Chapter 1 Water Relationship in Plant

P201-240 (W.G. Hopkins &N.P.A. Hner) Water absorption from environment , transportation and distribution in plant body, and water loss to atmosphere etcwater relationship Abundant or without harvest dependant on water

Section1 Role of water in plant life 1.1.Structure and physi-chemical characters of water

H -

O
H +

+ H hydrogen bond
H

104.9o

O
O
H

A.Water is a polar molecule with hydrogen bond among water molecules

 B. High specific heat and latent heat of evaporation (heat of vaporization, ) C. Great surface tension() and cohesion(

Figure 1-1 Water mounts along with grass wall by cohesion

D. High dielectric constant() and an extensive solvent ()

Table 10.2 Dielectric constants for some common solvents at 25 C

Water Methanol Ethanol Benzene Hexane

78.4 33.6 24.3 2.3 1.9

D. High dielectric constant() and an extensive solvent ()

+ +
colloidal particle

+ ++ + + colloidal + particle + + + + + + +

Figure 1-2 Hydrophilic colloidal particle and its hydration shell

 1.2 Water content and status in plant 1.2.1 Water content

1.2.1 Water content

1.Plant types Water plants (hydrophytes) >90% Land plants 40-90% Xerophilous plant () ~ 6% Herbs >trees

 1.2 Water content and status in plant 1.2.1 Water content

1.Plant typesWater plants (hydrophytes) >90%, Land plants 40-90%,Xerophilous plant ~ 6%, Herbs >trees 2.Growth environments shade plants>sun plants

 1.2 Water content and status in plant 1.2.1 Water content

3.Organs Stem tenders and root tips>90%, function leaves 70-90%, tree stem 40-50%, dormancy bud 40%, wind-dried seeds 8-14% The higher life activity, the higher water content.

 1.2.2 Status in plant. Free water and bound water free waterIt does not tightly bind to components of cell and it moves freely in the plant. Special characters: participate in metabolism, take as solvent and easily freeze

1.2.2 Status in plant. Free water and bound water free water bound waterIt tightly bind to components of cell and does not move in free in the plant. Special characters: not to participate in metabolism, not to take as solvent and not to freeze easily.

 In plant metabolic activity, growth situation and resistance are all dependent on the ratio of free water to bound water.
The higher ratio, the higher metabolism and the faster growth, but lower resistance because protoplasm is of sol. The lower ratio, the lower metabolism and the slower growth, but higher resistance because protoplasm is of gel.

1.3 Role of water in plant life

(1)Component of protoplasm Protoplasm in plant contains 70-90% water. (2)Substrate for plant metabolism Photosynthesis, respiration and biosynthesis or degradation of organic substance.

1.3 Role of water in plant life

(1)Component of protoplasm (2)Substrate for plant metabolism (3)Solvents for plant absorption and transportation (4)Keeping plant in shape (extension) (5) Balance plant temperature

Section2 Water absorption by plant cell

3 ways: Osmosis absorption(mainly) imbibition absorption; metabolism absorption.

2.1 Osmotic() absorption of water by plant cell

2.1.1 Free energy, chemical potential and water potential bound energy and free energy free energy can work and participate in chemical reaction. Chemical potential: the free energy per mole of that substance. Therefore, water chemical potential is the free energy per mole of water, which is called water potential in plant physiology.

 Water potentialWater potential is defined as the difference in free energy per unit volume, between matrically -bound, pressurized, or osmotically- constrained water and pure water.
w=(w / Vw) - 0wVw) =(w-0w)Vw =wVw

w reflects the capacity for chemical reaction and movement in plant system.

 Suppose: w0 of pure water is zero. w of solution water is minus. The higher concentration, the lower (minus) w . w Unit: MPa=106Pa=10bar Sea water: -2.5M Pa 1M NaCl:4.46 MPaPlant cells:-0.1~ -1.5MPa

 2.1.2 Osmosis and osmotic potential Diffusion():matter transfers from higher concentrations (energy) to lower concentrations (energy)

Figure 1-3 Solute diffuses

 Osmosis () is a diffusion in which solvent molecules pass through semipermeable membrane ().
semipermeable membrane Figure 1-4 See movie for osmosis

 Semipermeable membranevesica, seed coat,

dialysis bag etc.

