Flora (2000) 195, 83-94

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Distribution, development and structure of resin ducts

in Pistacia lentiscus var. chia DuHAMEL
THOMAS SAWIDIS 1, STEFANOS DAFNIS 1 and ELZBIETA WERYZKO-CHMIELEWSKA 2

1 Institute of Botany, University of Thessaloniki, GR-54006 Thessaloniki, Macedonia, Greece

2 Katedra Botaniki, Akademia Rolnicza, PL-20950 Lublin, Poland

Accepted: December 21, 1998

Summary
The distribution, anatomy and development of the resin duct system responsible for the resin exudation in Pistacia lentiscus
var. chia DuHAMEL were investigated. Resin ducts are present in the stem, leaf and root located only in the phloem of the vas-
cular bundles. They are tubular structures lined by a few layers of epithelial cells. Outwards the resin duct is well supported by
arcs of sclerenchyma fibres. In the petiole and petiolule 5 resin ducts were found whereas in the midvein of the leaflet 3 resin
ducts are present adjacent to vascular bundles. In the leaf primordium they are located in the mesophyll below the dense con-
ductive palisade parenchyma. Furthermore resin ducts were found in the leaf-scales protecting the dormant lateral buds. Ducts
are surrounded by numerous crystal idioblasts. All resin duct systems of the root and stem, though separated from one another,
form a continuous system with the corresponding ones in the leaf. The inner layers of epithelial cells seem to decompose releas-
ing their content into the duct lumen. In this space large masses of disorganized cytoplasm with many vesicles and organelles
and wall remnants of the disintegrated cells could be found.

Key words: Resin ducts, gum, mastic, Pistacia lentiscus var. chia

1. Introduction variety P lentiscus var. chia is a tree-like bush, known

The genus Pistacia of Anacardiaceae family comprises
some eleven species of trees in the warm temperate re-
gions of Europe, Asia, Southern and Central America.
According to ZoHARY (1972) the centre of the origin of
the genus is Central Asia, though the genus has a dis-
continous pattern of distribution. The various species of
Pistacia are spread mainly around the Mediterranean
basin and comprise important constituents of the natural
vegetation in this area. Many Pistacia species yield
resin in some degree. However, only two of them, name-
ly P atlantica and P lentiscus var. chia, are the major
sources of resin that can be collected, traded and used
and which has been an important article of commerce
for centuries. Both species are cultivated on the Greek
island of Chios and yield the resins known as Chian tur-
pentine and mastic gum respectively.
P lentiscus L. is a small dioecus woody evergreen
shrub 1-8m, with resinous bark, like all other species
of the genus Pistacia extending in the Mediterranean
region to Portugal (TunN et al. 1968). In contrast its
0367-2530/00/195/01-083 $ 12.00/0
since the ancient times. It is interesting that plantations
of mastic trees occur only on a single island, Chios.
Attempts to expand the cultivation to other neighbour-
ing islands or in the adjacent western Asia Minor, have
not been successful (HELDREICH 1862, PERNOT 1981).
Moreover, on Chios island itself the plantation is re-
stricted to its southern part (Fig. 1), which is distinctly
dry and hot.
Mastic is extracted after shallow incisions made on
the trunk and the thicker branches of the plant with spe-
cial tools (Fig. 2). This incision is repeated three times a
year, from July to September. The resinous exudate
flows out of the tree trunk and branches in the form of
tears (Fig. 3). The resin coagulation lasts about 15 days
and is collected by hand (PERIKOS 1993). The product
in the form of crystal-clear drops (Fig. 4) is a natural
resin of extreme adhesiveness. The information on
Chios mastic dates back to Herodotus, Hippocrates,
Thophrastus, Dioscurides and Galenius (BECKMANN
1793, HELDREICH 1862, HEHN 1894, MANUSAKIS 1984,
BROWICZ 1987).
FLORA (2000) 195 83

