The movement of materials within the body tissues or between tissues in multicellular organisms like high plants and animals occurs in transport systems. In unicellular or low plants and animals, such movement is by the transport mechanisms of molecules within a transport medium by diffusion, osmosis, active transport and mass flow, among others. In multicellular animals, the medium of transport is blood, while in plants, it is water in which the materials are dissolved and transported.
In case of diseases such as diarrhoea, dysentery, malaria, etc, the animal body may undergo dehydration. This causes serious effects on the body’s physiological process, which can lead to death. Just like dehydration in animals, in plants wilting or excess water loss results, especially during dry seasons or inadequate water supply or absorption by conducting tissues. Animals and plants have specialized cells or tissues performing special functions of the transport systems apart from transporting materials.
Transport in plants occurs on three levels:
- the uptake and release of water and solutes by individual cells
- absorption of water and minerals from the soil by root cells
- short-distance transport of substances from cell to cell
- loading of sucrose from photosynthetic cells into the sieve tube cells of the phloem
- long-distance transport of sap within the xylem and phloem
- this is a whole plant phenomena – transport of photosynthate from leaf to root
Survival of the plant depends on balancing water uptake and water loss.
In an animal cell, water flows from hypotonic to hypertonic solutions, but in a plant cell, there is the added presence of the pressure created by the cell wall
The combination of solute concentration differences and physical pressure are incorporated into water potential, abbreviated with the Greek letter psi ()
- Water will flow through a membrane from a solution of high water potential to a solution of low water potential
- Water potential is measured in units of megapascals (MPa)
- Pure water has a water potential of 0 MPa ( = 0 MPa)
- These two forces combine to form the following equation:
- = p + s
- = total water potential
- p = water potential due to pressure
- May be positive or negative
- s = water potential due solute concentration (also known as Osmotic Potential)
- Always negative or zero
Movement of Water Through Cells –
Two Routes, the Symplast and the Apoplast
- Movement of water and solutes through the continuous connection of cytoplasm (though plasmodesmata)
- No crossing of the plasma membrane (once it is in the symplast – however, if the solute was initially external to the cell, then it must have crossed one plasma membrane to enter the symplast)
- Movement of water and solutes through the cell walls and the intercellular spaces
- No crossing of the plasma membrane
- More rapid – less resistance to the flow of water
Absorption of Water and Minerals by Roots
Absorption is a surface area phenomenon – the more surface area there is, the more absorption there will be.
- Root hairs – extensions of the root epidermal cells to increase surface area
- Mycorrhizae – fungal associations with roots – greatly increase surface area
- as much as three meters of fungal hyphae can extend from each centimeter of root
- this is an ancient association – some of the oldest terrestrial plant fossils have fungal association.
- As water is drawn into the root, dissolved minerals are also brought into the root
- Water flows through the apoplast and the symplast on its way to the xylem
- The majority of the water, however, travels through the apoplast
The Endodermis – The Root’s Border Guard
Water flowing through the apoplast contains many minerals that the plant needs – it may also contains toxins and substances that the plant may not want. However, since the water is flowing through the apoplast, there is no way to prevent the passive transport of these toxins, until the water hits the endodermis.
Cells of the endodermis possess cell walls that are ringed by the Casparian Strip, a waxy layer (composed of suberin).
- The Casparian Strip is a wax and therefore prevents the apoplastic flow of water
- Water must pass through the plasma membrane and enter the symplast
- The plasma membrane of the endodermal cells contain many transport proteins to actively transport some molecules in and others to pump other molecules out
- Once water passes under the Casparian Strip in the endodermal cells, it is free to enter the apoplast again on its way to the xylem.
Two main types of plant tissue are used in transport
Xylem transports water and minerals. Phloem transports organic molecules such as the products of photosynthesis.
There are four types of xylem cells:
- Xylem vessels: Consist of dead hollow cells because the walls are lignified and the cell contents disintegrate. The lignin makes the cell wall impermeable so they are in effect waterproof. It also makes the vessels extremely strong and prevents them from collapsing. They have a wide lumen and are linked end to end to create a long, hollow tube since the end cell walls have one or many perforations in them. This allows the transport of large volumes of water. The sidewalls have bordered pits (unlignified areas) to allow lateral movement of water. Xylem vessels are found in angiosperms.
- Tracheids: Similar to vessels but with narrower lumens and connected by pits. They have tapered ends so that they dovetail together. Tracheids are found in conifers.
- Parenchyma: Living cells with thin cellulose walls. They can store water, which makes them turgid and so gives them a supporting role.
- Fibres: They provide strength because their walls are lignified (and therefore, dead).
Movement in the root
Water enters through the root hair cells and then moves across into the xylem tissue in the centre of the root. Water moves in this direction because the soil water has higher water potential, than the solution inside the root hair cells.
This is because the cell sap has organic and inorganic molecules dissolved in it. The root hairs provide a large surface area over which water can be absorbed.