Osmotic potential ( Solute potential s ) The decreased part of water potential caused by existence of the solute in the solution s(Mpa)= -0.0083iCT iosmotic coefficientNaCl: i=1.80CaCl2: i=2.60,
Sucrose: i=1.

Csolute concentration Tabsolute temperature

 2.1.3 Plant cell is an osmotic system Cell wall ( consists of cellulose,pectin and semi-cellulose)A permeable membrane Protoplastic layer (Plasmic membrane and tonoplast)A semipermeable(selective) membrane Plasmolysis () and Deplasmolysis (

In higher concentration of solution

Plasmolysis

Deplasmolysis
Return to the lower concentration of solution

The protoplast shrinks away from cell wall.

Figure 1-5 Plasmolysis and Deplasmolysis

 Significance for plasmolysis and deplasmolysis Protoplastic layer has selective permeability.

 Protoplastic layer has selective permeability. Judge cell alive or dead cell .

 Protoplastic layer has selective permeability. Judge alive or dead cell from this. Determine cell water potential, and resistance of crop to drought. Determine the entrance speed of substance into cell, easily or difficulty.

 2.1.4 Water potential elements of the plant cell w=s+p+m ssolute potentialDepending on sum of solute particles (molecules or ions)

Normal plant leaf: s=-1 -2 MPa xerophilous plant leaf:s reaches to -10 MPa s has diurnal and seasonal changes

p pressure potential
The increased part of water potential caused by turgor pressure. Normal conditions: Positive value (p>0) Herbs (warm weather+0.3 +0.5MPa in the afternoon +1.5MPa at night Special conditions: zero or minus Incipient plasmolysis=0, Over transpiration <0

turgor pressure

 mmatric potential
The decreased part of water potential results from cell components absorbing water. Minus Wind-dried seed, m -100MPa Obvious m in the cell before formation of

vacuole . Cell with large vacuole >-0.01MPa could be

neglected So, w=s+p in general cell .

1.5

Full water absorption

0.9 1 1.1 1.2 1.3 1.4 1.5

Exceptions: 1intensive transpiration

0.5

w (Mpa)

p<0
2incipient plasmolysis

-0.5

-1

p=0, w= s
3fully turgid cell

-1.5

-2

-2.5

w=0p= -s

incipient Cell volume(times) plasmolysis Figure 1-6 the relation among w, s and p

 2.1.5 Water movement between cells in plantdependent on w

s = -1.2MPa p = 1.0MPa s = -1.0MPa s = -0.8MPa p = 0.9MPa p = 0.4MPa

A s = -1.2MPa p = 1.0MPa w =-0.2MPa

s = -1.0MPa s = -0.8MPa p = 0.9MPa p = 0.4MPa w =-0.1MPa w =-0.4MPa

A C Figure 1-7 Water moves depending on water potential

Water flow direction

Isotonic solution, Hypertonic solution, Hypotonic solution

w-air

w-leaf

w-xylem

Fig 1-8 water potential in soilplantair continuous system

w-root w-soil

 2.2 Imbibing absorption of water of plant cell Imbibition () is a phenomena in which hydrophilic colloids enlarge with water absorption. Only depend on components (hydrophilic group) protein>starch>cellulose> >lipid and fat Soybean has extreme imbibition.

Imbibition is a power of water absorption for vacuole-unformed cell , such as wind-dried seed and meristematic cell. Imbibition is droved by m s=0p=0 w=m

 2.3 Metabolic absorption of water by plant cell The plant cell uses the energy produced in respiration and drives water absorption across plasmatic membraneMetabolic absorption of water Proofs: Respiratory inhibitors (dinitrophenol,DNP and azide, N3-) block water absorption and respiratory promoters (sugar) enhance water absorption.

