TRANSPORT IN PLANTS
MEANS OF TRANSPORT
Diffusion: Movement by diffusion is passive, and may be from one part of the cell to the other, or from cell to cell, or over short distances, say, from the intercellular spaces of the leaf to the outside. No energy expenditure takes place. In diffusion, molecules move in a random fashion, the net result being substances moving from regions of higher concentration to regions of lower concentration. Diffusion is a slow process and is not dependent on a ‘living system’. Diffusion is obvious in gases and liquids, but diffusion in solids rather than of solids is more likely. Diffusion is very important to plants since it is the only means for gaseous movement within the plant body. Diffusion rates are affected by the gradient of concentration, the permeability of the membrane separating them, temperature and pressure.
Facilitated Diffusion:- A gradient must already be present for diffusion to occur. The diffusion rate depends on the size of the substances; obviously smaller substances diffuse faster. The diffusion of any substance across a membrane also depends on its solubility in lipids, the major constituent of the membrane. Substances soluble in lipids diffuse through the membrane faster. Substances that have a hydrophilic moiety, find it difficult to pass through the membrane; their movement has to be facilitated. Membrane proteins provide sites at which such molecules cross the membrane. They do not set up a concentration gradient: a concentration gradient must already be present for molecules to diffuse even if facilitated by the proteins. This process is called facilitated diffusion.
In facilitated diffusion special proteins help move substances across membranes without expenditure of ATP energy. Facilitated diffusion cannot cause net transport of molecules from a low to a high concentration – this would require input of energy. Transport rate reaches a maximum when all of the protein transporters are being used (saturation). Facilitated diffusion is very specific: it allows cell to select substances for uptake. It is sensitive to inhibitors which react with protein side chains.
The proteins form channels in the membrane for molecules to pass through. Some channels are always open; others can be controlled. Some are large, allowing a variety of molecules to cross.
The porins are proteins that form huge pores in the outer membranes of the plastids, mitochondria and some bacteria allowing molecules up to the size of small proteins to pass through molecule bound to the transport protein; the transport protein then rotates and releases the molecule inside the cell, e.g., water channels – made up of eight different types of aquaporins.
(Ref: http://en.wikipedia.org/wiki/File:Scheme_facilitated_diffusion_in_cell_membrane-en.svg accessed on 10th Oct, 2009)
Passive symports and antiports: Some carrier or transport proteins allow diffusion only if two types of molecules move together. In a symport, both molecules cross the membrane in the same direction; in an antiport, they move in opposite directions. When a molecule moves across a membrane independent of other molecules, the process is called uniport.
Active Transport: Active transport uses energy to pump molecules against a concentration gradient. Active transport is carried out by membrane-proteins. Hence different proteins in the membrane play a major role in both active as well as passive transport. Pumps are proteins that use energy to carry substances across the cell membrane. These pumps can transport substances from a low concentration to a high concentration (‘uphill’ transport). Transport rate reaches a maximum when all the protein transporters are being used or are saturated. Like enzymes the carrier protein is very specific in what it carries across the membrane. These proteins are sensitive to inhibitors that react with protein side chains.
Water is essential for all physiological activities of the plant and plays a very important role in all living organisms. It provides the medium in which most substances are dissolved. The protoplasm of the cells is nothing but water in which different molecules are dissolved and (several particles) suspended.
Distribution of water within a plant varies – woody parts have relatively very little water, while soft parts mostly contain water. Terrestrial plants take up huge amount water daily but most of it is lost to the air through evaporation from the leaves, i.e., transpiration.
Because of this high demand for water, it is not surprising that water is often the limiting factor for plant growth and productivity in both agricultural and natural environments.
Water molecules possess kinetic energy. In liquid and gaseous form they are in random motion that is both rapid and constant. The greater the concentration of water in a system, the greater is its kinetic energy or ‘water potential’. Hence, it is obvious that pure water will have the greatest water potential. If two systems containing water are in contact, random movement of water molecules will result in net movement of water molecules from the system with higher energy to the one with lower energy.
Thus water will move from the system containing water at higher water potential to the one having low water potential. This process of movement of substances down a gradient of free energy is called diffusion. Water potential is denoted by the Greek symbol Psi or ψ and is expressed in pressure units such as pascals (Pa). By convention, the water potential of pure water at standard temperatures, which is not under any pressure, is taken to be zero.
