TRANSPIRATION

A small fraction (less than 5%) of the total water is used up by the plant and the remaining amount is lost in the environment in vapor form or liquid form. The loss of water in the vapour form from aerial organ of living plant is known as transpiration where as the loss of water in the liquid (droplet) form from aerial uninjured organ of living plant is known as Guttation. Rate of transpiration can be measure by Potometers while total pore area (stoma) measure by Porometer. Psychrometer is used to measure relative humidity and rate of transpiration. The loss of water is so great that it reduces water level in the soil and can lead to the death of plant, but transpiration is said to be necessary for water and mineral absorption, ascent of sap and lowering the temperature (cooling effect). So, transpiration is called as necessary evil (Curtis) or unavoidable evil (Steward).

Types of transpiration: Bases on the plant parts or structure involved, following types can be recognized. (i) Stomatal transpiration (ii) Cuticular transpiration (iii)Bark transpiration (iv) Lenticular transpiration.

(i) Stomatal transpiration/foliar transpiration: Surface of leaves possess many

minute pores called stomata, water vapour diffuse out of these stomata in to the atmosphere. It is called stomatal transpiration. It is the major form of transpiration, because 50-97% water is lost by stomatal transpiration. It however occurs only when stomatas are open. For performing stomatal transpiration water absorbed by root hairs reaches the xylem vessels & tracheids through the root cortex & then to xylem vessels & tracheids of the leaf due to which turgor pressure of its cells becomes more than of mesophyll cells. The intercellular spaces in mesophyll cells are filled with air. By transpiration water vapour enters the intercellular space and then passes on into the atmosphere through stomata.

(ii) Cuticular transpiration: - It is the loss of the water in the vapour form from the
general surface (leaves & young stem) through the layer of cuticle, but the quantity of loss will depend up on the thickness of the cuticle, means thicker the cuticle less the water loss MAMS ACADEMY FOR CAREER EXCELLENCE Page 1

TRANSPIRATION
(as in xerophytes) and thinner the cuticle more the water loss as in hydrophytes. Cuticular transpiration take place continues through out day & night however it is approximately 3 to 10% of total water loss but in herbs and fern it may be 50%.

(iii)Bark transpiration: - This occurs through the bark of woody stem. It is

approximately 1% of total lost.

(iv) Lenticular transpiration: -This is loss of water in vapour form from lenticels or
aerating pores present in the bark of stems or fruits etc. Through lenticels some water is lost continuously, however it is approximately 0.1 % of the total water lost.

Structure of stomata: - are minute pore complexes which occur on the soft aerial parts
of the plants, especially on the leaves. Each stomata is surrounded by two kidney shaped (dicots) or dumbbell shape (monocot as Gramineae and cypraceae) epidermal cells called guard cells. These cells are living having nucleus, chloroplast, and cytoplasm. Kidney shape guard cells have outer wall thinner and more elastic while the inner-wall thicker and less elastic. Dumb-bell shape guard cells have thin walled ends and thick walled middle region. These guard cells are surrounded by some specialized cells called subsidiary cells or accessory cells which support in the movement of guard cells. The size, shape, number and position of stomata and guard cells vary from plant to plant. When fully open, the stomatal pore measure 3-10 in width and 10-40 in length.

* Scotoactive stomata: - are special types of stomata which open in dark & close
during the day time so they prevent excess loss of water during transpiration. They occur in succulents xerophytic plants. E.g., Cactus, Bryophyllum, Pineapple etc. (Generally stomata are photoactive).

Number and distribution of stomata: -In most plants there are 50300 stomata
per sq. mm. or 1000-60000/sq. cm. of leaf area are present. (The least number is reported is 14 & max. no. 1038/sq.mm.). The number of stomata usually not equal on both surfaces of leaf and varies plant to plant. Xerophytes possess larger number of stomata than mesophytes, and higher in trees and shrubs than grasses. Stomata occupy 1-2% of total leaf area. According to Salisbury there is a co-relation between number of stomata and number of epidermal cells per unit area which is called stomatal index (I). I= S x 100 where S=no. of stomata /unit area. S+E E=no. of epidermal cells/unit area. Plants are divided into the following categories on the basis of the no. of stomata and its distribution-

MAMS ACADEMY FOR CAREER EXCELLENCE

Page 2

TRANSPIRATION
1. Apple type (mulberry type): -These plants have stomata only on the lower
surface of leaves. hypostomatic type. E.g., apple, peach, mulberry, walnut etc. such leaves are

2. Potato type: - have stomata on the both surfaces of leaf but more stomata on
the lower surface (multistomatic) and less stomata on upper surface of leaf (pauci stomatic), such leaves are amphi stomatic and aniso stomatic type. E.g., Potato, tomato, pea, cabbage, bean and many other dicot plants.