N leaf segments from the same tree were also prepared. Semi-
A thin sections of 0.5-1.0 llm thickness from resin embedded
GREECE
30km
Fig. 1. Map of the island ofChios (N. Aegean sea, Greece). In
the southern part of island occur 21 villages where mastic is
exclusively grown to produce gum mastic.
It has long been assumed that synthesis of resins takes
place in specialized anatomical structures, the resin
ducts. The presence of resin ducts, which are associated
in the stem with the primary and secondary phloem is
common in Anacardiaceae (JOEL & FAHN 1980).
Phloem serves principally as a conducting system for
transporting photosynthetic products throughout the
plant. The sap is maintained under pressure within the
sieve elements and damage elicits an abrupt release of
sap from severed veins. In many species resin ducts are
also found in leaf, root, flower or fruit.
Although the Chios mastic, and its by-products, have
been well known from ancient time, little information is
available on the morphology of various plant organs as
well as the distribution, anatomy and development of
the resin duct system responsible for the resin exuda-
tion. The present paper deals with the above subject, as
a prelude to forthcoming research work.
84 FLORA (2000) 195
2. Materials and methods
Stem portions, from apical (1 year old) and mature (2-3 years
old) regions containing bark attached to wood were taken from
a mature living tree. Stem segments were fixed in 2.5% glutar-
aldehyde and 2% paraformaldehyde in 0.05 M cacodylate buf-
fer for 2h. After dehydration in an ethanol series and post-
fixation in 1% osmium tetroxide, the tissue was embedded in
Spurr's epoxy resin (SPURR 1969). In the same way root and
tissue were heat-fixed to glass slides and stained with 0.5%
toluidine blue in 5% borax or 0.5% safranin in 50% ethanol or
1% pianese mixture in 30% ehtanol for preliminary light
microscope (LM) observation. In hand sections of fresh mate-
rial secondary xylem was stained with periodic acid-Schiff's
(PAS) reagent. Semithin sections were examined under a Zeiss
axioplan light microscope. Ultrathin sections were examined
using a Zeiss 9 S-2 electron microscope (TEM).
For scanning electron microscopy (SEM), specimens were
fixed in cacodylate-buffered glutaraldehyde without a post-
osmium fixation. After dehydration in ethanol series
(10-99.8% ), specimens were critical-point dried with liquid
C0 2 as an intermediate and coated with gold in CS 100 Sput-
ter Coater. Observations were made using a BS-340 Tesla
scanning electron microscope at various accelerating poten-
tials.
3. Results
3 .1. Distribution of the resin ducts
Resin ducts were present in the stem (Fig. 5, 6), leaf
(Fig. 7) and root (Fig. 8, 9). Fruits and flowers were not
investigated. In the stem (both primary and secondary
Fig. 2. Transverse cutting on the trunk of the tree caused by
special tool.
Fig. 3. Coagulated large mastic tears hanging from the tree
trunk.
Fig. 4. Mastic tears after collection and primary separation
from the leaves and soil.
Fig. 5. Hand cross-section of a young stem at the beginning of
secondary growth. Resin ducts (rd) in the metaphloem (ph) of
the vascular bundles. Xylem (xy) stained with periodic acid-
Scruff's reagent. ca = cambium, co = cortex, sc = scleren-
chyma. 100 f.!m.
Fig. 6. Resin duct in the primary phloem of a young stem.
Staining safranin. 50 f.lm.
Fig. 7. Hand transverse-section of a leaf resin duct. 100 f.lm.
Fig. 8. Longitudinal section of a root resin duct. Abundant
oxalate crystals mainly on the lower (inner) side of the duct.
Staining safranin, polarization optics. 20 f.lm.
Fig. 9. Idioblast containing oxalate crystals in the vicinity of
a root resin duct. Staining safranin, polarization optics. 5 f.lm.
FLORA (2000) 195 85
86 FLORA (2000) 195
longitudinal resin ducts are located only in the phloem
of the vascular bundles. They are unbranched and in
contact with each other only in the nodes. The resin duct
consists of a few layers of epithelial cells. After 10-15
cell layers of phloem tissue outwards, the resin duct is
well supported by arcs of sclerenchyma fibres (Fig. 10).
The sclerenchyma tissue consists of 3-4 cell layers and
surrounds the duct only from the outer side. Idioblasts
containing oxalate crystals are also present in the
surrounding area. The epidermis of the young stem
(1 year old) is covered by solid, non-glandular hairs
(Fig. 11). Glandular hairs were not observed. In the
secondary xylem vertical ducts are absent.
The inner layers of epithelial cells seem to decom-
pose releasing their content into the resin duct (Fig. 12).
The epithelial cells surrounding the resin duct are elon-
gated in the direction of the long axis of the duct
(Fig. 13). The other cells outwards are cortical paren-
chyma cells, many of which contain large vacuoles with
electron dense material. The latter cells are irregularly
distributed around the duct.