Minerals are also absorbed but, as you should be able to work out, their absorption requires energy in the form of ATP because they are absorbed by active transport. They have to be pumped against the concentration gradient.
Water taken up by the root hairs moves across the cortex of the root either via the cytoplasm of the cells in between the root hair cell and the xylem (the symplast pathway) or through the cell walls of these cells (the apoplast pathway). The root hair cell will have higher water potential than the cell next to it. As always, water moves by osmosis to where the water potential is lower. In this way, as water is always being absorbed by the root hairs, water will always move towards the centre of the root.
When the water reaches a part of the root called the endodermis, it encounters a thick, waxy band of suberin in the cell walls. This is the Casparian strip and it is impenetrable. In order to cross the endodermis, the water that has been moving through the cell walls must now move into the cytoplasm.
Once it has moved across the endodermis, it continues down the water potential gradient until it reaches a pit in the xylem vessel. It enters the vessel and then moves up towards the leaves.
Movement in the xylem
Water evaporates from the mesophyll cells into air spaces in the leaf. If the air surrounding the leaf has less water vapour than the air in the intercellular spaces, water vapour will leave the leaf through stomata.
This process is called transpiration and will continue as long as the stomata are open and the air outside is not too humid. On dry, windy days when water vapour is continually diffusing out and being removed, transpiration will increase in rate.
Although this loss of water can cool the plant, it is essential that the plant does not lose too much water. Therefore water must be continuously supplied to the leaves. The xylem ensures that this happens. Xerophytes are plants which are well adapted to living where conditions are very dry. They may have rolled up leaves – for example, Marram grass which exposes the waterproof cuticle on the outside and means the stomata open into an inner humid space. Other Xerophytes store water in their stems and reduce the surface area of their leaves, which become spines – for example, Cactus.
Water is removed from the top of xylem vessels into the mesophyll cells down the water potential gradient. This removal of water from the xylem reduces the hydrostatic pressure exerted by the liquid so the pressure at the top is less than at the bottom. This pushes the water up the tube. The surface tension of the water molecules, the thin lumen of the xylem vessels and the attraction of the water molecules for the xylem vessel wall (adhesion), helps to keep the water flowing all the time and to keep the water column intact.
Pressure to push water up can also be increased from the bottom. By actively pumping minerals from cells surrounding the xylem into the xylem itself, more water is drawn into the xylem by osmosis.
This increase in water pressure, called root pressure, certainly helps in the process but is less important than the simple movement of water down the water potential gradient, ultimately from the soil at the bottom, to the air at the top. This is because moving water this way does not require energy (it is passive).
There are four types of phloem cells:
- Sieve tube elements: These are living, tubular cells that are connected end to end. The end cell walls have perforations in them to make sieve plates. The cytoplasm is present but in small amounts and in a layer next to the cell wall. It lacks a nucleus and most organelles so there is more space for solutes to move. The cell walls are made of cellulose so solutes can move laterally a well as vertically. Next to each sieve tube element is a companion cell.
- Companion cell: Since the sieve tube element lacks organelles, the companion cell with its nucleus, mitochondria, ribosomes, enzymes etc., controls the movement of solutes and provides ATP for active transport in the sieve tube element. Strands of cytoplasm called plasmodesmata connect the sieve tube element and companion cell.
- Parenchyma: Provides support through turgidity.
- Fibres: Provides support for the sieve tube elements.
Movement in the phloem
This process is called translocation and involves the movement of organic substances around the plant. It requires energy to create a pressure difference and so is considered an active process.
Sucrose is loaded into the phloem at a source, usually a photosynthesizing leaf. For this to occur, hydrogen ions are pumped out of the companion cell using ATP. This creates a high concentration of hydrogen ions outside the companion cell. Sucrose is loaded (moved into companion cells) by active transport, against the concentration gradient.
However, the protein carrier involved in the loading, has two sites, one for sucrose and one for a hydrogen ion. When it is used to pump sucrose into the companion cell, hydrogen will move in the opposite direction, back down its concentration gradient. This is why a high concentration of ions is needed outside the cell.
The sucrose can then diffuse down the concentration gradient into the sieve tube element via the plasmodesmata that connects the companion cell with the sieve tube element. This lowers the water potential of the sieve element so water enters by osmosis.
At another point sucrose will be unloaded from the phloem into a sink (e.g. root). It is likely that the sucrose moves out by diffusion and is then converted into another substance to maintain a concentration gradient. Again, water will follow by osmosis.
This loading and unloading results in the mass flow of substances in the phloem. There is evidence to support this theory; the rate of flow in the phloem is about 10,000 times faster than it would be if it was due only to diffusion, the pH of the phloem sap is around 8 (it is alkaline due to loss of hydrogen ions), and there is an electrical potential difference across the cell surface (negative inside due presumably to the loss of positively charged ions).
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