In botany, a stoma (from Greek στόμα, "mouth", : stomata), also called a stomate (: stomates), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bordered by a pair of specialized parenchyma cells known as guard cells that regulate the size of the stomatal opening.
The term is usually used collectively to refer to the entire stomatal complex, consisting of the paired guard cells and the pore itself, which is referred to as the stomatal aperture. Air, containing oxygen, which is used in respiration, and carbon dioxide, which is used in photosynthesis, passes through stomata by gaseous diffusion. Water vapour diffuses through the stomata into the atmosphere as part of a process called transpiration.
Stomata are present in the sporophyte generation of all land plant groups except liverworts. In vascular plants the number, size and distribution of stomata varies widely. Dicotyledons usually have more stomata on the lower surface of the leaves than the upper surface. Monocotyledons such as onion, oat and maize may have about the same number of stomata on both leaf surfaces. In plants with floating leaves, stomata may be found only on the upper epidermis and submerged leaves may lack stomata entirely. Most tree species have stomata only on the lower leaf surface. Leaves with stomata on both the upper and lower leaf surfaces are called amphistomatous leaves; leaves with stomata only on the lower surface are hypostomatous, and leaves with stomata only on the upper surface are epistomatous or hyperstomatous. Size varies across species, with end-to-end lengths ranging from 10 to 80 µm and width ranging from a few to 50 µm.
Function
CO2 gain and water loss
Carbon dioxide, a key reactant in photosynthesis, is present in the atmosphere at a concentration of about 400 ppm. Most plants require the stomata to be open during daytime. The air spaces in the leaf are saturated with water vapour, which exits the leaf through the stomata in a process known as transpiration. Therefore, plants cannot gain carbon dioxide without simultaneously losing water vapour.
Alternative approaches
Ordinarily, carbon dioxide is fixed to ribulose 1,5-bisphosphate (RuBP) by the enzyme RuBisCO in mesophyll cells exposed directly to the air spaces inside the leaf. This exacerbates the transpiration problem for two reasons: first, RuBisCo has a relatively low affinity for carbon dioxide, and second, it fixes oxygen to RuBP, wasting energy and carbon in a process called photorespiration. For both of these reasons, RuBisCo needs high carbon dioxide concentrations, which means wide stomatal apertures and, as a consequence, high water loss.
Narrower stomatal apertures can be used in conjunction with an intermediary molecule with a high carbon dioxide affinity, phosphoenolpyruvate carboxylase (PEPcase). Retrieving the products of carbon fixation from PEPCase is an energy-intensive process, however. As a result, the PEPCase alternative is preferable only where water is limiting but light is plentiful, or where high temperatures increase the solubility of oxygen relative to that of carbon dioxide, magnifying RuBisCo's oxygenation problem.
C.A.M plants
A group of mostly desert plants called "C.A.M" plants (Crassulacean acid metabolism, after the family Crassulaceae, which includes the species in which the CAM process was first discovered) open their stomata at night (when water evaporates more slowly from leaves for a given degree of stomatal opening), use PEPcarboxylase to fix carbon dioxide and store the products in large vacuoles. The following day, they close their stomata and release the carbon dioxide fixed the previous night into the presence of RuBisCO. This saturates RuBisCO with carbon dioxide, allowing minimal photorespiration. This approach, however, is severely limited by the capacity to store fixed carbon in the vacuoles, so it is preferable only when water is severely limited.