 2.4 Water channel proteins or aquaporins Aquaporins in all living cell are a serious proteins which located in plasmatic membrane or tonoplast, and play an important role in water transmembrane transport because they have less resistance to water and speed up water transport across the membrane. About 80% of water entrance is controlled by aquaporins.

H2O
H2O Small neutral solutes

P N P A N P P A N A

Fig 1-9 Aquaporins facilitate the diffusion of water and small neutral solutes across plant cell membranes.
The putative structure or an aquaporin monomer with six tilted membrane-spanning domains

 Aquaporins have other possible functions Reproductive grow , cell elongation, guard cell behaviors, cell turgor and volume regulation, transpiration, water cycling in xylem and phloem, nutrition absorption and response to drought and salty.

 Section 3 Absorption of water by plant root 3.1 Absorption region

Main part for absorption of waterthe region of root hair

Lateral root

Root hair

Root cap

Figure 1-10 Model of root tip

1.Great numbers of root hair cells, large absorption area

 2.Thin cell wallbetter water conductivity 3.Well developed conduct tissues.

Figure 1-12 Anatomy of root tip

Why should tree root be maintained with a bulk of original soil when transplanted?

 3.2 Water absorption by rootactive

and passive 3.2.1 Active uptake of water A phenomenon in which water absorption is taken by the physiological activity of root.

Bleeding ()a phenomenon

that the sap flows out from the wounded (cut) partbleeding

sap (see

movie).
Root pressure () is a power which pushes water to mount along vessel, depending on physiological activity of root. 0.1-0.2MPa . Much or less depending on stronger or weaker physiological activities of the root (plant).

Figure 1-13 Guttation

 Guttation ()when soil has enough water and atmosphere is warm and higher relative humidity (RH), often in the early morning, unwounded leaf can secret sap from the tip or margin (water pore) of leaf. Guttation often appears in lotus, strawberry and gramineous crop. An index for healthy seedlings.

Why does root pressure occur?

Casparian band

 Apoplast: A continuous system is consist of cell wall, cell space (interplace) and vessel of xylem, except protoplast, considered as a non-life part in the past. Less resistance and higher speed of transport for water. Symplast: A continuous system is consist of protoplast, plasmodesma and plasmic membrane, considered as a life part except apoplast. Water enters symplast by osmosis and than water is transported across cell by cell.

 3.2.2 Passive uptake of water

Passive uptake is driven by transpiration of leaf

 Power-- Transpiration pull (). Transpiration pulla power driving water upward along xylem vessel is decided by a gradient of water potentials due to transpiration. Independent of root metabolism Main means for water absorption. Especially under the intensive transpiration. But plant can mainly absorb water by active absorption upon low transpiration or without transpiration, such as in the early spring and when the leaves unexpand.

 3.3 Factors affecting water absorption by

root Inner factors w , development degree,
water conductivity and respiration of roots

Outer factors air factors transpiration

water absorption (indirectly) .

Soil factors directly absorption of root.

influence

water

 (1) Soil available water (is referred as the water can directly be taken up and utilized by plants, whose water content is higher than wilting coefficient in the soil. wilting coefficient() is a soil water content (%) under which plant will occur wilting permanently . Under the condition of water deficiency, leaves and tender stems will loss their turgor, called wilting.

Figure 1-16 Plant wilting

 Temporary wilting () The wilting is caused by loss of equilibrium between water absorption and evaporation (main transpiration). Transpiration is larger than absorption. Normal status can be recovered by shading, or in the evening upon decreasing in transpiration, but not by watering. Permanent wilting ()The wilting is caused by no soil available water, plant can not absorb water from the soil. If the permanent wilting just happened, normal status can be recovered by watering or water spraying, but not by decreasing in transpiration.

 (2) Soil O2 CO2 N2 treatmentabsorption because O2 , respiration ,active absorption, anaerobic respiration, Ethanol accumulation, root toxication.

 (3) Soil temperature

Low temperature: water and plasma viscosity(), water conductance respirationenergy not enough root growth and root hair Too high Troot corkification High easily water conductance

Uptake rate of water

Low Temperature

Optimum

 (4) Soil solute concentration

w in root < w in soil, usually soil >-0.1MPa.

