Water potential •The potential energy of water is known as water potential. •The water potential is represented by Greek letter ‘Ψ’ (psi). •At normal temperature and pressure, the water potential of pure water is zero. •When solutes are added, the value of water potential decreases. Owing to this the water potential in a solution or in a plant cell has negative value. •The direction of cell to cell movement of water depends upon the difference in water potential. •The movement (flow) of water occurs from a less negative water potential to a more negative water potential. •Some solutes, when dissolved in pure water reduce the water potential. The magnitude of lowering of the water potential due to dissolution of solute is called solute potential (Ψs). •Solute potential is always negative. Permeability •The ability of a membrane to allow the passage of substances through it is called permeability. •There are four types of membranes, viz. semipermeable, impermeable, freely permeable and differentially permeable, depending upon their nature permeability. •The membrane which allows only the solvent to pass through it but not the solute is called semipermeable eg parchment paper. •The membrane which neither allows the solvent nor the solute to pass through it is called impermeable eg suberised wallsof cork cells. •The membrane which allows the passage of certain solutes but not all along with the solvent is called differentially permeable eg cell membrane, tonoplast etc •The membrane that allows the passage of both solutes and solvent is called freely permeable eg cell wall. Mechanism of water absorption: There are two methods of water absorption, viz. active absorption and passive absorption. (1) Active absorption: The absorption of water which involves the expenditure of metabolic energy is called active absorption of water. (2) Passive absorption: The absorption of water in which the living cells of root do not play any role is called passive absorption of water. In passive absorption there is no expenditure of metabolic energy. Path of water in root tissue during absorption The movement of water from the soil to the xylem of the root occurs through the epidermis (epiblema), cortex, endodermis and pericycle along two different pathways, viz. apoplast pathway and symplast pathway. (1) Apoplast pathway:- Apoplast movement of water in plants occurs exclusively through cell walls and intercellular spaces. (2) Symplast pathway:- In symplast movement of water, the water moves from one cell to other through cytoplasmic bridges called plasmodesmata.
The plasmodesmata interconnect the cytoplasm of cells
forming cytoplasmic network called symplast. Since the water moves through symplast, it is called symplast pathway. Ascent of sap •The upward movement of water (sap) from roots to aerial parts against the gravitational force is called ascent of sap. •Ascent of sap in vascular plants occurs through the xylem elements like tracheids and vessels. •Many theories have been put forth to explain the ascent of sap in plants •Cohesion- tension theory is the only theory which explains the ascent of sap in plants in the most satisfactory way. Objections/ limitations of root pressure theory: Although, ascent of sap takes place due to root pressure, there are certain objections raised, such as - i. It is not applicable to plants taller than 20 meters. ii. Ascent of sap can also occur even in the absence of root system. iii. Root pressure value is almost nearly zero in taller gymnosperm trees. iv. In actively transpiring plants, no root pressure is developed. v. Xylem sap under normal condition is under tension i.e. it shows negative hydrostatic pressure or high osmotic pressure. To sum up therefore, root pressure is not the sole mechanism explaining the ascent of sap in all plants of varying heights •Cohesion-tension theory was put forth by Dixon and Jolly (1894). •Cohesion-tension theory is based on four principles, viz.cohesion of water, adhesion of water, continuity of water column and transpiration pull. •According to the cohesion tension theory, transpiration pull augmented with cohesion and adhesion of water is responsible for ascent of sap Objections/Limitations of capillarity theory: Few important objections are: i. Capillary tube (xylem) must be continously and completely hollow from one end to the other end but tracheids in the xylem show closed end-walls. ii. The lower end of capillary tube i.e. xylem must be in direct contact with soil water. However, there exists a barrier of root cortex between xylem and soil water. iii. Narrower the capillary tube, greater is the height to which water column is raised. Thus, taller trees should show xylem vessels with very narrow bore (diameter). However, in nature the tall trees show xylem vessels having wider bore. Hence, to sum up capillarity can not be the sole mechanism to explain ascent of sap in all the plants of varying heights. Guttation: •The exudation of water in liquid form through intact plant parts is called guttation. •Guttation occurs when transpiration is low and absorption of water is more. •Guttation takes place through special glands called hydathodes. Transpiration pull: •The transpiration pull developed in the leaf vessel is transmitted down to root and thus accounts for the ascent of sap. •Excess water is lost in the form of vapour, mainly through the stomata found on leaf. •This water loss increases D. P. D. of mesophyll cells. •These cells withdraw water ultimately from xylem in the leaf. In otherwords, due to continous transpiration, a gradient of suction pressure (i.e. D. P. D.) is developed right from guard cells up to the xylem in the leaf. •This will create a tension (called negative pull or transpiration pull) in the xylem. Consequently, water column is pulled out of xylem. Thus, water is pulled upwards passively against the gravity leading to the ascent of sap. Objections/Limitations of transpiration pull theory: i. For transpiration pull to operate, water column should be unbroken and continous. However, due to temperature fluctuations during day and night, gas bubbles may enter in water column breaking the continuity. ii. This mechanism assumes that tracheids are more efficient than the vessels, as their end walls support water column. However, vessels are more evolved than tracheids and are more efficient.