Interxylary phloem and xylem rays are the

structural foundation of agarwood resin

formation in the stems of Aquilaria sinensis

Peiwei Liu, Xingli Zhang, Yun Yang,

Chun Sui, Yanhong Xu & Jianhe Wei

Trees
Structure and Function

ISSN 0931-1890

Trees
DOI 10.1007/s00468-018-1799-4

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https://doi.org/10.1007/s00468-018-1799-4

ORIGINAL ARTICLE

Interxylary phloem and xylem rays are the structural foundation

of agarwood resin formation in the stems of Aquilaria sinensis
Peiwei Liu1,2   · Xingli Zhang2,3 · Yun Yang1 · Chun Sui2 · Yanhong Xu2 · Jianhe Wei1,2

Received: 25 September 2018 / Accepted: 11 December 2018

© Springer-Verlag GmbH Germany, part of Springer Nature 2018

Abstract
Key message  The formation of resin in the wood of A. sinensis was observed, and the results showed that the interxy-
lary phloem, together with xylem rays, provided a structural foundation for the formation of agarwood resin.
Abstract  Agarwood is a costly resinous wood harvested from wounded Aquilaria trees and has been widely used in medi-
cine, incense, and perfumery. A defensive response of Aquilaria to various wounds has been shown to be the key reason
for agarwood formation; however, our understanding of the anatomical basis of agarwood formation is still fragmentary. In
this study, we examined the structural characteristics of A. sinensis wood and its relationship with agarwood formation. The
results showed that interxylary phloem together with xylem rays is the main tissue that contains living parenchyma cells in
the wood of healthy A. sinensis and that the main reserve substance in these parenchyma cells is in the form of starch grains.
After Agar-Wit treatment, these starch grains undergo a series of changes and are eventually converted into agarwood resin;
the non-starch polysaccharides and phenols are some of the intermediate products in the process of agarwood resin forma-
tion. The resin initially forms and mainly accumulates in the parenchyma cells of the interxylary phloem and xylem rays.
The results indicated that the interxylary phloem and xylem rays in the wood are not only the primary location of agarwood
resin formation but also the main accumulation site. The stored starch grains might be the material underlying the formation
of the resin.

Keywords  Aquilaria sinensis · Agar-Wit · Interxylary phloem · Xylem ray · Agarwood resin

Introduction

Agarwood or eaglewood is the most expensive wood in the

Communicated by Sano.
* Jianhe Wei
wjianh@263.net
1
Key Laboratory of State Administration of Traditional
Chinese Medicine for Agarwood Sustainable Utilization,
Hainan Branch of the Institute of Medicinal Plant
Development, Chinese Academy of Medical Sciences
and Peking Union Medical College, Haikou 570311, China
2
Key Laboratory of Bioactive Substances and Resources
Utilization of Chinese Herbal Medicine, Ministry
of Education & National Engineering Laboratory
for Breeding of Endangered Medicinal Materials, Institute
of Medicinal Plant Development, Chinese Academy
of Medical Sciences and Peking Union Medical College,
Beijing 100193, China
3
Qilu University of Technology (Shandong Academy
of Sciences), Jinan 250353, China
world with diverse applications in medicine, perfumery,
incense, and ornamentation across the Middle East, South
Asia, Japan and China (Xu et al. 2016; Elias et al. 2017).
As the main source of agarwood, Aquilaria trees have high
economic value; therefore, its natural resources have been
severely depleted. All Aquilaria species are listed in Appen-
dix II of the Convention on International Trade in Endan-
gered Species of Wild Fauna and Flora (Cites 2004).
Because only wounded trunks or branches of Aquilaria
trees can produce agarwood, people have developed vari-
ous techniques for producing agarwood, such as partly
breaking the trunk, making holes by drilling and inocu-
lating with fungi (Guangdong Institute of Plant Research
1976; Nobuchi and Siripatanadilok 1991; Pojanagaroon
and Kaewrak 2005; Faizal et al. 2016); however, these
methods do not have predictable results due to a lack of
understanding of the plant structure and response system