If some solute is dissolved in pure water, the solution has fewer free water and the concentration of water decreases, reducing its water potential. Hence, all solutions have a lower water potential than pure water; the magnitude of this lowering due to dissolution of a solute is called solute potential or ψs. ψs is always negative. The more the solute molecules, the lower (more negative) is the ψs. For a solution at atmospheric pressure
(water potential) ψw = (solute potential) ψs.
If a pressure greater than atmospheric pressure is applied to pure water or a solution, its water potential increases. It is equivalent to pumping water from one place to another. Pressure can build up in a plant system when water enters a plant cell due to diffusion causing a pressure built up against the cell wall, it makes the cell turgid; this increases the pressure potential. Pressure potential is usually positive, though in plants negative potential or tension in the water column in the xylem plays a major role in water transport up a stem. Pressure potential is denoted as ψp.
Water potential of a cell is affected by both solute and pressure potential. The relationship between them is as follows:
Ψw = ψs + ψp
Osmosis is the term used to refer specifically to the diffusion of water across a differentially- or semi-permeable membrane. Osmosis occurs spontaneously in response to a driving force. The net direction and rate of osmosis depends on both the pressure gradient and concentration gradient.
Water will move from its region of higher chemical potential (or concentration) to its region of lower chemical potential until equilibrium is reached. At equilibrium the two chambers should have the same water potential.
If a plant cell is placed in a hypertonic solution, the plant cell loses water and hence turgor pressure, making the plant cell flaccid. Plants with cells in this condition wilt. Further water loss causes plasmolysis: pressure decreases to the point where the protoplasm of the cell peels away from the cell wall, leaving gaps between the cell wall and the membrane. Eventually cytorrhysis – the complete collapse of the cell wall – can occur. There are some mechanisms in plants to prevent excess water loss in the same way as excess water gain, but plasmolysis can be reversed if the cell is placed in a weaker solution (hypotonic solution). Stomata help keep water in the plant so it does not dry out. Wax also keeps water in the plant. The equivalent process in animal cells is called crenation.
The liquid content of the cell leaks out due to diffusion. The cell collapse and cell membrane pulls away from the cell wall(in plants). Most animal cells consist of only a phospholipid bilayer and not a cell wall, therefore shrinking up under such conditions. Plasmolysis only occurs in extreme conditions and rarely happens in nature. It is induced in the laboratory by immersing cells in strong saline or sugar solutions to cause exosmosis, often using Elodea plants or onion epidermal cells, which have coloured cell sap so that the process is clearly visible.
Plasmolysis can be of two types. It can be either concave plasmolysis or convex plasmolysis. Convex plasmolysis is always irreversible while concave plasmolysis is usually reversible.
Imbibition is a special type of diffusion when water is absorbed by solids – colloids – causing them to enormously increase in volume. The classical examples of imbibition are absorption of water by seeds and dry wood. The pressure that is produced by the swelling of wood had been used by prehistoric man to split rocks and boulders. If it were not for the pressure due to imbibition, seedlings would not have been able to emerge out of the soil into the open; they probably would not have been able to establish! Imbibition is also diffusion since water movement is along a concentration gradient; the seeds and other such materials have almost no water hence they absorb water easily. Water potential gradient between the absorbent and the liquid imbibed is essential for imbibition. In addition, for any substance to imbibe any liquid, affinity between the adsorbant and the liquid is also a pre-requisite.
LONG DISTANCE TRANSPORT OF WATER
The bulk movement of substances through the conducting or vascular tissues of plants is called translocation.
Absorbption of Water
The responsibility of absorption of water and minerals is more specifically the function of the root hairs that are present in millions at the tips of the roots. Root hairs are thin-walled slender extensions of root epidermal cells that greatly increase the surface area for absorption. Water is absorbed along with mineral solutes, by the root hairs, purely by diffusion. Once water is absorbed by the root hairs, it can move deeper into root layers by two distinct pathways:
• apoplast pathway
• symplast pathway
Within a plant, the apoplast is the free diffusional space outside the plasma membrane. It is interrupted by the Casparian strip in roots, air spaces between plant cells and the cuticula of the plant.
Structurally, the apoplast is formed by the continuum of cell walls of adjacent cells as well as the extracellular spaces, forming a tissue level compartment comparable to the symplast. The apoplastic route facilitates the transport of water and solutes across a tissue or organ. This process is known as apoplastic transport.
