Transpiration - Major Plant Highlights
Root Detail– The major path for water movement into plants is from soil to roots. Water enters near the tip of a growing root, the same region where root hairs grow. The surface of the root hairs needs to be in close contact with the soil to access soil water. Water diffuses into the root, where it can take at least three different pathways to eventually reach the xylem, the conduit located at the interior of the root that carries the soil water to the leaves. View the next level of this animation to see the possible pathways that water can take across a root.
What path does water take to reach the leaf from the root hair? Once water has entered a root hair, it must move across the cortex and endodermis before it reaches the xylem. Water will take the path of least resistance through a root to reach the xylem.
Water can move across the root via three different pathways. One path is the apoplastic path where the water molecule stays between cells in the cell wall region, never crossing membranes or entering a cell. The other two routes, called cellular pathways, require the water molecule to actually move across a membrane.
The first cellular pathway is the transmembrane path where water moves from cell to cell across membranes; it will leave one cell by traversing its membrane and will re-enter another cell by crossing its membrane. The second cellular path is the symplastic path which takes the water molecule from cell to cell using the intercellular connections called the plasmodesmata which are membrane connections between adjacent cells. Regardless of the pathway, once the water molecule has traversed the cortex, it must now cross the endodermis. The endodermis is a layer of cells with a waxy inlay or mortar called the Casparian strip that stops water movement between cells. At this point, water is forced to move through the membranes of endodermal cells, creating a sieving effect. Once in the endodermal cells, the water freely enters the xylem cells where it joins the fast moving column of water or transpiration stream, headed to the leaves.
Xylem Details– The xylem is probably the longest part of the pathway that water takes on its way to the leaves of a plant. It is also the path of least resistance, with about a billion times less resistance than cell to cell transport of water. Xylem cells are called tracheids (cells with narrower diameters) or vessels (cells with wider diameters). Their cell walls contain cellulose and lignin making them extremely rigid. Xylem cells contain no membranes and are considered dead. These cells overlap to create a series of pathways that water can take as it heads to the leaves. There is no single column of xylem cells carrying water.
How do stomata open? Stomata sense environmental cues, like light, to open. These cues start a series of reactions that cause their guard cells to fill with water. Let’s follow a scenario where the sun is rising and a cotton plant is signaled to open its stomata:
- Signal received: The blue light at dawn is the signal that is recognized by a receptor on the guard cell.
- The receptor signals the H+-ATPases on the guard cell’s plasma membrane to start pumping protons (H+) out of the guard cell. This loss of positive charge creates a negative charge in the cell.
- Potassium ions (K+) enter the guard cell through channels in the membrane, moving toward its more negative interior.
- As the potassium ions accumulate in the guard cell, the osmotic pressure is lowered.
- A lower osmotic pressure attracts water to enter the cell.
- As water enters the guard cell, its hydrostatic pressure increases.
- The pressure causes the shape of the guard cells to change and a pore is formed, allowing gas exchange.
Cavitation– Cavitation is the filling of a xylem vessel or tracheid with air. It is also known as an ‘embolism’ or ‘air-lock’. Remember that during transpiration, the column of water is being pulled out of the plant by evaporation at the leaf cell surface. When this ‘pulling’ of water out of the plant becomes greater than the ability of the water molecules to stay together, the column of water will break.
Plants are particularly sensitive to cavitation during the hottest part of the day when there is not enough water available from the soil to keep up with the demand for water while it is evaporating off the leaf surface. Cavitation also occurs under freezing conditions. Because the solubility of gas in ice is very low, gas comes out of solution when the xylem sap freezes. Freezing of xylem sap is a problem in the spring when the ice thaws, leaving a bubble in a xylem vessel. These bubbles can block water transport and cause water deficit in leaves.
Plants avoid cavitation or minimize its damage through several mechanisms:
- Xylem cells possess pits or tiny holes that allow liquid water transport, but do not allow the gas bubble to escape; this structural characteristic helps keep the gas bubble in one cell, so the other xylem cells can continue to transport water up the plant.
- Water will detour around any xylem cell containing an air bubble through the pits as well.
- The gas bubble will re-dissolve into liquid water when the pulling of water through the xylem is reduced, such as during the night when water is not being pulled out of the leaf via transpiration because the stomata are closed.
- Xylem cells with narrower diameters (tracheids) compared to those with wider diameters (vessels) avoid cavitation because the column of water in a cell with a narrow diameter is better able to resist bubble formation or rupture.
Stomata Details– The stomata are the primary control mechanisms that plants use to reduce water loss and they are able to do so quickly. Stomata are sensitive to the environmental cues that trigger the stomata to open or close. The major role of stomata is to allow carbon dioxide entry to drive photosynthesis and at the same time allow the exit of water as it evaporates, cooling the leaf. Two specialized cells called ‘guard cells’ make up each stoma (stoma is singular for stomata). Plants have many stomata (up to 400 per mm2) on their leaf surfaces and they are usually on the lower surface to minimize water loss.
SIDE VIEW OF STOMATA– Environmental cues that affect stomata opening and closing are light, water, temperature, and the concentration of CO2 within the leaf. Stomata will open in the light and close in the dark. However, stomata can close in the middle of the day if water is limiting, CO2 accumulates in the leaf, or the temperature is too hot. If the plant lacks water, stomata will close because there will not be enough water to create pressure in the guard cells for stomatal opening; this response helps the plant conserve water.
Using sound-sensing equipment, one can actually hear a ‘click’ when the water molecules split from one another. Unique structural characteristics help the plant contain the air bubble so that it does not totally disrupt water movement up the plant. If the leaf’s internal concentration of CO2 increases, the stomata are signaled to close because respiration is releasing more CO2 than photosynthesis is using. There is no need to keep the stomata open and lose water if photosynthesis is not functioning. Alternatively, if the leaf’s CO2 concentration is low, the stomata will stay open to continue fueling photosynthesis. High temperatures will also signal stomata to close.
High temperatures will increase the water loss from the leaf. With less water available, guard cells can become flaccid and close. Another effect of high temperatures is that respiration rates rise above photosynthesis rates causing an increase of CO2 in the leaves; high internal CO2 will cause stomata to close as well. Remember that some plants may open their stomata under high temperatures so that transpiration will cool the leaves.
TOP VIEW OF STOMATA –
OPEN STOMATA– When stomata are signaled to open, potassium ions (K+) enter the guard cells. This causes water to enter down its water potential gradient, creating a hydrostatic pressure in the guard cell that changes the shape of the stoma. Guard cells expand on the outer edges of the stoma, but not on the inner side, resulting in kidney-shaped cells and an opening or pore between the two guard cells for gas exchange. Kidney-shaped guard cells are characteristic of dicots; however, many plant (e.g. grasses, other monocots) have dumbbell-shaped stomata. The shape taken by the guard cells is dependent on cellulose microfibrils that fan out radially from the pore, somewhat similar to radial tires. The cellulose microfibrils are rigid and do not stretch when water has entered the cell. The cell walls surrounding the stomatal opening are thickened, preventing that side of the guard cell from expanding. Therefore, when pressure in the cell increases due to water entry, guard cell does not widen, but rather the outer edge stretches disproportionately more than the inner edge. This unequal stretching allows the pore to form between the two guard cells.
CLOSED STOMATA– Stomata must be open for the plant to photosynthesize; however, open stomata present a risk of losing too much water through transpiration. Stomata close when the guard cells lose water and become flaccid. This occurs because potassium ions move back out of the guard cell, followed by water that lowers the pressure in the