Awais Ahmad Length of roots = 12 kilometre Surface area = 5 sq metres Length of roots + root hairs = 220 kilometre Surface area = 14 sq metres Degree of soil contact = 1% Maximum distance for H2O to move to a root = 10 millimetres There are two ways for water to enter into the plant roots Active Transport Passive Transport Active Transport: Water is absorbed due to activities going on in roots. Absorption of water occurs with the help of energy in the form of ATP. Absorption takes place against concentration gradient - even when the concentration of cell sap is lower than that of soil water. Passive Transport: Passive absorption is by osmosis. Passive absorption takes place along the concentration gradient - when the concentration of cell sap is higher than that of soil water. Water is absorbed when transpiration rate is high or soil is dry. Due to high transpiration rate, water deficit is created in transpiring cells. Rapid transpiration removes water and reduces turgor pressure in living cells of root. The suction force thus developed is transmitted to root xylem. It pulls water from surrounding root cells to make up water deficit. Diffusion: Diffusion is the movement of molecules from a region of high concentration to a region of low concentration by means of random molecular motion. Diffusion requires kinetic energy from the environment but does not require cellular energy. Hence diffusion is a form of passive transport. Osmosis: Omosis is diffusion of water across a semipermeable membrane. Again Osmosis like diffusion in general does not require any cellular energy but just the kinetic energy related to the heat on either side of the membrane. Hence its also a passive transport. Aquaporins: Are transport proteins in the cell membrane that allow the passage of water. Most of the water absorbed by plants comes in through root hairs -Collectively provide enormous surface area -Almost always turgid because their water potential is greater than that of soil 10 An expenditure of energy is required for ions to accumulate in root cells -Once in the roots, the ions are transported via the xylem throughout the plant Surface area for water and mineral absorption is further increased by mycorrhizal fungi -Particularly helpful in phosphorus uptake 11 Three transport routes exist through cells -Apoplast route = Movement through the cell walls and the space between cells -Symplast route = A cytoplasm continuum between cells connected by plasmodesmata -Transmembrane route = Membrane transport between cells and across the membranes of vacuoles within cells -Permits the greatest control 12 Cont… Apoplast route Symplast route Transmembrane route Plasma membrane Cell wall Plasmodesma Vacuole 13 Eventually on their journey inward, molecules reach the endodermis -Any further passage through the cell walls is blocked by the Casparian strips -Molecules must pass through the cell membranes and protoplasts of the endodermal cells to reach the xylem 14 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. H2O and minerals H2O and minerals Endodermis apoplastic route symplastic route Phloem Xylem Casparian strip Cell membrane H2O and minerals H2O and minerals Endodermal cell 15 Lateral transport of water in roots Casparian strip Endodermal cell Pathway along apoplast Pathway through symplast 1 Uptake of soil solution by the hydrophilic walls of root hairs provides access to the apoplast. Water and minerals can then soak into the cortex along this matrix of walls. Casparian strip 2 Minerals and water that cross the plasma membranes of root hairs enter the symplast. 3 As soil solution moves along the apoplast, some water and minerals are transported into the protoplasts of cells of the epidermis and cortex and then move inward via the symplast. 1 Plasma membrane Apoplastic route Vessels (xylem) 2 Symplastic route Root hair Epidermis 4 Within the transverse and radial walls of each endodermal cell is the Casparian strip, a belt of waxy material (purple band) that blocks the passage of water and dissolved minerals. Only minerals already in the symplast or entering that pathway by crossing the plasma membrane of an endodermal cell can detour around the Casparian strip and pass into the vascular cylinder. Cortex EndodermisVascular cylinder 5 Endodermal cells and also parenchyma cells within the vascular cylinder discharge water and minerals into th walls (apoplast). The xylem vessels transport the wate and minerals upward into the shoot system. Water first enters the roots and then moves to the xylem, the innermost vascular tissue. Water rises through the xylem because of a combination of factors. There are to types of movement on the basis of distance covered. Short-distance movement: Movement of water at the cellular level plays a major role in bulk water transport. Water can diffuse through cell membranes. (Herbaceous) Long-distance movement: Water movement in most of the vascular plants from roots to leaves. (Sometimes more than 100m) Potentials are a way to represent free energy Water potential (yw) is used to predict which way water will move -Measured in units of pressure called megapascals (MPa) 19 Diffusion of water across a semi-permeable membrane is termed osmosis If a plant cell is placed in a solution with high water potential (low osmotic concentration) -It will