Course In-Charge: Dr. R.K.

Goswami Assistant Professor & Head of Crop Physiology

Presented by: SHISARENLA AIER M.Sc (agri) 5th year Redg. No- 2011-AMB-05 BNCA, AAU

INTRODUCTION:
Water absorbed by the root system of the plants, is

carried to the aerial green parts (green folia and stem, green fruits) of the plants and then lost to the atmosphere, either in the form of vapour or in the form of liquid. Out of the whole water absorbed, 97% is lost by different water loss mechanism by the plants. 3% is being utilized in different metabolic process of the plant.

Loss of water by the plants occurs by three main processes:

Transpiration

Guttation

Exudation/ Bleeding

What is Transpiration?
Loss of water in the form of water vapour by the aerial

parts of the plant(having stomata) is called transpiration.

 Transpiration occurs by day as well

Some important aspects of transpiration

as by night. 95% transpiration occurs by day(maximum) and 5% occurs by night. Transpiration phenomenon was first observed by Stephen Hales (1927). Transpiration replace body temperature of plants and causes cooling effect on the leaf surfaces. The amount of water loss varies with plants. Eg. In Egyptian cotton, it is 1 litre/day, in sunflower 2kg/day, in Birch 300400 Kg/day, in Elm it is a tonne/day

Types of transpiration (based on organs involved)

Stomatal transpiration(97-99%) through stomata.

Cuticular transriration( 1-3%) through cuticle.

Lenticular transpiration (o-1%) through lenticel.

Guttation:
Loss of water in the form

of liquid or water droplets through special pole, hydathode located in the leaf tip of plants is called guttation. Hydathodes may be active or passive. Active hydathodes are specialized epidermal cell which do not have a cuticle on the outside and are not connected directly with vascular strands internally. Passive hydathodes comparise loosely arranged, colourless and parenchymatous cells known as epithem. It lies inside the leaf below the vein end.

Exudation or bleeding
Gentle exudation of water sap from the cut(incised)

tissues of the plant.

Plants that exudes salts from its leaves

Exudation of reddish brown liquid through cracks from the stem of coconut

Peach fruit attacked with gummy exudation.

Transpiration ratio
It is the ratio of total water transpired to the amount of

dry matter accumulation by the same plant. Transpiration ratio = Amount of water transpired/ Total amount of dry matter accumulated Ex:-1. In millets 200-300 units 2. In alfa-alfa 700-1000 units 3. In pineapple 50 units The ratio is lower in desert plants. 4. In C4 plants it is 100-200 units of water for the manufacture of single unit of dry matter. 5. In mesophytes under normal condition it is 300500 units for unit dry matter accumulated.

ADVANTAGES OF TRANSPIRATION

Maintenance of turgidity. Removal of excess water. Cooling effect. Regulates rate of absorption of water. Formation of mechanical tissues. Regulates mineral salt absorption. Controls ascent of cell sap.

97% of water absorbed is being loss through the process of

transpiration. Excess transpiration results in wilting and under severe condition death of the plant. Deciduous trees shed their leaves in summer to minimize transpiration, i.e. minimization of transpiration occurs at the cost of photosynthesis.

FACTORS AFFECTING THE RATE OF TRANSPIRATION

1. 2. 3. 4. 5. 1. 2. 3. 4. 5.

EXTERNAL FACTORS Light (transpiration through stomates) Temperature (more temperature more transpiration) Relative humidity. Soil temperature. Air currents. INTERNAL FACTORS Shape, size, structure and orientation of leaves. Stomatal structure, size and number. Water status of the leaves. Root/ shoot ratio. Effect of various biotic and abiotic stress.

 Depends upon the time of opening of stomata. Stomata open during day time i.e. Photo-active movement and

MECHANISM OF STOMATAL MOVEMENT

scoto-active movement open at night. Majority of plants show photo-active movement and only few plants like CAM plants show scoto-active movement. When there is more water, the guard cells become turgid and stomata open and when the amount of water decreases the guard cells become flaccid and stomata close. Several theories were proposed by different plant physiologists to explain the mechanism of stomatal movement.

Two such theories are explained below 1. Steward's Starch hydrolysis theory:
Lloyd(1908), Sayre (1926) & Scarth (1932) explained the opening and

closing of stomata through the starch-sugar hypothesis. The guard cells store starch molecules during night and this starch is converted into sugar during day time. The interconversion of starch to sugar is catalyzed by an enzyme called starch phosphorylase secreted by the chloroplasts of guard cells.

This theory is based on the following experimental conclusions. The starch content of the guard cells is high during night and low during day time. The high pH in guard cells favours the stomata to open and the low pH its closure.