Why should we not apply a large number of fertilizer to plant in one time?

 Section 4 Transpiration More than 95% of water loss in air, and only1-5% for plant metabolism. (1)liquid form--guttation (2)gas form--transpiration Transpiration ()is a process of loss water from plant in a form of water vapor.

 4.1 Organs for transpiration Lenticular transpiration ( ) about 0.1% Most of transpiration passes throughout leaf of plantcalled Leaf transpiration.

4.1 Organs for transpiration Lenticular transpiration *Leaf transpiration : Cuticular transpiration ( ), 5-10 Stomatal transpiration (),90-95%

 4.2 Stomatal transpiration 4.2.1 Size, number and distribution of stomata Stomata ()pore for gas exchange (main CO2, O2 , Water vapor)

Figure 1-14 Stomata in the lower epidermis of potato leaf

Table1-2 The number, size and distribution of stomata in plants

Plants Wheat Maize Oat Sun flower Tomato Bean Apple Lotus Number/ epiderm (mm2) Size() Single area Total area The upper The lower Lengthwidth ( 2) /leaf area (%) 33 14 387 209 0.52 52 68 195 75 0.82 25 23 388 239 0.98 58 156 228 136 3.13 12 130 136 61 0.85 40 281 7 3 17 0.84 0 400 1412 132 5.28 46 0 -

 Upper epidermis typehydrophyteslotus Lower epidermis type most trees:apple and

peach trees.
Both epidermis type most herbs including crops. But stomata are in the lower epidermis more than in the upper epidermis. In grain plants, those distribution is nearly equal in the lower epidermis to in the upper epidermis.

 4.2.2 Stomatal diffusionLaw of micropore diffusion perimeter diffusion

Table 1-3 The relationship between rate of water diffusion and pore area or perimeter
D of pore (mm) 2.64 0.95 0.35 Relative area 1.00 0.13 0.01 Relative Water loss perimeter (g) 1.00 0.36 0.13 2.655 0.928 0.364 Relative water loss 1.00 0.35 0.14

 Law of micro-pore diffusion: diffusion rate of

water vapor throughout poly micropore is not proportional to the area, but is proportional to the perimeter.

In the margin less chance of collision. Diffusion rate is larger in the margin than in the middle.

Diffusion by macropore

Diffusion by poly micropore

 4.2.3 Mechanism for stomatal opening and closing (p90-98) Opening in daytime and closure at night resulted from the swelling by water absorption or shrinking by water loss in guard cells. Stomatal complex Guard cell , subsidiary cell and substomatal space.

 Figure 1-15 Stomatal complex structure Guard cell

Dumbbell shape

subsidiary cell

Guard cell

Stomatal complex in dicot

Subsidiary cell Stomatal complex in monocot

Figure 1-16 The stomatal complex in dicot and monocot

Orientation of microfibrils allows the expansion of the cell only in the direction shown by the dashed arrows.

 (1) Starch-sugar conversion theory Starch phosphorylase (SPLase) plays an important role in stomata opening and closing.

pH>5.0 hydrolysis activity , pH<4.6 synthesis activity

 (1) Starch-sugar conversion theory

Light darkness

Photosynthesis in guard cell(GC) Respiration in GC Consume CO2Cell pH SPLase hydrolysis activity Starch becomes G-1-P Water potential GC absorbs water and turgor Stomata opening Produce CO2cell pH SPLase synthesis activity G-1-P to starch Water potential Loss of water and turgor Stomata closure

