Imbibition is a special type of diffusion when water is absorbed by solids – colloids – causing them to 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.

 

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 to long 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 flow system. 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).

The bulk movement of substances through the conducting or vascular tissues of plants is called translocation.

Do you remember studying cross sections of roots, stems and leaves of higher plants and studying the vascular system? The higher plants have highly specialised vascular tissues – xylem and phloem. Xylem is associated with translocation of mainly water, mineral salts, some organic nitrogen and hormones, from roots to the aerial parts of the plants. The phloem translocates a variety of organic and inorganic solutes, mainly from the leaves to other parts of the plants.

 

How do Plants Absorb Water?

 

We know that the roots absorb most of the water that goes into plants; obviously that is why we apply water to the soil and not on the leaves. 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 :

1. apoplast pathway
2. symplast pathway

The apoplast is the system of adjacent cell walls that is continuous throughout the plant, except at the casparian strips of the endodermis in the roots (Figure 11.6).

 

Figure 11.6 Pathway of water movement in the root

 

The apoplastic movement of water occurs exclusively through the intercellular spaces and the walls of the cells. Movement through the apoplast does not involve crossing the cell membrane. This movement is dependent on the gradient. The apoplast does not provide any barrier to water movement and water movement is through mass flow. As water evaporates into the intercellular spaces or the atmosphere, tension develop in the continuous stream of water in the apoplast, hence mass flow of water occurs due to the adhesive and cohesive properties of water.

 

The symplastic system is the system of interconnected protoplasts. Neighbouring cells are connected through cytoplasmic strands that extend through plasmodesmata. During symplastic movement, the water travels through the cells – their cytoplasm; intercellular movement is through the plasmodesmata. Water has to enter the cells through the cell membrane, hence the movement is relatively slower. Movement is again down a potential gradient. Symplastic movement may be aided by cytoplasmic streaming. You may have observed cytoplasmic streaming in cells of the Hydrilla leaf; the movement of chloroplast due to streaming is easily visible.

 

Most of the water flow in the roots occurs via the apoplast since the cortical cells are loosely packed, and hence offer no resistance to water movement. However, the inner boundary of the cortex, the endodermis, is impervious to water because of a band of suberised matrix called the casparian strip. Water molecules are unable to penetrate the layer, so they are directed to wall regions that are not suberised, into the cells proper through the membranes. The water then moves through the symplast and again crosses a membrane to reach the cells of the xylem. The movement of water through the root layers is ultimately symplastic in the endodermis. This is the only way water and other solutes can enter the vascular cylinder.

 

Once inside the xylem, water is again free to move between cells as well as through them. In young roots, water enters directly into the xylem vessels and/or tracheids. These are non-living conduits and so are parts of the apoplast. The path of water and mineral ions into the root vascular system is summarised in Figure 11.7.

 

 

Figure 11.7 Symplastic and apoplastic pathways of water and ion absorption and movement in roots

 

Some plants have additional structures associated with them that help in water (and mineral) absorption. A mycorrhiza is a symbiotic association of a fungus with a root system. The fungal 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.

 

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?

 

ROOTPressure

 

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.

Transpiration pull

Despite the absence of a heart or a circulatory system in plants, the upward flow of water through the xylem in plants can achieve fairly high rates, 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.

 

TRANSPIRATION

Transpiration is the evaporative loss of water by plants. It occurs mainly through stomata (sing. : stoma). Besides the loss of water vapour in transpiration, exchange of oxygen and carbon dioxide in the leaf also occurs through these stomata. Normally stomata are open in the day time and close during the night. The immediate cause of the opening or closing of stomata is a change in the turgidity of the guard cells.

Figure11.8 A stomatal aperture with 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 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:

 

1. Cohesion – mutual attraction between water molecules.

 

2. Adhesion – attraction of water molecules to polar surfaces (such as the surface of tracheary elements).

 

3. 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).

 

 

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.

 

Measurements reveal that the forces generated by transpiration can create pressures sufficient to lift a xylem sized column of water over 130 metres high.

 

Transpiration and Photosynthesis – a Compromise

 

Transpiration has more than one purpose; it

 

1. creates transpiration pull for absorption and transport of plants

2. supplies water for photosynthesis

3. transports minerals from the soil to all parts of the plant

4.cools leaf surfaces, sometimes 10 to 15 degrees, by evaporative cooling

5.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 dioxide (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

Plants obtain their carbon and most of their oxygen from CO₂ in the atmosphere. However, their remaining nutritional requirements are obtained from water and minerals in the soil.

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. Transport proteins of endodermal cells are control points, where a plant adjusts the quantity and types of solutes that reach the xylem. Note that the root endodermis because of the layer of suberin has the ability to actively transport ions in one direction only.

 

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. Hence, it is not that we can clearly make a distinction and say categorically that xylem transports only inorganic nutrients while phloem transports only organic materials, as was traditionally believed.

 

 

 

​​​​​​​PHLOEM TRANSPORT: FLOW FROM SOURCE TO SINK

 

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. (see Figure 11.10).

 

 

Figure11.10 Diagrammatic presentation of mechanism of translocation

 

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 sieve tube of phloem 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 as complex carbohydrates. The loss of solute produces a high water potential in the phloem, and water passes out, returning eventually to xylem. A simple experiment, called girdling, was used to identify the tissues through which food is transported. On the trunk of a tree a ring of bark up to a depth of the phloem layer, can be carefully removed. In the absence of downward movement of food the portion of the bark above the ring on the stem becomes swollen after a few weeks. This simple experiment shows that phloem is the tissue responsible for translocation of food; and that transport takes place in one direction, i.e., towards the roots. Thisexperiment can be performed by you easily.