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(Chong et al. 2014). Based on our current understanding
of the mechanism of agarwood formation (Zhang et al.
2010; Zhang 2013; Gao and Wei 2016), we invented an
efficient technique for producing agarwood, called the
whole-tree agarwood-inducing technique (Agar-Wit), in
which the agarwood inducer is transported throughout
the Aquilaria tree, creating wounds within the whole
tree. Agarwood forms around wounds over a period of
several months (Zhang et al. 2012a; Liu et al. 2013). The
agarwood induced over 6 months by this technique in A.
sinensis meets the requirements of the Chinese Pharmaco-
poeia, and the yield is 4–28 times higher than that obtained
by conventional agarwood-inducing methods (Liu et al.
2013). For these reasons, this efficient technique has been
widely used worldwide.
Considering the high value of agarwood, several groups
have focused on the anatomical basis of agarwood for-
mation, which can help in agarwood sourcing. Previous
reports have shown that the most obvious feature of the
wood of Aquilaria is the interxylary phloem (IP) (Rao
and Dayal 1992; Nobuchi and Siripatanadilok 1991; Zhang
et al. 2012b). But the exact role of the IP in agarwood for-
mation is still not clear. In addition, there are distinct ideas
about agarwood resin formation. Nobuchi and Siripatan-
adilok (1991) believed that agarwood resin is synthesized
by living parenchyma cells of Aquilaria, but Adams et al.
(2016) concluded that the resin is the product of fungi that
uses Aquilaria wood cells as raw material. Considering
the problems above, the process of agarwood formation
induced by Agar-Wit at the anatomical level was studied.
This information will contribute to clarify the anatomi-
cal basis of agarwood formation and may help for further
improve techniques for producing agarwood.
Materials and methods
Plant materials and artificial agarwood induction
The experiment was carried out in Xinglong county, Wan-
ning city in Hainan Province, China (18°44′N,110°12′E).
Twenty-seven 3-year-old healthy A. sinensis trees with
uniform base diameters (3 ± 0.3 cm) were used. Accord-
ing to the Agar-Wit technique (Liu et al. 2013), a 50 mL
agarwood inducer was injected into the xylem tissues
through a small hole with a diameter 5 mm, 20 cm above
the ground. After 5  days, 10  days, 15  days, 1  month,
2 months, 3 months, 5 months, and 12 months, 2-cm thick
wood slices were collected from 20 cm above the hole.
Three trees were repeated in each time. Three healthy A.
sinensis tree served as the controls.
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Scanning electron microscope (SEM)
The wood slices were cut into blocks (2–3 mm3) with a
razor blade to expose the transverse, radial or tangential
surface. The blocks were fixed in 4% paraformaldehyde for
4 h and then washed three times by 0.1 mol/L phosphate
buffer solution(PH = 7.2). Finally the blocks were air-
dried for approximately 7 days at 37 °C, and then observed
under an SEM (JSM 6510LV, Japan) after gold coating.
Histochemical staining
The wood slices were cut into blocks (2 × 1 × 1 cm3) and
immediately fixed in 4% paraformaldehyde. Then, 60-µm
thick sections were sliced from the blocks using a freezing
microtome (Leica CM1900, Germany). Five stains were
used according to the method of Soukup (2005) and Li
(1996). The sections were stained with iodine–potassium
iodide ­(I2–KI) to detect starch grains, whereas polysaccha-
ride compounds were detected by staining with Periodic
acid-Schiff (PAS) reagent. The sections were stained with
Sudan Black B to detect the lipid compounds, whereas
the phenolic compounds were detected using Millon’s
reagent. To detect living cells, the sections were stained
with 4,6-diamidino-2-phenylindole (DAPI). All sections
were observed under a Nikon 80i light microscope (Nikon,
Japan).
Semi‑thin and ultra‑thin sections
The wood slices were cut into blocks (1 mm 3) and fixed
in 3% paraformaldehyde plus 2% glutaraldehyde for 12 h
at 4 °C, followed by three washes in PBS. The specimens
were post-fixed in 1% osmium tetroxide for 8 h at room
temperature followed by three washes, and then dehy-
drated using an ethanol gradient and embedded in Epon
