Vol.:(0123456789) 1 3
Trees
https://doi.org/10.1007/s00468-020-01992-x
ORIGINAL ARTICLE
Stomatal features and desiccation responses of Persian walnut leaf
as caused by in vitro stimuli aimed at stomatal closure
Sasan Aliniaeifard
1
· Zeinab Maleki Asayesh
1
· John Driver
2
· Kourosh Vahdati
1
Received: 12 August 2019 / Accepted: 20 May 2020
© Springer-Verlag GmbH Germany, part of Springer Nature 2020
Abstract
Key message Functional stomata can be generated by in vitro ABA applications. This mitigates the desiccation
responses of Persian walnut leaves in their early stage of ex vitro exposure. This fnding paves the way for discovering
a solution for the uncontrollable water loss problem of in vitro-generated walnut plantlets.
Abstract Most in vitro-generated plants are susceptible to wilting when they are transferred to an ex vitro environment,
which is characterized by a higher evaporative nature. In this study, two stomatal closing stimuli [abscisic acid (ABA) and
polyethylene glycol (PEG)] were added to in vitro culture medium of Persian walnut (Juglans regia L.) to evaluate their leaf
ability to control water loss during exposure to an ex vitro environment. Diferent concentrations of PEG (0, 3 and 5%) and
ABA (0, 1 and 10 µM) were tested, and their efects were evaluated in two separate experiments. In the frst experiment, the
proline concentration decreased in the leaves grown on media containing PEG. In contrast, glycine betaine (GB) increased
only in leaves grown on a culture medium with 5% PEG. Leaf osmotic potential (ψ
s
) increased parallel to the increase in
PEG concentration. Adding PEG to the culture medium caused a decrease in stomatal aperture, and, as a result, the water
conservation capacity of the in vitro-grown leaves was increased during detached leaf ex vitro desiccation. However, PEG-
induced osmotic stress caused a negative infuence on the morpho-physiologic characteristics and chlorophyll content of
in vitro-grown nodal segments. In the second experiment, ABA was tested in the culture medium with the eventual aim
of making in vitro leaves lose less water when they were transferred to an ex vitro environment. ABA (especially 10 µM)
improved the leaf water conservation capacity during ex vitro desiccation of detached leaves. Small stomata having narrow
apertures were increased in number by supplementing the cultures with ABA, although the length of shoots decreased with
its use; no serious side efect was observed on morpho-physiologic parameters. Our fndings are signifcant because they can
ultimately be useful for preventing the uncontrollable water loss problem of in vitro-generated walnut plantlets.
Keywords Abscisic acid (ABA) · Tissue culture · Polyethylene glycol (PEG) · Stomata · Acclimatization
Introduction
The acclimation of micropropagated plantlets to ex vitro
conditions involves exposing them to environmental con-
ditions that are very diferent from those in their in vitro
environment. Plantlets naturally fail to be established suc-
cessfully after shifting from an in vitro environment to an
ex vitro one because the evaporative demand is higher in
the latter. Therefore, a period of acclimatization to the ex
vitro environment is necessary for the successful establish-
ment and survival of plantlets (Chandra et al. 2010). Aseptic
culturing media and unusual environmental conditions such
as light, temperature, nutrient levels as well as the gase-
ous composition [e.g., CO
2
, ethylene and relative humid-
ity (RH)] in the in vitro media and vessel head space can
Communicated by S.A. Merkle.
Electronic supplementary material The online version of this
article (https://doi.org/10.1007/s00468-020-01992-x) contains
supplementary material, which is available to authorized users.
* Kourosh Vahdati
kvahdati@ut.ac.ir
1
Department of Horticulture, College of Aburaihan,
University of Tehran, 3391653755 Tehran, Iran
2
Driver Consulting, Inc, Waterford, CA, USA