Citation: Busont, O.; Durambur, G.;
Bernard, S.; Plasson, C.; Joudiou, C.;
Baude, L.; Chefdor, F.; Depierreux, C.;
Héricourt, F.; Larcher, M.; et al. Black
Poplar (Populus nigra L.) Root
Extracellular Trap, Structural and
Molecular Remodeling in Response
to Osmotic Stress. Cells 2023, 12, 858.
https://doi.org/10.3390/cells12060858
Academic Editor: Andrei Smertenko
Received: 16 February 2023
Revised: 4 March 2023
Accepted: 6 March 2023
Published: 9 March 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
cells
Article
Black Poplar (Populus nigra L.) Root Extracellular Trap, Structural
and Molecular Remodeling in Response to Osmotic Stress
Océane Busont
1
, Gaëlle Durambur
2
, Sophie Bernard
2,3
, Carole Plasson
2
, Camille Joudiou
1
, Laura Baude
1,4
,
Françoise Chefdor
1
, Christiane Depierreux
1
, François Héricourt
1
,Mélanie Larcher
1
, Sonia Malik
1
,
Isabelle Boulogne
2
, Azeddine Driouich
2
, Sabine Carpin
1
and Frédéric Lamblin
1,
*
1
Laboratoire de Biologie des Ligneux et des Grandes Cultures, Université d’Orléans, INRAE, USC 1328,
CEDEX 2, F-45067 Orléans, France
2
GLYCOMEV UR 4358, SFR Normandie Végétal FED 4277, Innovation Chimie Carnot, University of Rouen
Normandie, IRIB, F-76000 Rouen, France
3
INSERM, CNRS, HeRacLeS US 51 UAR 2026, PRIMACEN, University of Rouen Normandie,
F-76000 Rouen, France
4
Department of Biology, University of Fribourg, CH-1700 Fribourg, Switzerland
* Correspondence: frederic.lamblin@univ-orleans.fr; Tel.: +33-(0)2-3841-7127
Abstract: The root extracellular trap (RET) consists of root-associated, cap-derived cells (root AC-DCs)
and their mucilaginous secretions, and forms a structure around the root tip that protects against
biotic and abiotic stresses. However, there is little information concerning the changes undergone
by the RET during droughts, especially for tree species. Morphological and immunocytochem-
ical approaches were used to study the RET of black poplar (Populus nigra L.) seedlings grown
in vitro under optimal conditions (on agar-gelled medium) or when polyethylene glycol-mediated
(PEG
6000
—infused agar-gelled medium) was used to mimic drought conditions through osmotic
stress. Under optimal conditions, the root cap released three populations of individual AC-DC
morphotypes, with a very low proportion of spherical morphotypes, and equivalent proportions of
intermediate and elongated morphotypes. Immunolabeling experiments using anti-glycan antibodies
specific to cell wall polysaccharide and arabinogalactan protein (AGP) epitopes revealed the presence
of homogalacturonan (HG), galactan chains of rhamnogalacturonan-I (RG-I), and AGPs in root AC-
DC cell walls. The data also showed the presence of xylogalacturonan (XGA), xylan, AGPs, and low
levels of arabinans in the mucilage. The findings also showed that under osmotic stress conditions,
both the number of AC-DCs (spherical and intermediate morphotypes) and the total quantity of
mucilage per root tip increased, whereas the mucilage was devoid of the epitopes associated with the
polysaccharides RG-I, XGA, xylan, and AGPs. Osmotic stress also led to reduced root growth and
increased root expression of the P5CS2 gene, which is involved in proline biosynthesis and cellular
osmolarity maintenance (or preservation) in aerial parts. Together, our findings show that the RET is
a dynamic structure that undergoes pronounced structural and molecular remodeling, which might
contribute to the survival of the root tip under osmotic conditions.
Keywords: AC-DCs; mucilage; drought stress; root cap; xylogalacturonan; xylan; arabinogalactan
protein; rhamnogalacturonan-I
1. Introduction
Drought is one of the most severe environmental stresses affecting plant growth
and productivity [1]. The black poplar (Populus nigra L.) is an emblematic alluvial forest
species that grows along large rivers with considerable seasonal flow variation. The
tree is regularly exposed to periods of flooding, followed by periods of increasingly low
water levels and thereby drought conditions. In the context of global warming, wild and
cultivated poplars are being subjected to increasingly prolonged and intense periods of
drought. The restoration and protection of riparian forests is one of the key priorities in
Cells 2023, 12, 858. https://doi.org/10.3390/cells12060858 https://www.mdpi.com/journal/cells