The symplast of a plant is the inner side of the plasma membrane in which water (and low-molecular solutes) can freely diffuse.
The plasmodesmata allow the direct flow of small molecules such as sugars, amino acids, and ions between cells. Larger molecules, including transcription factors and plant viruses, can also be transported through with the help of actin structures.
This allows direct cytoplasm to cytoplasm flow of water and other nutrients along concentration gradients. In particular, it is used in the root systems to bring in nutrients from soil. It moves these solutes from epidermis cells through the cortex into the endodermis and eventually the pericycle, where it can be moved into the xylem for long distance transport. It is contrasted with the apoplastic flow, which uses cell wall transport.
As various ions from the soil are actively transported into the vascular tissues of the roots, water follows (its potential gradient) and increases the pressure inside the xylem. This positive pressure is called root pressure, and can be responsible for pushing up water to small heights in the stem.
Effects of root pressure is also observable at night and early morning when evaporation is low, and excess water collects in the form of droplets around special openings of veins near the tip of grass blades, and leaves of many herbaceous parts. Such water loss in its liquid phase is known as guttation.
Root pressure can, at best, only provide a modest push in the overall process of water transport. They obviously do not play a major role in water movement up tall trees. The greatest contribution of root pressure may be to re-establish the continuous chains of water molecules in the xylem which often break under the enormous tensions created by transpiration. Root pressure does not account for the majority of water transport; most plants meet their need by transpiratory pull.
Transpiration is the evaporative loss of water by plants. It occurs mainly through the stomata in the leaves. Besides the loss of water vapour in transpiration, exchange of oxygen and carbon dioxide in the leaf also occurs through pores called stomata (sing. : stoma). Normally stomata are open in the day time and close during the night. The immediate cause of the opening or closing of the stomata is a change in the turgidity of the guard cells. The inner wall of each guard cell, towards the pore or stomatal aperture, is thick and elastic. When turgidity increases within the two guard cells flanking each stomatal aperture or pore, the thin outer walls bulge out and force the inner walls into a crescent shape. The opening of the stoma is also aided due to the orientation of the microfibrils in the cell walls of the guard cells. Cellulose microfibrils are oriented radially rather than longitudinally making it easier for the stoma to open. When the guard cells lose turgor, due to water loss (or water stress) the elastic inner walls regain their original shape, the guard cells become flaccid and the stoma closes.
Factors Affecting Transpiration:
• Wind speed,
• Number and distribution of stomata,
• Number of stomatal aperture with guard cells stomata open,
• Water status of the plant,
• Canopy structure etc.
The transpiration driven ascent of xylem sap depends mainly on the following physical properties of water:
• Cohesion – mutual attraction between water molecules.
• Adhesion – attraction of water molecules to polar surfaces (such as the surface of tracheary elements).
• Surface Tension – water molecules are attracted to each other in the liquid phase more than to water in the gas phase.
These properties give water high tensile strength, i.e., an ability to resist a pulling force, and high capillarity, i.e., the ability to rise in thin tubes. In plants capillarity is aided by the small diameter of the tracheary elements – the tracheids and vessel elements.
As water evaporates through the stomata, since the thin film of water over the cells is continuous, it results in pulling of water, molecule by molecule, into the leaf from the xylem. Also, because of lower concentration of water vapour in the atmosphere as compared to the substomatal cavity and intercellular spaces, water diffuses into the surrounding air. This creates a ‘pull’.
Transpiration and Photosynthesis
Transpiration has more than one purpose; it
• creates transpiration pull for absorption and transport of plants
• supplies water for photosynthesis
• transports minerals from the soil to all parts of the plant
• cools leaf surfaces, sometimes 10 to 15 degrees, by evaporative cooling
• maintains the shape and structure of the plants by keeping cells turgid
An actively photosynthesising plant has an insatiable need for water. Photosynthesis is limited by available water which can be swiftly depleted by transpiration. The humidity of rainforests is largely due to this vast cycling of water from root to leaf to atmosphere and back to the soil. The evolution of the C4 photosynthetic system is probably one of the strategies for maximising the availability of CO2 while minimising water loss. C4 plants are twice as efficient as C3 plants in terms of fixing carbon (making sugar). However, a C4 plant loses only half as much water as a C3 plant for the same amount of CO2 fixed.
