Studies in Stomatal Behaviour: VII. EFFECTS OF ANAEROBIC CONDITIONS UPON STOMATAL MOVEMENT--A TEST OF WILLIAMS'S HYPOTHESIS OF STOMATAL MECHANISM

An experiment was carried out to investigate stomatal responsesin wheat to four ‘closing treatments’, viz. highcarbon dioxide concentration, darkness, dry air and nil, eachgiven under both aerobic and anaerobic conditions. Thus theeffect of lack of oxygen on the closing (or opening) tendencywas estimated. Changes in calculated from resistance porometer readings were used as data and reasonsare given for thinking this is the best available measure forinvestigating stomatal dynamics in wheat. Williams's hypothesisdemands that lack of oxygen should cause stomatal opening orprevent closure; the present experiment shows that anaerobicconditions significantly increase the closing tendency when‘closing treatments’ are first applied. There isalso some suggestion that oxygen-lack itself tends to causeclosure in the absence of any other ‘closing treatment’.Williams's hypothesis in its original form is thus disproved(for wheat) but the present results would be consistent withan ‘active’ uptake of water by the guard cells contributingto stomatal opening. A nearly significant interaction betweencarbon dioxide and oxygen suggests that under anaerobic conditionsa ‘closing substance’ may perhaps be formed, forexample, by the union of some intermediate in glycolysis withcarbon dioxide.