Investigation of volatile oil glands of Satureja hortensis L. (summer savory) and phytochemical comparison of different varieties


Svoboda, K.P.; Greenaway, R.I.

International Journal of Aromatherapy 13(4): 196-202

2003


A study was conducted to determine which intrinsic factors (high fresh weight yield, high dry matter yield, large leaf area, or density, structure and size of oil glands on the plant surfaces) contribute to the high oil yielding performance of summer savory (S. hortensis) cultivars Aromata, Saturn and Compacta. There were no significant differences among the cultivars in terms of fresh weight and dry weight. Volatile oil yield showed clear differences between seed lots. It ranged from 0.9% for the classical type of savory to 3.5% for the 3 high-yielding cultivars. There were no statistical differences in oil composition between the cultivars. On all calyces of all cultivars, the glands were mainly on the surface, not in depressions. There were very few volatile oil glands on the petals. These were mainly on the surface, approximately 5-10 per corolla, evenly distributed and arranged in small groups. On the stems there were very few volatile oil glands, approximately 5 or 6 per cm length for all cultivars. The structure and size of all the volatile oil glands was similar on all surfaces and in all cultivars. The diameter of 20 glands was 123, 123, 124 and 130 micro m for Classic, Aromata, Saturn and Compacta, respectively.

K.P.
Svoboda
SAC
Ayr,
Department
of
Plant
Biology,
Ayr,
Scotland
KA6
511W,
UK;
E-mail:
INVESTIGATION
OF
VOLATILE
OIL
GLANDS
OF
SA'T'UREJA
HOR'T'ENSIS
L.
(SUMMER
SAVORY)
AND
PHYTOCHEMICAL
COMPARISON
OF
DIFFERENT
VARIETIES
K.P.
SVOBODA
AND
R.I. GREENAWAY
INTRODUCTION
p
lants
produce
primary
me-
tabolites
during
photosyn-
thesis
and
respiration.
These
are
two
fundamental
processes
of
life
and
are
termed
primary
metabolism;
the
organic
compounds
produced
during
the
processes
are
called
pri-
mary
metabolites
and
fall
into
four
distinctive
groups:
carbohydrates,
lipids,
proteins
and
nucleic
acids.
In
addition
to
these
processes,
plants
use
some
of
these
primary
metabolites
to
carry
out
additional
processes
which
we
call
secondary
metabolism.
Basically
they
are
called
`secondary'
because,
as
far
as
we
know,
they
are
not
essential
for
the
plant's
life.
They
encompass
a
wide
range
of
chemical
compounds,
such
as
volatile
oils,
tannins,
coumarins,
flavonoids,
saponins,
alkaloids,
steroids,
glycosides
and
polyamines.
Their
function
is
not
completely
un-
derstood;
it
is
presumed
that
they
act
to
attract
insects
and
animals
for
pollination,
to
deter
predators,
to
inhibit
growth
of
fungi
and
bacteria,
and
to
heal
plant
organ
wounds.
Essential
oils
are
known
to
be
produced
by
over
17,000
plant
spe-
cies,
distributed
all
over
the
world.
They
include
trees,
such
as
eucalyp-
tus,
and
humble
annual
plants,
such
as
water
mint.
Essential
oils
are
produced
and
stored
in
specialised
plant
secretory
tissues.
These
tissues
are
divided
into
two
main
types:
those
which
occur
on
the
plant
surfaces
and
usually
secrete
substances
directly
to
the
outside
of
the
plant
(exogenous
secretion)
and
those
which
occur
within
the
plant
body
and
secrete
substances
into
specialised
intercellular
spaces
(endogenous
secretion).
Secretion
is
a
common
feature
of
living
cells
and
the
secreted
material
may
contain
various
salts,
latex,
waxes,
fats,
flavo-
noids,
sugars,
gum,
mucilages,
as
well
as
essential
oils
and
resins.
An
example
of
an
inner
secre-
tory
resin
duct
is
in
the
leaf
of
Cedrus
brevifolia
(Fig.
1),
or
vittae
under
the
seed
coat
of
caraway
seed,
Carum
carpi
(Fig.
2).
This
picture
is
followed
by
a
close
up
of
the
vittae
in
the
same
species
(Fig.
3).
Another
example
of
an
inner
secretory
structure
is
a
se-
cretory
cavity
in
citrus
species
under
the
rind
(Fig.
4).
Glands
on
the
sur-
face
include
simple
sessile
glandular
trichomes
such
as
in
Oregano
species
(Fig.
5),
covered
by
a
comparatively
thick
cuticle.
These
glands
can
be
accompanied
by
the
dense
occur-
rence
of
trichomes,
such
as
in
Lavandula
angustifolia
(Fig.
6);
other
0962.4562/6
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Fig.
1
Cross
section
through
the
needle
of
Cedrus
brevifolia
with
detail
of
resin
duct
(rd).
Duct
is
lined
with
epithelium
of
secretory
cells
(sc)
[x3331.
Vittae
oils
in
which
one
main
component
makes
