title: Fern Stomatal Responses to ABA and CO2Depend on Species and Growth Conditions.
abstract: Changing atmospheric CO2levels, climate, and air humidity affect plant gas exchange that is controlled by stomata, small pores on plant leaves and stems formed by guard cells. Evolution has shaped the morphology and regulatory mechanisms governing stomatal movements to correspond to the needs of various land plant groups over the past 400 million years. Stomata close in response to the plant hormone abscisic acid (ABA), elevated CO2concentration, and reduced air humidity. Whether the active regulatory mechanisms that control stomatal closure in response to these stimuli are present already in mosses, the oldest plant group with stomata, or were acquired more recently in angiosperms remains controversial. It has been suggested that the stomata of the basal vascular plants, such as ferns and lycophytes, close solely hydropassively. On the other hand, active stomatal closure in response to ABA and CO2was found in several moss, lycophyte, and fern species. Here, we show that the stomata of two temperate fern species respond to ABA and CO2and that an active mechanism of stomatal regulation in response to reduced air humidity is present in some ferns. Importantly, fern stomatal responses depend on growth conditions. The data indicate that the stomatal behavior of ferns is more complex than anticipated before, and active stomatal regulation is present in some ferns and has possibly been lost in others. Further analysis that takes into account fern species, life history, evolutionary age, and growth conditions is required to gain insight into the evolution of land plant stomatal responses.
authors.affiliation: Plant Signal Research Group, University of Tartu, Institute of Technology, Tartu 50411, Estonia.
authors.forename: Hanna
authors.identifier: http://orcid.org/0000-0002-6392-859X
authors.initials: H
authors.lastname: Hõrak
authors.affiliation: Plant Signal Research Group, University of Tartu, Institute of Technology, Tartu 50411, Estonia.
authors.forename: Hannes
authors.identifier: http://orcid.org/0000-0002-6895-3583
authors.initials: H
authors.lastname: Kollist
authors.affiliation: Plant Signal Research Group, University of Tartu, Institute of Technology, Tartu 50411, Estonia ebe.merilo@ut.ee.
authors.forename: Ebe
authors.identifier: http://orcid.org/0000-0003-1767-6413
authors.initials: E
authors.lastname: Merilo
mesh_terms: D000040:Abscisic Acid / Q000378:metabolism / Q000494:pharmacology; D002245:Carbon Dioxide / Q000378:metabolism*; D029624:Ferns / Q000187:drug effects / Q000254:growth & development / Q000502:physiology*; D006813:Humidity; D054046:Plant Stomata / Q000502:physiology*; D013045:Species Specificity; D055548:Vapor Pressure
chemical_list: D002245:Carbon Dioxide; D000040:Abscisic Acid