812 (SPI Supplies, USA). Polymerization was conducted
at 35 °C, 40 °C, and 60 °C for 24 h each. Subsequently,
the specimens were sliced into 2-µm sections using a ultra-
microtome (Leica EM UC7, Germany). After staining
with Toluidine Blue O or PAS or Sudan black B, the sec-
tions were observed under a Nikon 80i light microscope.
According to the results of the semi-thin sections, ultrathin
sections (90 nm) were cut using diamond knives (DiA-
TOME, Switzerland), stained with lead citrate and uranyl
acetate, and observed by transmission electron microscopy
(FEI Tecnai G2, USA).
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Three‑dimensional structure of the wood
A three-dimensional structure of A. sinensis wood was
created using Adobe Illustrator CS5 based on the images
and statistical data.
Results
Structural characteristics of healthy A. sinensis wood
Similar to other species of Aquilaria, A. sinensis wood is
mainly composed of four parts: interxylary phloem (IP),
xylem rays (XR), vessels, and wood fibers (Fig. 1a, 2). The
IP is a typical anomalous tissue that is island-shaped and
evenly distributed in A. sinensis wood (Fig. 1a, b). Fully dif-
ferentiated IP may consist the following typical structures:
Fig. 1  Structural characteristics of healthy A. sinensis wood. a
Transverse section showing island-shaped IP and multiple radial
vessels(V). b Transverse section showing phloem fibers (PF), phloem
rays (PR) and crystal (C) in IP. c Tangential section showing stripped
IP, fusiform XR and vessels(V). d, e Tangential sections show-
parenchyma cells, sieve tubes, phloem rays, and phloem fib-
ers (Figs. 1b, 6a). The outer region of the phloem is com-
prised of 1–4 layers of larger polygonal parenchyma cells
known as the connective tissue (Fig. 6a). The thin-walled
cells that connect with some of the xylem rays are the interx-
ylary phloem rays (Figs. 1b, f and 3b). The xylem rays and
interxylary phloem rays represent the lateral conductive tis-
sue in wood, and the cell wall of the latter is thinner and
unlignified compared to the former (Fig. 1c–e). In addition
to starch grains (Fig. 1d, e), crystals were also observed in
the thin-walled cells described above (Fig. 1b). Addition-
ally, thick-walled fibers in the inner region of the IP were
observed (Figs. 1b, 3a, 6b). Moreover, the inner regions of
the IP also contained several sieve tubes (Fig. 2).
Safranin and fast green staining showed that the IP and
XR were the major tissues containing parenchyma cells in A.
sinensis wood (Fig. 3a, b). DAPI staining revealed that these
ing parenchyma cell with starch grains (S) in IP and fusiform XR. f
Radial section showing XR connected with IP. C crystal, IP interxy-
lary phloem, PF phloem fiber, PR phloem ray, S starch grain, V ves-
sel, WF wood fiber, XR xylem ray. Scale bar = 500 µm in a; 100 µm
in b, c, e, f; and 10 µm in d 
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Fig. 2  3D reconstruction of the
wood
parenchyma cells were alive based on the positive staining
in the nucleus (Fig. 3c); SEM and ­I2–KI staining revealed
that there were many starch grains contained within these
parenchyma cells (Figs. 1d, e, 3d, e). PAS staining showed
that some parenchyma cells also contained non-starch pol-
ysaccharides (Fig. 3f). In the transverse section, approxi-
mately 3–7 phloem rays were connected with XR in one IP
(Figs. 1b, 3d). In the radial section, the IP was strip-shaped
and intersected by numerous ribbon-shaped XR (Figs. 1f,
3b, e). In other words, IP and XR combined organically to
form a living parenchyma cell network in the wood.
Stratified characteristics of A. sinensis wood
after Agar‑Wit treatment
Healthy A. sinensis wood is yellowish-white. After Agar-Wit
treatment, part of the wood in the centre of A. sinensis stems
is altered significantly, with the degree of discoloration pro-
portional to the treatment time (Fig. 4a). The wood can be
divided into four layers from the inside to the outside based
on colour and relative position: decayed layer, agarwood
layer, transitional layer, and healthy layer (Fig. 4b).
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The structural features of each layer were observed. In the
