpbell shape, with bulbous ends
(Figure 1).
These guard cells are always flanked by a pair of differentiated epidermal cells
calledsubsidiary cells,whichhelptheguardcellscontrolthestomatalpores.
In dicots and nongrass monocots, guard cells have an elliptical contour (often
calledkidney-shaped)withtheporeattheircenter.
Subsidiary cells are often absent, the guard cells are surrounded by ordinary
epidermalcells.
Adistinctivefeatureofguardcellsisthespecializedstructureoftheirwalls.
Thealignmentofcellulosemicrofibrils,whichreinforceallplantcellwallsandare
an important determinant of cell shape, play an essential role in the opening and
closingofthestomatalpore.

An increase in guard cell turgor pressure opens the stomata


Guard cells function as multisensory hydraulic valves.
Environmental factors such as light intensity and quality,
temperature, leaf water status, and intracellular CO 2
concentrations are sensed by guard cells, and these signals
are integrated into well-defined stomatal responses.
The early aspects of this process are ion uptake and other
metabolic changes in the guard cells.
The decrease of osmotic potential (s) resulting from ion
uptake and from biosynthesis of organic molecules in the
guard cells. Water relations in guard cells follow the same
rules as in other cells. As s decreases, the water potential
decreases, and water consequently moves into the guard
cells. As water enters the cell, turgor pressure increases.
Because of the elastic properties of their walls, guard cells
can reversible increase their volume by 40 to 100%,
depending on the species. Such changes in cell volume lead
to opening or closing of the stomatal pore.
Subsidiary cells appear to play an important role in allowing
stomata to open quickly and to achieve large apertures.

Dailypatternsofstomatalactivity

Responseofstomatatoenvironmentalconditions(top),andtypicaldailypatterns
ofstomatalopening.(FromSalisburyandRoss,1992)

Stomatal Regulation
How do plants decide on the most appropriate
stomatal aperture for any given situation?
Stomata seem to sense a variety of
environmental parameters and respond
accordingly
The precise response depends on the crop
species and the history of the plants
Environmental factors discussed here: light, CO2,
leaf water status & temperature

Sensing the environment


Feedback from internal CO2 and leaf water content (sensed partly
by carbohydrate supply; hydropassive feedback due to direct
effects on water supply); Abscisic acid a key signal from roots and
mesophyll.
Feedforward : guard cells have chloroplasts (sense light) and
water is evaporated directly around the guard cell complex to

Light
Blue light
Has a direct effect (perceived by guard cells)- similar to the new theory of
stomatal opening
*photoreceptor (flavin, caretenoid or phytochrome, Hinckley & Braatre,
1994)) change the permeability of plasma membrane stomata open
Has indirectly effect PAR increasing photosynthesis and decreasing leaf
internal CO2 (Ci). This will cause K+ ions to be actively pumped into guard
cells, in turn decreasing solute potential in guard cell which causes the water
to enter the guard cells, thus increasing turgor and causing the stomatal
pore to increase in size
A decrease in leaf water potential causes the opposite to occur induce
stomatal closure
Red light
red light photoreceptor (chlorophyll, Zeiger, 1990) modification of
chloroplast metabolism (sugar >) stomata open
similar to the old theory of stomatal opening, which is not totally wrong

CO2

CO2 and stomata opening are inversely related, regardless intensity of


light (Mansfield et al., 1980; Morrison, 1987)
Stomata is more sensitive to Ci, not Ca (Mott, 1988; Mansfield et al.,
1990)
photosynthesis decreases, (i) leaf internal CO2 increases Ci )
(ii) stomatal close
(iii) low rate of water loss

Overall effect : [CO2] WUE


Drastically increasing [CO2] in the air around a leaf will usually cause at
least transient stomatal closure.

Changesinphotosynthesis(AN),stomatalconductance(g)and
leafinternalCO2concentration(ci)ofacornleafinresponseto
fluctuationsinincidentPPFD.(DataofH.J.Earl)

Leaf water status


As leaf water potential drops, stomata tend to
close. Again, this effect may be direct or indirect.
If water potential is very low, the guard cells
themselves, and the surrounding subsidiary cells,
may lose turgor, causing stomatal closure directly.
Much more often, the response to leaf water
potential is indirect. As leaf water potential drops,
the content of ABA in the leaves increases. This
ABA appears to sensitize the stomata to other
signals that would normally cause closing, and so
the average stomatal aperture is decreased.

Evidence that guard cells respond to vapour pressure


independent of leaf water status other evidence?
Shoot water potential is constant, but stomatal
conductance declines in drying air

Relative humidity (RH)


Observation showed when leaf is exposed to low RH,
transpiration rate increased. But this mechanism is still
unclear.
Postulation:

Influence of RH through L (leaf water potential)


RH L gs

E
(Schulze et al., 1987)
Direct effect of RH on i.e. guard cell which is sensitive towards
surrounding RH (ea) and respond accordingly (Turner, 1987).
But the problem with this postulation is that cutical layer is
RH (water) proof.

RH(ea)

Cutical layer (guard


cell is protected from
external RH)
ei cannot
influence
stomata ~
saturated

ei

Substomatalspace

Influence of RH through metabolism processes which provide


energyrequiredforstomatalopeningprocess.e.g.*K +intoguard
cell requires energy, * changes in permeability towards H + also
requiresenergy(Famouspostulation:Tallman,1993)
Stomata has a capability to trace and regulate the rate of water
loss metering volumetric fluxes of H2O (Meinzer & Grantz,
1991).

Leaf temperature

Leaf temperatures may indirectly affect stomatal opening in


several ways:

Changes in temperature affect the photosynthetic rate, and


therefore alter leaf internal CO 2 stomata
High temperatures cause leaf internal vapor pressure and
therefore transpiration to increase.
lead to a reduction in leaf water potential, causing stomata to
close.
Temperature may also have a more direct effect on stomata.
In most species, deleteriously high leaf temperatures may induce
stomatal opening, even when leaf internal CO 2 is not limiting to
photosynthesis. (The effect occurs even in darkness.)
This appears to be a strategy designed to decrease leaf
temperatures through evaporative cooling.