The secondary stem is surrounded by a multilayer
cork (phellem) for mechanical protection. The cork of
the short trunk or the thicker branches are frequently
covered by the thalli of lichen Xanthoria parietina
which is specifically associated with the mastic trees
and not on other tree species growing in the vicinity.
The leaves of Pistacia are compound, imparipinnate,
alternate and estipulate. The leaflets are lanceolate and
sclerophyllous. In leaves only longitudinal ducts were
found joining the primary (meta-) and secondary
phloem (when produced). In the petiole and petiolule 5
resin ducts were found joining the individual vascular
bundles. The epithelial cells have thick cell walls with-
out any intercellular space (Fig. 14).
In the mid vein of the leaflet 3 resin ducts were found
Fig. 10. Scanning electron micrograph of a stem resin duct.
The sclerenchyma fibres (sc) protect the duct from the outer
side. 30 11m.
Fig. 11. Scanning electron micrograph of numerous non-glan-
dular solid hairs on the epidermis of a young stem. 30 11m.
Fig. 12. Electron micrograph of a stem resin duct showing
disintegration of epithelial cells. In the duct lumen remnants of
a disintegrated cell (large osmiophilic droplets) are seen. 3 11m.
Fig. 13. Longitutinal section of a stem resin duct. The epithel-
ial cells are elongated in the direction of the long axis of the
duct. 50 11m.
Fig. 14. Electron micrograph of an epithelial (swollen) and
neighbouring (flattened) cells of a petiole resin duct. Thick cell
walls lacking any intercellular space. 2 11m.
Fig. 15. Light micrograph of a mature leaf cross-section.
Resin duct is located below the dense palisade parenchyma.
100 !lm.
adjacent to vascular bundles. In the leaf primordium
they were located in the mesophyll below the dense con-
ductive palisade parenchyma (Fig. 15). Idioblasts were
frequent in the loose spongy parenchyma.
Both upper and lower epidermis of the leaf are
relatively thick consisting of small epidermal cells.
Numerous, relatively large, stomata are uniformly
arranged in the lower epidermis. A thick waxy cuticle
covers both lower and upper epidermis which addition-
ally presented a coarsely striated ornamentation
(Fig. 16). The cuticle on the upper side of the observed
leaves is rather smooth.
Resin ducts were also present in the young leaves.
Furthermore, relatively well-developed resin ducts were
found in the leaf-scales supporting the dormant lateral
buds (Fig. 17). The protective solid stem trichomes were
already formed in the lateral bud.
In young roots (primary construction) resin ducts
were observed neither in the central cylinder (Fig. 18)
nor in the cortex (Fig. 19). The cortical parenchyma
showed large intracellular spaces. A number of cells
with visible dark content were observed. In longitudinal
and cross sections these cells were elongated containing
raphide crystals (Fig. 20, 21).
In the secondary construction of the root a number of
longitudinal resin ducts of different size were found in
the cross section of the root. Ducts occurred in second-
ary phloem and in adjacent parenchyma, but not in wood
rays. In some of these ducts secretion was also observed
around the epithelial cells (Fig. 22). Ducts were sur-
rounded by numerous crystal idioblasts (Fig. 8, 9).
3.2. Development of resin ducts
Ducts initiate and develop schizogenously in the prim-
ary phloem. The formation of an interceiJular space
among a group of densely stained cells with conspi-
Fig. 16. Scanning electron micrograph of large stomata in the
abaxial (lower) leaf surface. Striate ornamentation of cuticula.
10 !lm.
Fig.17. Lateral bud. Scales bear resin ducts. Non-glandular
(solid) trichomes are already formed (arrow heads). The inset
shows details of the scale resin duct and trichomes. I 00 11m.
Fig. 18., 19. Light micrographs of a primary root cross section
of the central cylinder and cortex respectively. Resin ducts are
not yet developed. Cortex parenchyma cells with many intra-
cellular spaces, some of them (arrows) with a visible dark con-
tent. 200 11m.
Fig. 20., 21. Longitudinal- and cross-section of raphide idio-
blasts. 50 11m.
Fig. 22. Secondary root cross section. Resin ducts in the
secondary phloem and the adjacent parenchyma. 50 11m.
FLORA (2000) 195 87
88 FLORA (2000) 195
Fig. 23. Resin duct in the metaphloem at stage of differentiation. The intercellular space among the epithelial cells is already
formed. 5 IJm.
Fig. 24. Scanning electron micrograph of a stem resin duct lined by epithelial cells. 20 IJm.
Fig. 25. (Detail from Fig. 12). Swollen epithelial cell of a stem resin duct showing large nucleus and osmiophilic droplets (aste-
risk) in the cytoplasm. Similar osmiophilic droplets are also shown in the neighbouring cells and into the duct lumen. Vesicular
structures (arrows) occurred between plasmalemma invaginations and cell wall. lj.im.
Fig. 26. A vesicle between plasmalemma and cell wall. 0,2 11m.