become swollen or turgid If a plant cell is placed in a solution with low water potential (high osmotic concentration) -It will exhibit shrinkage or plasmolysis 20 Pressure potential (yp): Turgor pressure against the cell wall -As turgor pressure increases, yp increases Solute potential (ys): Pressure arising from presence of solute in a solution -As solute concentration increases, ys decreases (< 0 MPa) The total potential energy of water in the cell yw = yp + ys 21 When a cell is placed in pure water, water moves into the cell because the water potential of the cell is relatively negative When a cell is placed in a solution with a different ys, water moves in the direction that eventually result in equilibrium -Both cell and solution have the same yw 22 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Pressure Potential p Turgor pressure p=-0.5MPa Wall pressure – Cell wall + Cell membrane Pure water a. Solute Potential s s=-0.2MPa s=–0.7MPa Sucrose molecules b. Water Potential water movement = s + p cell = –0.7 MPa + 0.5 MPa = –0.2 MPa solution = –0.2 MPa (solution has no pressure potential) c. 23 Cell Initially Introduced into Solution Solution ys = –0.7 MPa yp = 0 MPa ysolution = –0.7 MPa Cell wall Cell membrane Cell ys = –0.2 MPa yp = 0.5 MPa ycell = 0.3 MPa a. b. Cell at Equilibrium Is Plasmolyzed ycell = ysolution = 0.7 MPa Cell yp = 0 –0.7 MPa = ys + 0 MPa ys = –0.7 MPa Cell membrane Cell wall 24 Aquaporins are water channels that exist in vacuole and cell membranes -They speed up osmosis, without changing the direction of water movement 25 Water potential regulates movement of water through the whole plant as well -Water moves from the soil into the roots only if the soil’s water potential is greater -It then moves along gradients of successively more negative water potentials in the stems, leaves and air 26 Evaporation of water in a leaf creates negative pressure or tension in the xylem -This “negative water potential” literally pulls water up the stem from the roots The driving force for transpiration is the gradient in vapor pressure -From 100% relative humidity inside the leaf, to much less than 100% outside the stomata 27 There are three main forces for water movement trough xylem Transpiration Pull Cohesion Adhesion Forces (Cohesive Tension Theory) Root Pressure Transpiration Pull: It is the pulling force responsible for lifting the water column. As water is lost in form of water vapour to atmosphere from the mesophyll cells by transpiration, a negative hydrostatic pressure is created in the mesophyll cells which in turn draw water from veins of the leaves. The negative tension is then gradually transmitted downwards via xylem tissues of the leaf, stem and finally to the roots. As a result there is a continuous upward movement of water column in the plant. Thus the transpiration pull acts as pull from above on the-whole of water column of the plant which pushes the water column of xylem vessels of roots lowers leaves i.e. in an upward direction. This is how ascent of sap is affected in plants. Cohesion Adhesion Forces: (Cohesive Tension Theory) The water molecules in the chain are held together by hydrogen bonds which exist between neighboring water molecules. (cohesion) The chain of molecules is prevented from being pulled down because each water molecule in the chain is attracted to the walls of the xylem by hydropyllic attraction between water and the cellulose in the cell walls. (Adhesion) Hence the water column which is held together by cohesion and prevented from lowering by adhesion is pulled up by the tension generated from above by transpiration. It is valuable both for herbaceous grasses as well as vascular plant. Root Pressure: Root pressure is caused by active transport of mineral nutrient ions into the root xylem. Without transpiration to carry the ions up the stem, they accumulate in the root xylem and lower the water potential. Water then diffuses from the soil into the root xylem due to osmosis. Root pressure is caused by this accumulation of water in the xylem pushing on the rigid cells. Root pressure provides a force, which pushes water up the stem, but it is not enough to account for the movement of water to leaves at the top of the tallest trees. Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Water exits plant through stomata. Air Smooth Rippled surface surface H2 O The water film that coats mesophyll cell walls evaporates. Rippled cell surfaces result in higher rate of transpiration than smooth cell surfaces. Water moves up plant through xylem. Plant Adhesion due to polarity of water molecules Cohesion by hydrogen bonding between water molecules Water enters plant through roots. Soil H2 O Soil Soil Cytosol 0 –0.5–1.0 –100 y w Water potential (MPa) Proton pumps contribute to the H+ y w gradient that Mineral determines the ions directional flow of water. Proton pump Water 37 1. Physical Factors: a) Soil Factors Soil water contents Soil Temperature Soil Aeration Flooding Texture and Structure Speed of Water Movement Effective Root Zone b) Atmospheric Factors: Temperature Relative Humidity VDP Wind Speed Stress (chemical) 2. Biological Factors Plant Class (Herbaceous or Vascular ) Root Length Root System Plant Health Biological Stresses Tolerance to Stresses (Salinity) Genetic Makeup Growth Rate Growth Hormones