3. Oat type: - Plants have equal number of stomata on the both surfaces of the
leaf. E.g.,. oat, wheat, maize, grasses and many other monocots. Such leaves are amphi stomatic and iso stomatic type.

4. Water lily type: - Have stomata on the upper surface of leaves only because lower
surfaces of leaves make contact with water surface. E.g.,. Water lily, nymphaea & most floating plants. Such leaves are epistomatic type.

5. Potamogeton type: - stomatas are absent or vestigial because the entire body of
plant dip in water so gaseous exchange take place through general body surfaces. E.g., Potamogeton, hydrila, valisnaria and other submersed plants. Such leaves are called astomatic. Loftfield has classified stomata into following main groups on the basis of their daily opening and closing movement-

(i)Alfa alfa type- the stomata remain opens through out the day and closed all night.
E.g., pea, bean, mustard, radish, turnip, apple, grapes etc.

(ii)Potato type- the stomata close for a few hours in the evening. E.g., Onion, potato,
banana, cabbage etc.

(iii)Barley type- the stomata open only for a few hours in a day. E.g., Barley, maize,
wheat and other cereals.

(iv)Equisetum type- stomata remain always open. Mechanism of opening & closing of stomata: - It depends upon the turgid or
flaccid state of the guard cells. When guard cells are in turgid state the stomatal aperture become open and when guard cells are in flaccid state the stomatal aperture become closes. The inner wall of guard cell is thick while outer wall is thin. When the turgor pressure of the guard cells is increased the outer thinner wall of the guard cell is pushed out due to which a tension is created on the inner thicker wall thus pulling the inner thicker wall towards periphery thus leading to the opening of stomatal aperture & water vapours go outside. When the guard cells are in a flaccid state the outer thinner wall of guard cells returns to original position or move towards pore, due to which tension on the inner thick wall is released which also returns to its original position and stomatal aperture gets closed again. MAMS ACADEMY FOR CAREER EXCELLENCE Page 3

TRANSPIRATION

Theories explaining opening and closing of stomata: (i)Active K+ ion uptake theory (Levit) or Importance of K+ in opening & closing of stomata: -Acc to Imamura & Fujino (1967) When a leaf is exposed to
sunlight, pH of guard cells rises due to assimilation of CO2 in photosynthesis and uptake of H+ ions by chloroplast and mitochondria from cytoplasm. At this higher pH starch contents of the guard cells are converted to PEP. PEP combine with CO2 with the help of PEPcase and form OAA which get changed into malic acid which dissociate in to malate anion & H+. In exchange for H+, K+ are taken into the guard cells, some Cl- ions are also taken into guard cells to neutralize a part of k+. As H+ ions leave the guard cells (efflux), K+ions enter the guard cells (influx). These K+ ions react with malate anions and form potassium malates which migrate into the vacuole of guard cells. Due to presence of potassium malate in the guard cells, the water potential with in the guard cells become decreased so water from adjoining epidermal cells enters in to guard cells as a result of which their turgor pressure is increased & the stomata open. During stomatal closing, reversal of H+ & K+ pump occurs where by the K+ ions lost from the guard cells in to the surrounding cells resulting in increased water potential within the guard cells so water comes out due to exosmosis & the guard cells become flaccid there by promoting the closure of stomata. Stomatal opening is an active process where as closing is a passive process.

In succulent plants during night there is incomplete oxidation of carbohydrate

(respiration) and accumulation of organic acids, the CO2 is therefore not released and as a result stomata remain open in dark. During night(dark) 2C6 H12 O6 + 3O2 3C4 H6 O5 + 3H2 O Malic acid Where as during day time, the accumulated organic acids break down rapidly releasing excess amount of CO2, sufficient for photosynthesis as well as to keep the stomata closed. During day time (light) C4 H 6 O 5 Malic acid + 3O2 4 CO2 + 3 H2 O

Light formation of malic acid from starch in guard cells dissociation of malic acid in to malate anions & H+ ions influx of K+ & efflux of H+ formation of potassium malate and enters into the vacuoles water potential decrease in to the guard cell resultant endosmosis of water into guard cells increase of T.P. in guard cells Stomata opens.

Photosynthetic theory (Von Mohl and Schwendener): -Theory proposes that in

the morning as soon as light is available chloroplast of guard cell start photosynthesis as a result sugars are produced, which increase the osmotic concentration of guard cells, water MAMS ACADEMY FOR CAREER EXCELLENCE Page 4

TRANSPIRATION
comes in, guard cells become turgid and stomata are open. This theory is not accepted because photosynthetic activity of guard cell chloroplast seems to be negligeable and sugar does not occur in detectable quantity in guard cells.

