Contents lists available at ScienceDirect
Industrial Crops & Products
journal homepage: www.elsevier.com/locate/indcrop
Resin exudation profile, chemical composition, and secretory canal
characterization in contrasting yield phenotypes of Pinus elliottii Engelm
Franciele A. Neis, Fernanda de Costa, Márcia Rodrigues de Almeida, Luana Caroline Colling,
Camila Fernanda de Oliveira Junkes, Janette Palma Fett, Arthur G. Fett-Neto
⁎
Center for Biotechnology and Department of Botany, Federal University of Rio Grande do Sul (UFRGS), C.P. 15005, CEP 91501-970, Porto Alegre, RS, Brazil
ARTICLE INFO
Keywords:
Pinus
resin canals
tapping
resin composition
high yield
microCT
ABSTRACT
In conifer stems, secretory canals synthesize and store resin for defense against herbivores and pathogens. Resin
terpenes are used as raw material by an array of industrial sectors. Most forest stands operationally used in resin
extraction are derived from seeds, showing high genetic variation, which reflects in yield. The objective of this
study was to identify adult slash pine (Pinus elliottii Engelm.) trees of high yield of resin in a short timeframe by
resin mass flow rate analysis, aiming at the establishment of elite forests for tapping prior to its start. In addition,
the anatomical basis of resin yield was investigated by examining the correlation between parameters such as
number, shape, area and internal volume of wood canals with resin production. Monoterpene composition in
resin of high and low yielding trees was also compared. The resin flow-based selection method was reliable for
resin yield phenotype detection, confirming this property in trees formerly identified as being of high and low
resin production by conventional tapping. The reverse test for identification of high and low yield resin features
in previously untapped younger plants was in good agreement with their yields after subsequent standard
tapping procedure. To evaluate and quantify the three-dimensional structure of resin canals, we used microCT
scans. High yielding trees had more axial resin canals when compared with low yielding ones. Frequency of
putative anastomosed canals and canal diameter were also superior in the former. Chemical analyses of resin
monoterpenes revealed that the ratio of α-pinene/ β-pinene was lower in more productive trees, which also had
more limonene in total terpenes compared with their low yield counterparts. Data support the use of short-term
resin mass flow rate analysis as a tool to identify and select high yield trees for the establishment of elite slash
pine forests for resin tapping operations. Strong correlation of the superesinous phenotype with canal density
and structure was also evident.
1. Introduction
Conifers have developed a series of adaptive strategies to deal with
herbivore and pathogen attacks (Franceschi et al., 2005; Keeling and
Bohlmann, 2006; Geisler and Jensen, 2016). Resin is considered the
major defense of conifers, and its composition consists of various ter-
penoids such as monoterpenes, sesquiterpenes, and diterpenes (Phillips
and Croteau, 1999; Martin et al., 2002; Zulak and Bohlmann, 2010).
Resin is synthesized and accumulated in specialized secretory
structures (isolated resin cells, multicellular resin blisters and net-
worked resin canals), which may appear as a normal feature of devel-
opment in tissues (constitutive defense) or may result by the induction
of external factors (Bannan, 1936; Lewinsohn et al., 1994; Wu and Hu,
1997; Hudgins et al., 2003; Langenheim, 2003). A commonly induced
response to mechanical damage, insect attack, fungal invasion,
application of hormones and chemical stimulants is the production of
traumatic resin canals in the xylem (Lombardero et al., 2000; Nagy
et al., 2000; Franceschi et al., 2002; Arbellay et al., 2014). The for-
mation of traumatic resin canals represents an important induced de-
fense that enhances resin production and flow in response to environ-
mental perturbations in tissues close to the wounded zone (Franceschi
et al., 2005; DeRose et al., 2017).
Resin canals of Pinaceae are differentiated into radial canals and
axial canals, depending on their orientation, creating a complex net-
work (Bannan, 1936; Lewinsohn et al., 1991; Rodríguez-García et al.,
2014). Resin flow can be influenced by an array of factors such as ir-
radiance, temperature, season, and edaphic conditions, as well as by
genetics, age, and wounding (Peñuelas and Llusià, 1999; Ayres and
Lombardero, 2000; Knebel et al., 2008; Rodrigues and Fett-Neto, 2009;
Hood and Sala, 2015; Neis et al., 2018). Selection of high resin yield
https://doi.org/10.1016/j.indcrop.2019.02.013
Received 24 August 2018; Received in revised form 27 December 2018; Accepted 7 February 2019
⁎
Corresponding author.
E-mail address: fettneto@cbiot.ufrgs.br (A.G. Fett-Neto).
Industrial Crops & Products 132 (2019) 76–83
0926-6690/ © 2019 Elsevier B.V. All rights reserved.
T