decayed layer, the IP was almost destroyed; often, only holes
with some mycelia were observed in the original locations
(Fig. 4c, d). In the agarwood layer, the IP and XR, as well
as the vessels and wood fibers, were filled with brownish or
dark-brown resin (Fig. 4d). In the transitional layer, although
no resin was visible in the IP, occlusion was found in some
vessels (Fig. 4d). The healthy layer showed a structure simi-
lar to that of healthy A. sinensis wood (Figs. 1a, 4d).
Histological formation process of agarwood resin
in the agarwood layer
Agarwood resin mainly exists in the agarwood layer; there-
fore, histological changes in the agarwood layer were inves-
tigated. No brown substance was detected in healthy A. sin-
ensis wood (Fig. 5a). After 15 days of Agar-Wit treatment,
a small amount of yellowish substance began to appear in
some parts of the interconnected phloem rays and xylem rays
(Fig. 5b). After 30 days of treatment, a significant increase
was observed in the amount of yellowish substance in the
IP and XR (Fig. 5c). With prolonged treatment time, the IP
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Fig. 3  Histochemical staining of healthy A. sinensis wood. a, b Safra-
nin and fast green stains showing IP and XR are the main tissues that
contain living parenchyma cells and their connected network. c DAPI
staining showed bright blue fluorescence (arrow) in the nuclei of the
parenchyma cells in IP and XR. d, e ­I2–KI staining showed starch
and XR in the agarwood layer were filled with an abundance
of the yellowish substance, and the surrounding vessels and
fibers were filled with the brown substance (Fig. 5d). After
1 year of Agar-Wit treatment, the colour and lustre of the
brown substance (Fig. 5e) were similar to those of high-
quality agarwood; hence, it was considered agarwood resin.
Transformation of substances during the formation
of agarwood
Some histochemical tests were used to follow substance
changes during the formation of agarwood resin. The first
and most typical change after Agar-Wit treatment was a sig-
nificant reduction in the abundance of starch grains (Fig. 6a,
b). After 15  days, the starch grains nearly disappeared
(Fig.  6b). Moreover, a new substance appeared, which
reacted with PAS, resulting in a burgundy colour, indicating
grains in the parenchyma cells of IP and XR. f PAS staining showed
starch grains (S) and non-starch polysaccharides (NP) in the cross
section of IP parenchyma cells. Scale bar = 100 µm in a–e and 20 µm
in f 
that it might be non-starch polysaccharide (Fig. 6d). The
amount of this substance in the IP and XR increased gradu-
ally and was highest 60 days after Agar-Wit (Fig. 6e). Sub-
sequently, this substance reacted with PAS and became less
abundant until it finally was no longer detectable (Fig. 6f).
Another new substance appeared in the IP and XR paren-
chyma cells after Agar-Wit. This substance exhibited yel-
low autofluorescence and reacted with Millon’s reagent,
resulting in a dark red colour (Fig. 5m, n), indicating the
possible presence of phenols. After 30 days of Agar-Wit,
this substance began to appear in the IP and XR (Fig. 5g)
and gradually increased to a maximum level approximately
3 months after Agar-Wit (Fig. 5h). The level of this sub-
stance by gradually decreased until it was no longer detect-
able (Fig. 5i, j).
Unlike the above substances, one substance increased
continuously after Agar-Wit. This substance might have
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Fig. 4  Stratified structures of
A. sinensis after Agar-Wit treat-
ment. a A. sinensis stems after
15 (upper figure) and 90 days
(lower figure) of treatment; the
arrow points to the agarwood
layer. b Transversal surface
of the A. sinensis stem after
150 days of treatment, where
the layers from the inside to
outside are the decayed layer
(DL), agarwood layer (AL),
transitional layer (TL), and
healthy layer (HL). c Void IP
with mycelia (long arrow) in
the decayed layer. d Transverse
section showing the structural
features of the above four layers.
DL decayed layer, AL agarwood
layer, TL transitional layer,
HL healthy layer. Scale bar
= 400 µm in b, 100 µm in c and
200 µm in d 