FLORA (2000) 195 89

90 FLORA (2000) 195
cuous nuclei and nucleoli in the phloic procambium and
ground tissue signals the initiation of the duct. The inter-
cellular space enlarges into a distinct lumen by separa-
tion of walls of the lining cells and their subsequent re-
arrangement forming the lumen. They constitute the
epithelial sheath cells which divide anticlinally and
show tangential flattening in the resin duct (Fig. 23).
These epithelial cells are compactly arranged in radial
rows, thick walled and devoid of intercellular spaces
(Fig. 14). The resin duct develops through either the
separation or breakdown of epithelial sheat cells. The
elongate spaces connect or ramify to form a complex
system throughout the whole plant.
3.3. Structure of resin ducts
Tangential resin ducts are tubular structures lined by
epithelial cells (Fig. 24). A young duct shows the sepa-
ration of cells along their radial walls leading to the
enlargement of the intercellular space and participation
of neighbouring cells to form an epithelium. Later
epithelial cells show disintegration of inner tangential
walls and the subsequent release of the cell contents into
the duct lumen. The neighbouring cells become the new
epithelial cells.
The epithelial cells, which are about to disintegrate,
usually contain a large nucleus and the cytoplasm be-
comes gradually osmiophilic. Large lipid droplets are
present in the cytoplasm similar to those released al-
ready into the duct lumen (Fig. 25). At this stage nume-
rous vesicles occur around the walls of these cells for-
med by invagination of the plasmalemma (Fig. 25, 26).
Further widening of the duct occurs by disintegration
and/or separation of epithelial cells. However this
widening of the duct is not a continuous process and the
cells lining the duct lumen are not specialized as epi-
thelial cells. The mode of development of a duct varies
Fig. 27. A regularly developed midrib resin duct. Staining
safranin. 3 !Jill.
Fig. 28. A leaf midrib resin duct irregularly developed. Abun-
dant phenolics containing cells in the duct epithelium. Staining
safranin followed by pianese mixture. 3 !Jill.
Fig. 29. A mass of disorganized cytoplasm in a petiolule resin
duct lumen. Free cytoplasm and oil droplets into the duct ori-
ginated from the former epithelial cells. Staining pianese mix-
ture. 5 !Jill.
Fig. 30. Whole cell walls within a midrib resin duct. Staining
safranin. 3 !Jill.
Figs. 31, 32. Parts of longitudinal section of a root resin duct.
Disintegrated cells are almost removed from the duct epitheli-
um. Their walls show abundant vesicles (arrows). Staining
safranin. Fluorescence optics. 3 !Jill.
among different resin ducts of the same plant organ.
When the duct development is continuous, the shape is
more or less round (Fig. 27). However in developing sta-
ges the epithelial cells are ruptured and so the sheath is
not well defined. In some cases the abrupt growth of
neighbouring cells, which form the epithelium, causes a
partial filling of the duct lumen. Such cells show intense
vacuolation or accumulation of phenolics and change
the duct shape into a branched irregular slit (Fig. 28).
In the duct lumen itself masses of disorganized cyto-
plasm and organelles of disintegrated cells were fre-
quently found (Fig. 29). Cell wall remnants or whole
cells, after releasing their content were also present
(Fig. 30). In some cases the disintegrated cells remote
early from the epithelial tissue (Fig. 31, 32).
4. Discussion
There is much confusion in the literature concerning the
mode of origin of the resin ducts. Different opinions
were often expressed, concerning ducts of the same
plant species. This may depend on the species and more
specifically on the considered tissue (VENKAIAH &
SHAH 1984). According to ENGLER (1896) within the
Anacardiaceae all secretory ducts are developed schizo-
genously. Later several authors described secretory
ducts of schizogenous, lysigenous or schizo-lysigenous
origin depending on the plant organ and considered
species (JOEL & FAHN 1980, DEN OUTER & VAN
VEENENDAAL 1986).
In Pistacia lentiscus var. chia it is obvious that the
ducts originate schizogenously and later become
lysigenous because the epithelial cells lining the duct
swell, disintegrate and are fed from their neighbouring
cells as is likely in some other genera of Anacacardia-
ceae (JOEL & FAHN 1980, VENKAIAH & SHAH 1984,
