Translocation of Water

(BOT-202)
 Ascent of SAP

• The water after being absorbed by the roots is distributed to all parts of the plant
(excess of which is lost through transpiration). In order to reach the topmost parts
of the plant, the water has to move upward through the stem. This upward
movement of water is called as Ascent of Sap.
• Ascent of sap can be studied under the following two headings

(A) Path of Ascent of Sap

(B) Mechanism of Ascent of Sap.
 Path of Ascent of Sap

• It is well established that the ascent of sap takes place through xylem.
• Its cleared from the following experiments:

I. A leafy twig of balsam plant is cut under water (to avoid entry of air-bubbles
through the cut end) and placed in a beaker containing water with some eosin (a
dye) dissolved in it. After sometime coloured lines will be seen moving upward in
the stem. If sections of stem are cut at this time, only the xylem elements will
appear to be filled with coloured water.
(ii) Ringing Experiment

A leafy twig from a tree is cut under water and placed in a beaker filled with water.
A ring of bark (all the tissues outer to vascular cambium) is re­moved from the stem.
After sometime it is observed that the leaves above the ringed part of the stem
remain fresh and green. It is because wa­ter is being continuously supplied to the
upper part of the twig through xylem.
 Xylem Sap

• The fluid translocated in xylem is dilute (0.1%) solution of mineral ions absorbed
from the soil.

• These dissolved mineral ions are carried passively in water in the transpiration
stream and when these reach leaf, they are absorbed by leaf cells by active
transport mechanism.

• In addition to mineral ions, small amount of organic substances, amides, sugars,

cytokinin, abscisic acid etc. are also present.

• This water solution translocated in the xylem is referred to as xylem sap.

 Transpiration and Ascent of Sap

• Translocation of xylem sap is quite rapid during daytime especially when

transpiration rates are high.

• Lowest rates are found in evergreen conifers, intermediate rats in deciduous trees
and herbaceous plants and highest in vines and lianas.

• Ascent of sap in transpiring plants is initiated by transpiration pull.

• The liquid water filling tracheid and vessels of xylem shows bulk flow.
 Mechanism of Ascent of Sap

• In small trees and herbaceous plants the ascent of sap can be explained easily

• In tall trees like Australian Eucalyptus, some conifers such as mighty Sequoias e.g.
Sequoia sempervirens (California redwood) is the tallest tree of the world reaching a
height of 111.6 m.
• Where the water has to rise up to the height of several hundred feet, the ascent of sap,
in fact, becomes a problem.

• Although the mechanism of ascent of sap is not well understood, a number of theories
have been put forward to explain it.
 Theories of Ascent of SAP

• (A) Vital Theories:

• Supporters of vital theories think that the ascent of sap is under the control of vital activi­ties in the
stem.
• Two such theories are common but they are not very convincing:

• Many physiologists suggested that the living cells of stem especially xylem parenchyma cells are
responsible for upward translocation of water. They were of the view that living cells act as
pumps and tracheid and vessels act as water reservoir. Experimental evidence does not support
these theories and no such pumping cells were observed in plats when anatomy of plants was
studied in detail.

• Eduard A. Strausurger cut a trunk of 20 m tall tree and suspended in a bucket containing copper
sulphate, picric acid and other poisons. He found that the fluid moved upward killing all the living
cells (bark and ray cells). The movement continued until the leaves were killed and transpiration
ceased.
• (B) Root Pressure Theory:
• Although, root pressure which is developed in the xylem of the roots can raise water to a
certain height but it does not seem to be an effective force in ascent of sap due to the
following reasons:

• (i) Magnitude of root pressure is very low (about 2 atms).

• (ii) Even in the absence of root pressure, absent of sap continues. For example, when a
leafy twig is cut under water and placed in a beaker full of water it remains fresh and
green for sufficient long time.

• (iii) In gymnosperms root pressure has rarely been observed.

(111) Atmospheric Pressure:

• Some physiologists are of the view that atmospheric pressure is responsible for upward
translocation of water, but to raise water from the ground level to the top of tallest trees (116.6
m), a pressure of 10.63 atm would be required. At the same time same amount of pressure
would be needed to overcome the resistance offered by the pathway of water, thus a total of
about 22 atm is required.
• The plants are rooted in the soil and for operation of atmospheric pressure an open surface at
the base is required.
• (i) water cannot be lifted more than 34 ft even when the suction pump is used. Thus clearly
water is not pushed to the top of tall trees by atmospheric pressure.
• (iv) Capillarity:

• Most physiologists are of the view that capillarity is responsible for rise of water to
the top of trees.
• Capillarity causes the rise of liquids in small tubes. It occurs because the liquid wets
the sides of the tube by adhesion and is pulled up, which is evident from the curved
meniscus at the top of liquid column.
• Xylem vessels are placed one above the other forming a sort of continuous channel
which can be compared with long capillary tubes and it was thought that as water
rises in capillary tube due to capillary force, in the same manner ascent of sap takes
place in xylem vessels.
• There are many objections to this theory:

• (i) Water in capillary tube rise because of the open meniscus at the top of water
surface, but the xylem cells in plants do not have open meniscus

• (ii) The magnitude of capillary force is low.

• (iii) In Gymnosperms usually the vessels are absent. Other xylem elements do not
form continuous channels.
• (D) Transpiration Pull and Cohesion Theory:

• This theory was originally proposed by Dixon and Joly (1894). This theory is very
convincing and has now been widely supported by many work­ers.

It is based on the following features:

• Cohesive and Adhesive properties of water mol­ecules to form a continuous water

column in the xylem.
• Transpiration pull exerted on this water col­umn.
• Water molecules remain joined to each other due H-bonds between water
molecules, to the presence of H-bonds between them.
• Although H-bond is very weak but when they are present in enormous
numbers as in case of water, a very strong mutual force of attraction or
cohesive force develops between water molecules and hence they remain in
the form of a continuous water column in the xylem. The magnitude of this
force is very high (some­times up to 350 atmos.), therefore the continuous
water column in the xylem cannot be broken easily due to the force of gravity
or other obstructions offered by the internal tis­sues in the upward movement
of water.
• The adhesive properties of water i.e. the attraction between the water
molecules and the container’s walls (here the walls of xylem) further ensure
the continuity of water col­umn in xylem.
• When transpiration takes place in leaves at the upper parts of the plant, water evapo­
rates from the intercellular spaces of the leaves to the outer atmosphere through the
sto­mata. More water is released into the intercellular spaces from the mesophyll cells.
In turn, the mesophyll cells draw water from the xylem of the leaf.

• Due to all this, a tension is created in water in the xylem elements of the leaves. This
tension is transmitted down­ward to water in xylem elements of the roots through the
xylem of petiole and stem and the water is pulled upward in the form of continuous
unbroken water column to reach the transpiring surfaces—up to the top of the plants.
• According to some workers, the main objection against this theory is that certain
air bubbles present in the conducting channels will break the continuity of the
water column. This is known as cavitation.

• This has been counteracted by others who say that there are no air bubbles and if
at all are present, they will not break the water column which will remain
continuous through other elements of the xylem