 (2) Potassium ion pump or inorganic ion uptake theory

Inner
0.10 pH5.19

pH5.56

pH5.60

Open
Guard cell

0.45 0.29 0 .1

Outer Subsidiary cell Close

Figure 1-17 the change in K+ and pH of guard cell and subsidiary cell during stomata opening and closing

pH5.78

Subsidiary cell
0.2
0.16

 (2) Potassium ion pump or inorganic ion uptake theory outer

Lightdark Guard cell(GC) photosynthesisrespiration H+ H+ ATP and malate K+ ATPase hydrolysis ATP, malate dissociates H+ H+ pump out of GCK+pump into GC Water potential GC absorbs water and turgor Stomatal opening
ATPase

inner ATP K+ + K K+

Figure 1-18 Stomata opens and closes

http:/Bio.fsu.edu/ ~outlaw/assorted/ k-salts.html

 4.2.4 Factors affecting stomatal opening and closure (1) light: form sugar and malate, acumulate K+ and Cl About 2.5% of full sun light Sensitive to blue light, UV-A receptor (blue light receptor) Lot of gene relevant to stamatal behavior

Positive

NPQ1

PLA2

PP1/PP2A

Light Negative
abi1-1 abi2-1

ABA ozone Ca2+ PP2B InsP3 InsP6

Actin filament Protein kinase/ CDPK

K+in channel

Fig 1 A simplified working model for proposed function of positive and negative regulators in light-induced stomatal opening. For simplicity, parallel signaling branches are not included here.

Stomatal opening

OA sensitive linolenic acid/ (blue light) arachidonic acid

Protein Kinase

14-3-3 protein fusicoccin

H+ pump

 (2) CO2: Low CO2 ,stomatal openinghigh CO2 , stomatal closure because of acidification and K+ leakage from guard cell. (3) Relative humidity in atmosphere: higher RH, larger opening. Low RH, loss of water of Guard cell. (4) temperature. In arrange of T, T rises and opening increases. Optimum 30 the opening become smaller at >35

 (5) leaf water and potassium contents The higher water and enough potassium, the opening larger. Too much water condition blocks stomatal opening (6) plant hormones ABA---close, ABA promotes Ca2+ increase in cytosolindirectly makes K+Cl- flow out of GC and inhabits entrance of K+into GC. IAA CTK result in stomatal opening

K+in channel
Ca2+ ADP+Pi

ABA
Ca2+permeable channel

H+
ATP Ca2+

Ca2+

S-type

AK+ pH
De po la

anion channel
R-type

Vacuole

ri z e

anion channel

K+out channel Fig 2 A guard cell model, illustrating the proposed functions of ion channels in ABA signaling and stomatal closing. The
right of the stomatal shows ion channels and regulators that mediate ABAinduced stomatal closing. The left cell shows the parallel effects of ABAinduced [Ca2+]cyt increases that inhibit stomatal opening mechanisms.

 4.3 Internal and environmental conditions affecting transpiration

Boundary layer

Substomatal space

eleaf-eair T

rleaf+rair

e=vapor pressure r=resistance

4.3.1 Effect of internal factors on transpiration Stomatal density (number/leaf cm2) Opening degree Leaf waterCO2 and ionspotassium contents ABA The areas of leaves or leaf cells; The transplanted plants are often cut the branches and leaves!

 4.3.2 Effect of environmental factors on transpiration (1) light lighttranspiration openingresistanceTleaf and Tair transpiration The difference of vapor pressure between in the leaf and air (2) Atmosphere relative humidity RH transpiration RH too lowstomatal closure transpiration

 (3) Air temperature In arrange T transpiration Too low or high transpiration (4) Wind Breeze transpiration the thickness of boundary layer (5) Air CO2 transpiration (6) Other factors which affect water absorption

 4.3.3
Relative transpiration(%)

Diurnal change of transpiration

120 100 80 60 40 20 0 4 8 12 Time of day 16 20

Dry and hot day

Clean day

4.4 Role and index of transpiration 4.4.1 Role (1) It decreases in leaf temperature; (2)It is a power for water absorption and transportation. (3) It enhances the transfer and distribution of mineral nutrition and other solutes in plant body.