DEN OUTER & VAN VEENENDAAL 1986). In Rhus glabra
the ducts were found to develop schizogenously (FAHN
& EvERT 1974). In Trichoscypha in some tissues secre-
tory ducts originate schizogenously and later become
lysigenous but in some other tissues they develop and
expand lysigenously (DEN OUTER & VAN VEENENDAAL
1986). The same secretory duct development also
occurs in Mangifera indica (JOEL & FAHN 1980). In
investigated species of the Cactaceae family it is
suggested that schizolysigenous ducts are ancestral to
strictly lysigenous ones (WITTLER & MAUSETH 1984).
In Anacardiaceae, secretory ducts, when present, are
always found in the stem. The studied Pistacia lentiscus
var. chia shares the general anatomical feature of the
axis of all Anacardiaceae, namely the oc~urrence of gum
resin ducts (ENGLER 1896), or resin canals, laticiferous
canals or secretory ducts in the phloem (METCALFE &
CHALK 1983, FAHN & EVERT 1984). Resin ducts are
FLORA (2000) 195 91
generally present in the primary phloem immediately on
the inside of the arcs of sclerenchyma fibres and in the
secondary phloem. In some other genera resin ducts
occurring in the rays were also reported but much less
frequent in the cortex.
Pistacia leaves belong to the nanophyll type accord-
ing to 0RSHAN (1986). Small leaves are considered an
adaptation to mediterranean conditions. The structural
diversity of the plant organs may enable this plant to
withstand the drought and high summer temperatures as
well as the frost of not-so-mild winters. The duct is
located rather in the lower part of the mesophyll (spongy
parenchyma), for better protection from the light and
overheating. For the same reason the spongy paren-
chyma is very loose for better aeration.
The large, uniformly arranged stomata on the lower
surface (Fig. 16) may complement the above adaptation.
Such abnormally large stomata with relatively well-
developed outer rims (ledges) were quite common in sev-
eral other genera of Anacardiaceae (WILKINSON 1979).
SITHOLEY & PANDEY (1971) proposed the term "giant"
stomata to distinguish them from true water-stomata on
the one hand and normal stomata on the other. The pre-
sence of water-stomata, secreting drops of water, in Pi-
stacia lentiscus var. chia which is spread in the dry and
full sunlight Mediterranean region is out of the question.
The striate ornamentation of cuticles (Fig. 16) is a
common characteristic of Anacardiaceae and other
xerophytic families (WILKINSON 1979). Such striated or
wrinkled cuticles frequently occur in shade leaves or the
lower epidermis of the sunny ones. The morphological
and anatomical leaf features enable Pistacia lentiscus to
be one of the most tolerant shrubs to drought and high
temperature among many sclerophyllous plants of
Mediterranean climate (GRATANI 1995).
Cells containing phenolic coumpounds are more fre-
quent in the leaf ducts. The role of these parenchy-
matous cells around the duct is the chemical defense
against pathogens and folivorous insects. Thus when
these cells are ruptured the content deposited in their
specialized vacuoles flows to the side of damage. A her-
bivore biting into a resin duct soon encounters a copious
outflow of unpleasant and often toxic secretion. Pheno-
lic compounds provide effective coverage throughout
the resin duct. They have the advantage the defense can-
not be readily dismantled. The prize for this defense is
susceptibility to herbivores with appropriate behaviour-
al counteradaptations. Thus the chemicals may also be
less effective since herbivores encounter them in dilute
form. Another role of the above cells could be the
adsorption of solar radiation, which may cause an
overheating and polymerization of the resin. This prob-
ably affects the viscosity and the flow of the secreted
product within the resin ducts.
In young stems (primary construction) where the
92 FLORA (2000) 195
periderm is not yet developed the sclerenchyma arcs
around the duct as well as the numerous hard solid hairs
which are present on the epidermis at this stage obvious-
ly support the defense against insect attack. This hypo-
thesis unfortunately is lacking enough observational or
experimental support.
The extensive root system of Pistacia explores shal-
low soils horizons, as an adaptive strategy in the arid
mediterranean region, in order to use up short seasonal
rainfalls. Therefore the protection of the root and espe-
