Chapter 11 Transport in Plants Chapter 12 Mineral Nutrition Chapter 13 Photosynthesis in Higher Plants Chapter 14 Respiration in Plants Chapter 15 Plant Growth and Development UNIT 4 PLANT PHYSIOLOGY The description of structure and variation of living organisms over a period of time, ended up as two, apparently irreconcilable perspectives on biology. The two perspectives essentially rested on two levels of organisation of life forms and phenomena. One described at organismic and above level of organisation while the second described at cellular and molecular level of organisation. The first resulted in ecology and related disciplines. The second resulted in physiology and biochemistry. Description of physiological processes, in flowering plants as an example, is what is given in the chapters in this unit. The processes of mineral nutrition of plants, photosynthesis, transport, respiration and ultimately plant growth and development are described in molecular terms but in the context of cellular activities and even at organism level. Wherever appropriate, the relation of the physiological processes to environment is also discussed. TRANSPORT IN PLANTS 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. 11.3 LONG DISTANCE TRANSPORT OF WATER At some earlier stage you might have carried out an experiment where you had placed a twig bearing white flowers in coloured water and had watched it turn colour. On examining the cut end of the twig after a few hours you had noted the region through which the coloured water moved. That experiment very easily demonstrates that the path of water movement is through the vascular bundles, more specifically, the xylem. Now we have to go further and try and understand the mechanism of movement of water and other substances up a plant. Long distance transport of substances within a plant cannot be by diffusion alone. Diffusion is a slow process. It can account for only short distance movement of molecules. For example, the movement of a molecule across a typical plant cell (about 50 µm) takes approximately 2.5 s. At this rate, can you calculate how many years it would take for the movement of molecules over a distance of 1 m within a plant by diffusion alone? In large and complex organisms, often substances have to be moved across very large distances. Sometimes the sites of production or absorption and sites of storage are too far from each other; diffusion or active transport would not suffice. Special long distance transport systems become necessary so as to move substances across long distances and at a much faster rate. Water and minerals, and food are generally moved by a mass or bulk flowsystem. Mass flow is the movement of substances in bulk or en masse from one point to another as a result of pressure differences between the two points. It is a characteristic of mass flow that substances, whether in solution or in suspension, are swept along at the same pace, as in a flowing river. This is unlike diffusion where different substances move independently depending on their concentration gradients. Bulk flow can be achieved either through a positive hydrostatic pressure gradient (e.g., a garden hose) or a negative hydrostatic pressure gradient (e.g., suction through a straw). BIOLOGY filaments form a network around the young root or they penetrate the root cells. The hyphae have a very large surface area that absorb mineral ions and water from the soil from a much larger volume of soil that perhaps a root cannot do. The fungus provides minerals and water to the roots, in turn the roots provide sugars and N-containing compounds to the mycorrhizae. Some plants have an obligate association with the mycorrhizae. For example, Pinus seeds cannot germinate and establish without the presence of mycorrhizae. 11.3.2 Water Movement up a Plant We looked at how plants absorb water from the soil, and move it into the vascular tissues. We now have to try and understand how this water is transported to various parts of the plant. Is the water movement active, or is it still passive? Since the water has to be moved up a stem against gravity, what provides the energy for this? 184.108.40.206 Root Pressure 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. How can we see that root pressure exists? Choose a small soft-stemmed plant and on a day, when there is plenty of atmospheric moisture, cut the stem horizontally near the base with a sharp blade, early in the morning. You will soon see drops of solution ooze out of the cut stem; this comes out due to the positive root pressure. If you fix a rubber tube to the cut stem as a sleeve you can actually collect and measure the rate of exudation, and also determine the composition of the exudates. 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. 220.127.116.11 Transpiration pull Despite the absence of a heart or a circulatory system in plants, the flow of water upward through the xylem in plants can achieve fairly high rates, TRANSPORTIN PLANTS up to 15 metres per hour. How is this movement accomplished? A long standing question is, whether water is ‘pushed’ or ‘pulled’ through the plant. Most researchers agree that water is mainly ‘pulled’ through the plant, and that the driving force for this process is transpiration from the leaves. This is referred to as the cohesion-tension-transpiration pull model of water transport. But, what generates this transpirational pull? Water is transient in plants. Less than 1 per cent of the water reaching the leaves is used in photosynthesis and plant growth. Most of it is lost through the stomata in the leaves. This water loss is known as transpiration. You have studied transpiration in an earlier class by enclosing a healthy plant in polythene bag and observing the droplets of water formed inside the bag. You could also study water loss from a leaf using cobalt chloride paper, which turns colour on absorbing water. 11.4 TRANSPIRATION 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. Usually the lower surface of a dorsiventral (often dicotyledonous) leaf has a greater number of stomata while in an isobilateral (often monocotyledonous) leaf they are about equal on both surfaces. Transpiration is affected by several external factors: temperature, light, humidity, wind speed. Plant factors that affect transpiration include number and distribution of stomata, per cent of open Figure11.8 A stomatal aperture with guard cells 188 BIOLOGY stomata, 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. The process of photosynthesis requires water. The system of xylem vessels from the root to the leaf vein can supply the needed water. But what force does a plant use to move water molecules into the leaf parenchyma cells where they are needed? 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’ (Figure 11.9). Measurements reveal that the forces generated by transpiration can create pressures sufficient to lift a xylem sized column of water over 130 metres high. Palisade Xylem Phloem Guard Cell StomatalStomaDiffusion into poresurrounding air Figure11.9 Water movement in the leaf. Evaporation from the leaf sets up a pressure gradient between the outside air and the air spaces of the leaf. The gradient is transmitted into the photosynthetic cells and on the water-filled xylem in the leaf vein.