Transpiration

Transpiration is the process of water movement through a plant and its evaporation from aerial parts, such as leaves, stems and flowers. Water is necessary for plants but only a small amount of water taken up by the roots is used for growth and metabolism. The remaining 97–99.5% is lost by transpiration and guttation.[1] Leaf surfaces are dotted with pores called stomata (singular "stoma"), and in most plants they are more numerous on the undersides of the foliage. The stomata are bordered by guard cells and their stomatal accessory cells (together known as stomatal complex) that open and close the pore.[2] Transpiration occurs through the stomatal apertures, and can be thought of as a necessary "cost" associated with the opening of the stomata to allow the diffusion of carbon dioxide gas from the air for photosynthesis. Transpiration also cools plants, changes osmotic pressure of cells, and enables mass flow of mineral nutrients and water from roots to shoots. Two major factors influence the rate of water flow from the soil to the roots: the hydraulic conductivity of the soil and the magnitude of the pressure gradient through the soil. Both of these factors influence the rate of bulk flow of water moving from the roots to the stomatal pores in the leaves via the xylem.[3]

Overview of transpiration:
  1. Water is passively transported into the roots and then into the xylem.
  2. The forces of cohesion and adhesion cause the water molecules to form a column in the xylem.
  3. Water moves from the xylem into the mesophyll cells, evaporates from their surfaces and leaves the plant by diffusion through the stomata
Transpiration of Water in Xylem
Stoma in a tomato leaf shown via colorized scanning electron microscope
The clouds in this image of the Amazon Rainforest are a result of evapotranspiration.

Mass flow of liquid water from the roots to the leaves is driven in part by capillary action, but primarily driven by water potential differences. If the water potential in the ambient air is lower than the water potential in the leaf airspace of the stomatal pore, water vapor will travel down the gradient and move from the leaf airspace to the atmosphere. This movement lowers the water potential in the leaf airspace and causes evaporation of liquid water from the mesophyll cell walls. This evaporation increases the tension on the water menisci in the cell walls and decrease their radius and thus the tension that is exerted on the water in the cells. Because of the cohesive properties of water, the tension travels through the leaf cells to the leaf and stem xylem where a momentary negative pressure is created as water is pulled up the xylem from the roots.[4] As evaporation occurs at the leaf surface, the properties of adhesion and cohesion work in tandem to pull water molecules from the roots, through xylem tissue, and out of the plant through stomata.[5] In taller plants and trees, the force of gravity can only be overcome by the decrease in hydrostatic (water) pressure in the upper parts of the plants due to the diffusion of water out of stomata into the atmosphere. Water is absorbed at the roots by osmosis, and any dissolved mineral nutrients travel with it through the xylem.

The cohesion-tension theory explains how leaves pull water through the xylem. Water molecules stick together, or exhibit cohesion. As a water molecule evaporates from the surface of the leaf, it pulls on the adjacent water molecule, creating a continuous flow of water through the plant.[6]