 4.4.2 Index
(1) Transpiration rate ()Water loss of plant through transpiration per unit leaf area and per unit time (g/m2s) . daytime1.5-7.5 g/m2s night<0.3 g/m2s MeasurementWeight loss and gas exchange(GE).
Devices for GE Steady pore meterLI-1600

Photosynthetic systemLI-6400

 (2) Transpiration efficiency or transpiration ratio() Plant produces quantity (g) of dry mater when it consumes 1kg of water by transpiration. Wild types 1-8g/kg crops 2-10g/kg

Water utilization efficiencyWUE, Special definition Photosynthetic rate (CO2mol/m2s ) WUE= Transpiration rate (mmol H2O/m2s ) Intensive definition= Transpiration efficiency

 (3) Transpiration coefficient or water requirement ( Water requirement is a reciprocal of transpiration efficiency, means that plant consumes water quantity (g) for making 1g of dry matter. Wild types:125-1000 g crops:100-500 g .

 Section5 Water transport in plant 5.1 Pathway of water transport

soil root hairs root cortex parenchymaroot pericycle root vessels (tracheids) stem vessels (tracheids) potile vessels (tracheids)leaf vessels and tracheids mesophyll cells mesophyll cell space substomatal space stomataatmosphere
Fig Pathway of water transport in plant --- Soil-plant-atmosphere continuum

 5.1.1 Short distance transport

Transport form root hairs to root vessels. Largest resistance is in endodermis.
Casprian band blocks water transport through apoplast.

 Transport form terminal vessels(tracheids) of leaf to substomatal space

Leaf tracheids

stomata Air

 5.1.1 Long distance transport

Transport in root vessels (or tracheids) to leaf vessels (or tracheids) Vessels in angiosperm and tracheids in gymnosperm. Less resistance to water transport

 5.2 Power of water transport Root pressure in bottom, Transpiration pull in top

 Transpiration-cohesion-tension theory( )water can be transported in a continuous water column because water cohesion is larger than its tension.

5.3 Rate of water transport Symplast1mm/h. Xylem3-45m/h. Angiosperm:1-40m/h Gymnosperm <0.6m/h.

 Section6 Effective irrigation based on water physiology Maximum efficiency with the least water! 6.1 Law of plant water requirement 6.1.1 Plant types
Table 1-5 Water requirements for different crops
Crops Maize Sorghum Barley Rice Bean Potato Cotton Water requirements 370 322 520 680 700 640 570

 6.1.2 Growth stages

Relative water requirement

Seedling

Tillering or branching

Flowering and setting

Ripening

Growth stages

 Critical period of water (): a period during which plant is most sensitive to water deficiency and is most easily injured by this, but the water requirement is not always largest at that period. a period from pollen mother cell meiosis to pollen tetrad(4).
Two Critical periods of water for grain crops: Stem elongation from pollen mother cell meiosis to pollen tetrad and filling stage.

6.2 Index for effective irrigation 6.2.1 Morphological index (1)The tender stems and leaves wilt. (2)Stem and leaf appear in darkness or reddish. (3) Growth delay.

 6.2.2 Physiological index (1)Leaf relative water contents() A percentage of the actual water content to the water content of the leaf with water-saturated.
Leaf relative water contents (%)=
FW-DW

FW SFW-DW SFW

100%

FW=fresh weight of leaf, DW= dry weight of leaf , SFW=the water content of leaf with water-saturated

if Leaf relative water contents <80%, irrigation!

 (2)Diurnal change in leaf water potential

Water potential (MPa) normal -0.4 -0.8 -1.2 -1.6 Water deficient

Recovered in the eveningnot necessary to irrigate Not recovered in the dawn, necessary to irrigate

 6.2.3 Irrigation methods (1)ground irrigation. Alternative irrigation (2)sprinkling irrigation. (3)dropping irrigation.

 1. Cloze
(1)When the cell is bathed by a solution, water will enter the cell as it moves down the water potential gradient; When the cell is solution, which has more negative osmotic bathed by a potential than the cell, the protoplast will shrink away from the cell . wall. It is known as (2) Transpiration is defined as , by in the which water loss passes mainly through the epidermis of the leaves.

2.Question:
(1) What are water potential, osmotic potential and pressure potential? Describe the relations among them. (2) what is driving force for water movement in the xylem? (3) Why do we not irrigate plant with cold water at noon in the sunny summer day?