been a lipid because it reacted with Sultan black B, resulting
in a black colour. After 15 days of treatment, a small amount
of lipids appeared in the corner of parenchyma cells in the
IP and XR (Fig. 6g). After 60 days of treatment, additional
lipids appeared in the centre of the parenchyma cells in the
IP and XR (Fig. 6h). After 1 year of treatment, the IP and
XR were completely filled with lipids (Figs. 5o, 6i).
It should be noted that the substance changes above
mainly occurred in the agarwood layer around the decayed
layer. In fact, the substance in one agarwood layer also
showed a similar change pattern. For example, in the 1-year-
old agarwood layer, the IP, XR, and some vessels surround-
ing the decayed layer were dominated by lipids (Fig. 5o);
however, polysaccharides (Fig. 5l) or phenols (Fig. 5j) were
predominant around the transition layer.
To further observe the above transformation of sub-
stances, transmission electron microscopy on the paren-
chyma cells in IP was carried out during the early forma-
tion of agarwood. In the healthy trees, a great deal of starch
grains were discovered in the parenchyma cells of the IP
(Fig. 7a). After 30 days of Agar-Wit treatment, these starch
grains disappeared and cytoplasm increased. At the same
time, many mitochondria (Fig. 7b), plastids (Fig. 7c) and
black osmiophilic substances were observed in these paren-
chyma cells.(Fig. 7c). After 60 days of treatment, black
osmiophilic droplets appeared in the plastids (Fig. 7d). In
addition, many grey osmium substances first appeared in
small vacuoles (Fig. 7e) and then accumulated in the large
central vacuole(Fig. 7f).
Discussion
The xylem structure of Aquilaria is the most important part
to study because it is the basis of agarwood resin formation
and accumulation (Chong et al. 2014). The wood structure
has been reported for A. sinensis (Zhang et al. 2012b), A.
crassna (Nobuchi and Siripatanadilok 1991), A.agallocha
(Rao and Dayal 1992), A.malaccensis (Adams et al. 2016;
Mohamed et al. 2013), and A. beccariana (Wheeler 2011;
Insidewood 2004). These results suggested that the wood
structures of Aquilaria trees are similar, and the most obvi-
ous feature is the IP. Although the physiological role of the
anomalous tissue in plants is not clear (Robert et al. 2011;
Carlquist 2013), the present study indicate that the IP plays
an important role in agarwood formation.