Steward (1964) modified this hypothesis and proposed the starch hydrolysis theory and outline that as given below.
ACTIVITIES AT DAY TIME Starch converts to glucose -1-phosphate Glucose-6- phosphate dissociates Glucose and inorganic phosphate dissolved in water ACTIVITIES AT NIGHT TIME Glucose + ATP Hexokinase Glucose-1-phosphate pH5 phosphorylase Conversion of glucose into polysaccharide (starch) leads to dilution of cell sap

Increased concentration of solute in guard Decreased concentration of solutes in cell (i.e. OP & DPD increases) guard cell(i.e. DPD of cell sap decreases) Influx of water in guard cells Increased turgor pressure in guard cell Stomata open Efflux of water in guard cells Decreased turgor pressure in guard cell Stomata closed

The biochemical reactions involved in the opening and closing of stomata

Objections of classical theory: 1. Starch # Sugar interconversion is a slow process which cannot account for rapid stomatal movement. 2. Starch molecules are not observed in onion plant where stomatal movement occurs. 3. Glucose is not detectable in the guard cells of all plants when stomatal opening occurs. 4. The theory could not explain the effectiveness of blue - light in photosynthesis at the time of stomatal opening.

2. K+ pump theory: Proposed by Levitt (1974). The theory is also called Proton transport theory or Phytohormone theory or Ion-flux theory. The theory is based on the efflux of H+ from the guard cells into the subsidiary cells and influx of K+ from the subsidiary cells into the guard cells and describe the maintenance of turgidity in cells. Stomatal opening is an energy requiring active process whereas stomatal closure is regulated by certain hormones.

Conti

 A brief account of this theory is given below.

Conti.

DURING DAY (IN LIGHT) Accumulation of K+ ions in guard cells Increased concentration of the solute Water potential decreases Endosmosis of water

DURING NIGHT (IN DARK) Loss of K+ ions in guard cells Decreased concentration of the solute Water potential increases Exosmosis of water

Increased turgor pressure in guard cells

Stomata open

Decreased turgor pressure in guard cells

Stomata close

Opening of stomata Light > starch ->Production of malic acid in guard cells -> Dissociation into hydrogen and malate ions -> Influx of K+ and efflux of H+ -> Formation of potassium malate ->Transport of potassium malate into the cell sap -> Endosmosis of water into the guard cells -> Increase of turgor pressure -> Stomata open. Closing of stomata During closing of stomata all the above processes are reversed. Dark -> Potassium malate transported into cytoplasm -> Dissociate into K+ and malate > Influx of H+ and efflux of K+ -> Malate combine with H+ to form malic acid -> converted into starch -> Exosmosis of water from the guard cells -> Decrease of turgor pressure -> Stomatal closure

Mid day closure of stomata:

Stomates may close even by day when the environmental temperature exceeds normal at noon. This phenomenon of closure of stomates at noon is termed as mid day stomatal closure and Transient phenomenon. This is a protective mechanism of some plants against excess water loss.

Any substance which is being used to check or control the rate of transpiration is called anti-transpirant. Practical utility of anti-transpirants : 1. To check the water loss from the crops particularly during water deficit condition. 2. To increase the water use efficiency (WUE). Characteristic of anti-transpirants: 1. Non-toxic, more durability. 2. It should not disturb the normal gas exchange phenomenon. 3. Easy in application. 4. Availability in market. 5. Reasonable cost. Examples of anti-transpirants : 1. Colourless plastics , silicon oil, low viscosity waxes act as film forming and they reduce cuticular transpiration. 2. Natural antitranspirants are abscisic acid, Phenolic acid which act as stomatal inhibitors.

CONCLUSION
The main requirement of water within the plant is to keep the

protoplasm in its correct physical state. Water is also very essential to keep the cells turgid which is necessary condition for growth. Deficiency of water causes wilting and permanent wilting consequently leading to death. Water acts as a medium for movement of various types of gases, minerals, organic compounds etc. in xylem and phloem inside the plant body. It takes part in many important metabolic activities like reduction of CO2 in photosynthesis through reduced coenzyme II (NADPH2) All hydrolyzing enzyme needs water for the completion of enzymatic action. Water helps in break down of starch during respiration. A uniform temperature in the plant tissues is also maintained by water.

References
Bora, D. & Rajkhowa, R.C.2010.Water relation of plant

life. A Text Book of Higher Secondary Biology. Assam book depot, 141-149. Sinha, R.K.2007.Loss of water from plants. Modern Plant Physiology. Narosa Publishing house, 84-97.