Starch Sugar Interconversion hypothesis (Classical theory): - The theory

was proposed by Sayre and Scarth and was modified by Steward on the basis of activity of phosphorylase enzyme. Starch + iP
pH=7 Phosphorylase enzyme

Glucose -1-phosphate

According to this theory guard cells contain starch and phosphorylase enzyme, which causes conversion of starch into glucose at higher pH which is developed by utilization of CO2 in photosynthesis. Glucose increases osmotic concentration of guard cells, raising the turgor pressure and causing opening of stomata. In the night, CO2 accumulate in the cell and intercellular spaces, thus lowering the pH at which phosphorylase causes conversion of glucose to starch. Osmotic concentration of guard cells decreases water moves out therefore, stomata are closed. This theory is not accepted because in some families starch is absent as in onion. Glucose does not appear in detectable quantity in guard cell of open stomata rather malic acid accumulates.

Transpiration ratio/water requirement/efficiency of transpiration: -It is the

amount of water transpired by a plant for the synthesis of a unit dry matter. The value gives an idea of the requirement of water by crops, shrubs and trees. The ratio varies from one plant species to another and from one condition to another. Desert plants have highest transpiration ratio.

Guttation (term by Burgerstein): -Loss of water in the liquid state from uninjured parts of
plant is known as Guttation. It usually occurs from tips & margins of leaves during night or early morning when there is high atmospheric humidity as during wet seasons. Guttation occurs in some plant only (345 genera). E.g.,. Amorphophallus (max.), Oat, Garden Nasturtium, Saxifraga, Cucurbits, potato, tomato, colocasia & many grasses etc. In the regions of Guttation, the leaves possess special pores called hydathodes. Hydathodes/water pores are permanently open pores as their guard cells are immobile. Hydathodes are present at the tips of veins in leaves. A hydathode consists of a pore in the epidermis followed by large intercellular spaces and loosely arranged parenchyma called epithem and blindly ending xylem elements. Root pressure is probably the main cause of Guttation. Gutteted water contains minute quantities of both inorganic & organic salts and is not pure. Guttation is observed frequently during warm-humid night but normally it occur maximum during day and minimum during night. In moist and humid conditions, the rate of absorption of water greatly exceeds than transpiration.

Factors effecting transpiration: - are of two types

(i) External factors (ii) Internal factors

(i) External Factors: -are of followings

MAMS ACADEMY FOR CAREER EXCELLENCE Page 5

TRANSPIRATION
1. Light: - light is very important factor of transpiration. In most plants stomata open during the day. Blue light is most effective in causing stomatal opening followed by Red light. 2. Relative humidity (R.H.) of atmosphere: - R.H. is the ratio of actual amount of water vapour present in the atmosphere and the amount of water vapour required to saturate it completely. If R.H. of a place is low then the transpiration rate become high and if R.H. is high then the rate of transpiration become low. R.H. decreases with increasing temperature and increases with decreasing temperature. Psychrometer is used to measure relative humidity and rate of transpiration. 3.Wind: - If wind velocity is high the rate of transpiration become also high because wind removes humid air from around the leaves. 4. Temperature: -temperature directly effects the transpiration. Transpiration increases with increasing temperature. 5. Available water: - If available water in the soil is low the transpiration is also low. With the deficiency of available water leaves wilt & stomata are closed thus decreasing the rate of transpiration. 6. CO2 concentration: - increasing the CO2 concentration around the leaves should lead to wide opening of stomata but instead partial closure occurs. 7. Sayre (1926) observed that stomata remain open in neutral or alkaline pH (in the atmosphere of ammonia vapors) even in dark and closed in acidic pH (in the atmosphere of acetic acid vapors) even in light.

(ii) Internal factors: Structure of leaves: -Transpiration will decrease if on the leaves thick cuticle, waxy coating, sunken stomata, covering of dead hairs etc. present. These adaptations are found in xerophytes.

Significance of transpiration: -Transpiration is an unavoidable evil or necessary evil

(Curtis-1926), because it often produces water deficit in plants which check photosynthesis, reduce growth and if too severe may cause death from desiccation. It is probable that plant are injured & killed by excessive transpiration than by any other cause. Transpiration is considered necessary for the plants because of its supposed role in cooling the plant, in upward conduction of water, in absorption & transported minerals.

Plant anti-transpirants: These materials applied to plants for retarding transpiration. E.g.,. Colourless plastics, waxes, silicon oil, aspirin, abscisic acid and a fungicide phenyl mercuric acetate.

MAMS ACADEMY FOR CAREER EXCELLENCE

Page 6

TRANSPIRATION

MAMS ACADEMY FOR CAREER EXCELLENCE

Page 7