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Fig. 5  Histological observation of the formation of agarwood resin
after the Agar-Wit treatment. a–e Natural coloration of agarwood
showing the formation of a yellowish substance. a No brown resin
was visible in healthy A. sinensis. b After 15  days of treatment, a
small amount of yellowish substance began to appear in the IP and
XR. c After 30 days of treatment, yellowish substance (arrow) clearly
appeared in the IP and XR. d More brownish yellow substance was
found in the IP and XR and was also visible in some of the sur-
rounding vessels (arrow) after 90 days of treatment. e After 1 year of
treatment, the IP and XR, as well as some of the surrounding ves-
sels and wood fibers, were filled with the brown substance. f–j The
autofluorescence of agarwood layer under 450–490  nm excitation. f
No autofluorescence in the parenchyma cells of the IP in healthy A.
sinensis. g After 30  days of treatment, autofluorescence began to
IP and XR may be not only the primary location of agar-
wood resin formation but also the main accumulation site.
Following evidences are advanced in support of this per-
spective. First, several studies have reported that the liv-
ing parenchyma cells in Aquilaria wood are the only cells
appear in the IP and XR. h After 90  days of treatment, autofluores-
cence was higher in the IP and XR. i After 150  days of treatment,
autofluorescence began to decrease in the IP and XR (white arrow)
but appeared in some vessels and fibers (black arrow). j After 1 year
of treatment, almost no autofluorescence was observed near the
decayed layer (white arrow); however, bright autofluorescence (black
arrow) was detected near the transition layer. k ­I2–KI staining showed
the remaining starch grains (arrow) in IP after 10 days of treatment.
l PAS staining showed polysaccharides in some IP (arrow) near the
transition layer. m, n Millon’s reagent showed phenols in the ves-
sel (V) and parenchyma cells (arrow) after 150  days of treatment. o
Sudan black B staining showed lipid in IP, XR and some vessels after
1 year of treatment. Scale bar = 100 µm
that can synthesize agarwood resin (Mohamed et al. 2013;
Nobuchi and Siripatanadilok 1991). The present study fur-
ther shows that the living parenchyma cells in the wood of
A. sinensis almost distribute in the IP and XR, and these two
tissues combine organically to form a living cell network
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Fig. 6  Histochemical observation of the formation of agarwood resin
after the Agar-Wit treatment. a–c Toluidine blue staining. a After
5  days of treatment, the number of starch grains (S) in the paren-
chyma cells of the interxylary phloem (IP) and xylem rays (XR) was
significantly lower. b After 15  days of treatment, the starch grains
in the parenchyma cells disappeared, and a small amount of dark-
coloured substances appeared in the intersection of the parenchyma
cells in the XR and phloem ray (PR). c After 30 days of treatment, a
large volume of the darkly stained substance (arrow) was found in the
parenchyma cells and the surrounding fibers. d–f PAS staining. Red
coloration is indicative of polysaccharides. d After 15  days of treat-
ment, a small amount of non-starch polysaccharides (NP) appeared in
in the wood. Second, many reports have shown that the
brownish agarwood resin is concentrated mostly in the IP
and XR, lesser in the vessels and fibers (Rao and Dayal
1992; Mohamed et al. 2013; Faizal et al. 2016; Adams et al.
2016). The present study and other studies further found that
the agarwood resin is initially formed in the parenchyma
cells of the IP and XR and then plugged to the surrounding
vessels and fibers (Guangdong Institute of Plant Research
1976; Nobuchi and Siripatanadilok 1991; Rao and Dayal
1992). Importantly, in the early process of agarwood for-
mation, many mitochondria and plastids in the parenchyma
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the parenchyma cells in the XR and part of the IP. e After 60 days of
treatment, a large amount of NP appeared in the IP and XR. f After
150  days of treatment, the level of polysaccharides in the IP was
reduced significantly. g–i Sudan black B staining. Black coloration
indicates lipids. g After 15 days of treatment, a small amount of lipids
(L) appeared in the corners of parenchyma cells (arrow) and some
vessels (V). h After 60 days of treatment, a large amount of lipids (L)
appeared in the parenchyma cells in the IP and XR and in part of the
surrounding wood fibers. i After 1 year of treatment, the IP and XR