UPTAKE AND TRANSPORT OF MINERAL NUTRIENTS
Uptake of Mineral Ions
Unlike water, all minerals cannot be passively absorbed by the roots. Two factors account for this:
(i) minerals are present in the soil as charged particles (ions) which cannot move across cell membranes and
(ii) the concentration of minerals in the soil is usually lower than the concentration of minerals in the root.
Therefore, most minerals must enter the root by active absorption into the cytoplasm of epidermal cells. This needs energy in the form of ATP. The active uptake of ions is partly responsible for the water potential gradient in roots, and therefore for the uptake of water by osmosis. Some ions also move into the epidermal cells passively. Ions are absorbed from the soil by both passive and active transport.
Specific proteins in the membranes of root hair cells actively pump ions from the soil into the cytoplasms of the epidermal cells. Like all cells, the endodermal cells have many transport proteins embedded in their plasma membrane; they let some solutes cross the membrane, but not others.
Translocation of Mineral Ions
After the ions have reached xylem through active or passive uptake, or a combination of the two, their further transport up the stem to all parts of the plant is through the transpiration stream. The chief sinks for the mineral elements are the growing regions of the plant, such as the apical and lateral meristems, young leaves, developing flowers, fruits and seeds, and the storage organs. Unloading of mineral ions occurs at the fine vein endings through diffusion and active uptake by these cells.
Mineral ions are frequently remobilised, particularly from older, senescing parts. Older dying leaves export much of their mineral content to younger leaves. Similarly, before leaf fall in decidous plants, minerals are removed to other parts. Elements most readily mobilised are phosphorus, sulphur, nitrogen and potassium. Some elements that are structural components like calcium are not remobilised.
An analysis of the xylem exudates shows that though some of the nitrogen travels as inorganic ions, much of it is carried in the organic form as amino acids and related compounds. Similarly, small amounts of P and S are carried as organic compounds. In addition, small amount of exchange of materials does take place between xylem and phloem.
Food, primarily sucrose, is transported by the vascular tissue phloem from a source to a sink. Usually the source is understood to be that part of the plant which synthesises the food, i.e., the leaf, and sink, the part that needs or stores the food. But, the source and sink may be reversed depending on the season, or the plant’s needs. Sugar stored in roots may be mobilised to become a source of food in the early spring when the buds of trees, act as sink; they need energy for growth and development of the photosynthetic apparatus.
Since the source-sink relationship is variable, the direction of movement in the phloem can be upwards or downwards, i.e., bi-directional. This contrasts with that of the xylem where the movement is always unidirectional, i.e., upwards. Hence, unlike one-way flow of water in transpiration, food in phloem sap can be transported in any required direction so long as there is a source of sugar and a sink able to use, store or remove the sugar.
Phloem sap is mainly water and sucrose, but other sugars, hormones and amino acids are also transported or translocated through phloem.
The Pressure Flow or Mass Flow Hypothesis
The accepted mechanism used for the translocation of sugars from source to sink is called the pressure flow hypothesis. As glucose is prepared at the source (by photosynthesis) it is converted to sucrose (a dissacharide). The sugar is then moved in the form of sucrose into the companion cells and then into the living phloem sieve tube cells by active transport. This process of loading at the source produces a hypertonic condition in the phloem. Water in the adjacent xylem moves into the phloem by osmosis. As osmotic pressure builds up the phloem sap will move to areas of lower pressure. At the sink osmotic pressure must be reduced. Again active transport is necessary to move the sucrose out of the phloem sap and into the cells which will use the sugar – converting it into energy, starch, or cellulose. As sugars are removed, the osmotic pressure decreases and water moves out of the phloem.
To summarise, the movement of sugars in the phloem begins at the source, where sugars are loaded (actively transported) into a sieve tube. Loading of the phloem sets up a water potential gradient that facilitates the mass movement in the phloem.
Phloem tissue is composed of sieve tube cells, which form long columns with holes in their end walls called sieve plates. Cytoplasmic strands pass through the holes in the sieve plates, so forming continuous filaments. As hydrostatic pressure in the phloem sieve tube increases, pressure flow begins, and the sap moves through the phloem. Meanwhile, at the sink, incoming sugars are actively transported out of the phloem and removed Sugars leave sieve tube for metabolism and storage; water follows by osmosis as complex carbohydrates. The loss of solute produces a high water potential in the phloem, and water passes out, returning eventually to xylem.