The evolution,structure and functioning of stomata

Summary

The anatomy and ultrastructure of guard cells from a range of species varying from the primitive types, such as mosses, to the advanced grasses and orchids are described. An attempt is made to trace the lines along which stomata developed and to define what might be considered advanced stomata. Additionally, the differentiation of guard cells from guard mother cells is discussed. Of particular note is the preprophase band of microtubules which marks the zone where the future cell wall will form and the movement of the spindle and developing cell plate through 45 degrees. The structure and function of guard cells are intimately linked. Stomata are turgor regulated valves; the osmotica for absorbing water during opening are K⁺, Cl^- and malate anions which accumulate in the guard cell vacuoles. Upon stomatal closure, K⁺ and Cl^- exit from the guard cells while at least some of the carbon from malate is channelled into starch and there is a resultant loss of guard cell turgor. The Calvin cycle may be absent or of low activity in guard cell chloroplasts and under those circumstances a source of carbon and energy to sustain the guard cells is needed. Hence it is believed that sucrose is transported into the guard cells from mesophyll cells. A brief consideration of the mechanism by which the ions are transported across the plasma membrane and tonoplast is made: the driving force for the K⁺, Cl^- and malate movement across the membranes is the proton motive force set up by proton-pumping ATPases.