were filled with lipids. NP non-starch polysaccharides, L lipids. Scale
bars = 50 µm
cells were observed and we believe these organelles may
provide energy and sites for agarwood resin formation.
Meanwhile, in these parenchyma cells, we also observed
three kinds of osmiophilic substances which are considered
to be closely related with the agarwood formation (Nobuchi
and Siripatanadilok 1991). These results imply that IP and
XR may provide site for the agarwood resin formation and
accumulation.
The starch grains in the parenchyma cells of IP and XR
may provide raw material for the formation of agarwood
resin. Firstly, the parenchyma cells in the IP and XR of
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Fig. 7  Cytological observation of the parenchyma cells in the IP after
the Agar-Wit treatment. a Starch grains(S) in the parenchyma cells
in the unwounded IP. b After 30  days of treatment, the parenchyma
cells contained dense cytoplasm and more mitochondria (M). c After
30  days of treatment, black osmium substances (arrow) appeared
in cytoplasm. d After 60  days of treatment, black osmium droplets
healthy Aquilaria trees contained abundant starch grains
(Rao and Dayal 1992; Nobuchi and Siripatanadilok 1991;
Zhang et al. 2012b). After wounding or applying the Agar-
Wit treatment, these starch grains were gradually hydrolysed
and eventually disappeared (Guangdong Institute of Plant
Research 1976; Nobuchi and Siripatanadilok 1991). Adams
(2016) pointed out that the metabolic byproducts of starch
hydrolysis may be involved in agarwood resin synthesis
(Adams et al. 2016). On the other hand, Nobuchi (1991)
detected many phenolic compounds and a small amount
of lipids in A. crassna wood after the starch grains disap-
peared (Nobuchi and Siripatanadilok 1991). The present
study showed that after the starch grains were degraded,
non-starch polysaccharides initially appeared and then grad-
ually vanished, with a concomitant increase in the amount
of phenolic substances. Afterwards, the amount of phenolic
substances gradually reduced and finally, many lipid sub-
stances were detected. These results imply that the proper-
ties of the resin gradually changed during the process of
(arrow) were formed in the plastids. e After 60  days of treatment,
grey osmiophilic substance (arrow) accumulated in the vacuoles (Va).
f After 60 days of treatment, osmiophilic substance (arrow) accumu-
lated in a large vacuole(Va). ER endoplasmic reticulum, M mitochon-
dria, Va vacuole. Scale bar = 10 µm in a; 1 µm in b–e; 2 µm in f 
agarwood formation, and the starch grains in the IP and XR
may provide a raw material foundation for these changes.
Conclusion
In this study, the structural characteristics of A. sinensis
wood and its relationship with agarwood formation were
studied. The results showed that the IP and XR intercon-
nected to form a living parenchyma cell network in A. sin-
ensis wood. After Agar-Wit treatment, the starch grains in
the parenchyma cells of IP and XR underwent a series of
changes and were eventually converted into agarwood resin;
the non-starch polysaccharides and phenols were some of
the intermediate products in the process of agarwood resin
formation. The resin initially forms and mainly accumulates
in the parenchyma cells of the IP and XR. Therefore, we
speculate that the IP and XR in A. sinensis wood provide
a structural and material foundation for the formation of
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agarwood resin. Further research is required to uncover the
regulatory mechanisms of the transformation of these sub-
stances during the formation of agarwood.
Author contribution statement  PL: participated in the
experiments, analyzed the data, and drafted the manuscript.
XZ and YY: performed the experiments. CS and YX: par-
ticipated in the design of the study and analysis of the data.
JW: participated in the design of the study and revised the
manuscript.
Acknowledgements  This work was supported by the funding from
the Hainan province Key Scientific and Technological Research and
Development Special Project (ZDKJ2016004), the National Natural
Science Foundation of China (81673549, 81703651), the Ten Thousand
Talent Program (99950534),the CAMS Innovation Fund for Medical
Sciences (2016-I2M-2-003,2016-I2M-1-012).
Compliance with ethical standards  a developmental oddity or an adaptive structure? PLoS One
Conflict of interest  The authors declare that they have no conflict of
interests.
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