cially of the duct content from worms or other animals
seems to be vital (KUMMEROW 1973). This task is under-
taken by numerous idioblasts containing crystals in the
tissue which surrounds the duct (Fig. 8, 9). The presence
of raphides in young roots (Fig. 20, 21) is a surprising
observation since the Anacardiaceae are not listed as
having them in METCALFE & CHALK (1983). On the
other hand cells containing condensed phenolics found
in all tissues, and in the root as well, may also support
this adaptation (WALLACE & MANSELL 1975, SAWIDIS
1998).
The formation of resin involves two stages. Initially
secretion of the resin substance occurs from the epithe-
lial cells and then in the next stage, disintegration of
epithelial or neighbouring cells follows a process called
retinosis. The walls of these cells dissolve gradually,
beginning with the layers adjacent to the cytoplasm. It is
suggested that the numerous vesicles (Fig. 25, 26) which
seemed to load their content into the apoplast are carry-
ing the lytic enzymes for the expansion and finally dis-
integration of the cell wall. The so-called retinosis
employs in the elimination of resin components from the
cytoplasm of epithelial cells into the duct lumen.
The various cell contents from epithelial cells, as well
as the cell wall remnants, membranes, etc. form there an
amorphous component which is the prodromous resin
(Fig. 29). In the past such cell remains in the duct lumen
were interpreted as artifacts (Me NAIR 1918). JoEL &
F AHN ( 1980) reported that both cytoplasm and cell walls
disintegrated and that the only wall remains were those
attached to living epithelial cells. In our observations
not only cell walls remnants of disintegrating cells were
observed but also the whole cells almost free in the duct
lumen (Fig. 30, 31, 32).
Resins are generally a mixture of volatile compounds
and non-volatile ones, mostly diterpenses in the form of
resin acids. According to FAHN (1979) the secreted
material into the secretory ducts of Anacardiaceae, con-
tains terpenes and polysaccharides. In ducts of Rhus
glabra (FAHN & EvERT 1974) as well as in those of
Trichoscypha oba (DEN OUTER & VAN VEENENDAAL
1986) small lipophilic droplets were present. MARNER
et al. (1991) identified four novel triterpenoids in the
neutral faction of gum mastic, the resin of P. lentiscus.
According to recent analyses of acidic fraction of the
resin of P. lentiscus var. chia (PAPAGEORGIOU et al.
1997) ten triterpenoid acids were tentatively identified
as their methyl esters.
The presence of resin ducts as well as the general
structural characters of various plant organs is a part of
the strategy that Pistacia lentiscus var. chia has develop-
ed to deal with the stressful mediterranean climatic con-
ditions. All duct systems of the root and stem form a
continuous system with the corresponding ones in the
leaf. The presence of resin ducts in the form of a three-
dimensional network in leaves, stem and roots increases
the water-holding capacity of tissues, helping the plant
to prevent desiccation (KRAMER & KOZLOWSKI 1979,
VENKAIAH & SHAH 1984).
Fluid within resin ducts is typically stored under pres-
sure (BUTTERY & BOATMAN 1979). Thus when an insect
severs veins, it effectively ruptures the resin ducts, and
thereby diminishes the outflow of secretion beyond the
cuts. Additionally the adhesiveness of resins poses a
potentially lethal hazard to small insects, which may
become entrapped in exudate (DussouRD & DENNO
1991). On the other hand a mechanical injury to the bark
will immediately exude resin with the sap in order to
coat the cut surface.
The presence of resin at the wound area may serve the
plant to seal the wounds and thus the stem is rendered
less liable to infection (HABERLANDT 1914, BANKO &
HELTON 1974). On this feature of Pistacia lentiscus var.
chia is based the extraction of resin, used as mastic, on
Chios island. The number of cuttings for resin yielding
is proportional to the size and age of the tree and reaches
sometimes to 100 over a whole period.
5. Acknowledgements
The authors are thankful to the General Secretary of Research
and Technology of the Ministry of Research and Technology
for supporting this work by a grant (Code Nr. 1757). We are
also thankful to Mr. COSTAS STAMOULIS for providing us the
pictures nr. 2, 3 and 4.
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