up
more
than
80%
of
the
whole
oil
(eucalyptol),
whereas
mar-
joram
produces
a
mixed
oil
in
which
no
single
oil
component
dominates.
Finally,
individual
plant
species
are
also
classified
into
chemotypes
or
chemical
races,
e.g.,
Eucalyptus
citrio-
dora
or
Nepeta
cataria
variety
citriodora.
There
is
no
definite
threshold
for
when
a
compound
can
be
classed
as
a
major
component
and
used
in
che-
motaxonomy
classification.
Many
people
have
differing
opinions
about
this.
For
example,
in
our
recent
study,
when
we
looked
at
the
collec-
tion
of
Ocimum
species
from
the
USA
botanical
garden
in
Ames,
Iowa,
we
classified
components
over
30%
as
the
major
ones
for
differentiation
between
the
individual
acquisitions
in
the
collection.
OBJECTIVES
OF
THE
INVESTIGATION
Fig.
2
Seed
pair
of
caraway
(Carum
carvi)
showing
secretory
structures
called
vittae
[x481.
'
1 /4 .
-1
/4 -
'
species
can
have
both
stalked
and
sessile
glandular
hairs,
as
can
be
seen
in
clary
sage,
Salvia
sclarea
(Fig.
7),
and
there
are
glands
with
a
multi-
cellular
head,
such
as
in
yarrow,
Achillea
millefolium
(Fig.
8).
The
essential
oil
yield
produced
by
these
structures
and
their
neigh-
bouring
cells
is
variable,
with
low
yielding
plants,
such
as
Melissa
offici-
-'.
'
nalis,
producing
on
average
0.05%
v/w
and
high
yielding
plants,
such
as
frankincense,
which
can
yield
up
to
18%
v/w
of
essential
oil
from
bark,
or
clove
flower
buds
producing
up
to
15%
v/w.
In
addition,
individual
species
produce
oil
profiles
which
differ
both
qualitatively
and
quantitatively.
Species
such
as
eucalyptus,
produce
Summer
savory,
Saturt
ja
hortensis,
is
an
annual
herb
of
the
family
Labia-
tae
and
is
widely
used
as
a
culinary
herb.
Commercially
the
fresh,
dried
and
ground
plant
material
is
mar-
keted
and
the
dried
material,
the
volatile
oil,
oleoresin,
tincture
and
extracts
are
used
as
seasoning
in
the
food
industry,
the
oils
and
extracts
in
the
perfume
and
toiletries,
and
in
recent
years,
in
aromatherapy.
Savory
belongs
to
the
low
yield-
ing
plant
species,
usually
producing
between
0.5
and
0.9%
v/w.
However,
a
search
for
higher
yielding
individ-
uals
within
the
population
resulted
in
the
selection
and
breeding
of
several
varieties
with
an
oil
yield
of
approximately
4%,
representing
a
considerable
economic
advantage
in
commercial
production.
The
aim
of
this
investigation
was
to
determine
which
intrinsic
factors
contributed
to
the
higher
perfor-
mance
of
these
varieties.
Is
it
higher
Fig.
3
Seed
of
caraway
showing
detail
of
primary
cotta
[x4061.
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Fig.
5
Sessile
secretory
gland
on
upper
leaf
surface
of
Greek
oregano
(Origanum
heracleoticum)
showing
oil-filled
subcuticular
space
resulting
in
fully-extended
cuticle
[x4201.
.4,
6
ra
,
t
1
k..4
6
.'?
s
i
d
it4Vf?^
v
irt
1
.
7
,6
"'FAL
o
w—
Fig.
6
Lower
leaf
surface
of
Lavandula
angustifolia
showing
sessile
secretory
gland
and
non-secretory
trichomes
[x
6381.
(a)
(b)
Oil
eon
ones
endocarp
memory
(pith)
ormaip
epidermis
Orange
Lemon
.re
f
'
Fig.
4
(a)
Transverse
section
through
orange
and
lemon,
showing
a
general
structure
of
citrus
fruit
with
oil
cavities
located
in
the
exocarp
[x61.
(b)
Detail
of
secretory
cavity
(transverse
section)
in
the
flavedo
of
orange
fruit
peel
[x961.
fresh
weight
yield,
higher
dry
matter
yield,
perhaps
larger
leaf
area,
or,
finally,
and
most
importantly,
the
density,
structure
and
size
of
oil
glands
on
the
plant
surfaces.
MATERIALS
AND
METHODS
Savory
has
been
bred
in
several
Eu-
ropean
countries
and
released
into
41k.
cultivation
as
distinct
varieties:
Aro-
mata
(Germany,
1986),
Saturn
(Poland,
1994)
and
Compacta
(Ger-
many,
1997).
We
obtained
seeds
of
these
va-
rieties,
together
with
the
'classical'
type
of
savory.
All
seed
lots
were
observed
for
germination,
vigour
and
growth.
Plants
were
measured
at
regular
intervals,
phenological
observations
and
detailed
observa-
tion
of
flowering
habit
and
general
growth
were
conducted.
All
plants
were
harvested
at
flowering
stage,
and
the
following
characteristics
were
measured:
fresh
weight,
dry
weight,
volatile
oil
yield
and
volatile
oil
composition.
Plant
material
was
distilled
using
a
British
Pharmacopoeia
Hydrodi-
stillation
Apparatus
(British
Stan-
dards,
1985).
GC
and
GCMS
were
carried
out
as
described
in
Galam-
bosi
et
al.
(1999).
Detailed
observations
of
volatile
oil
glands
included a
count
of
the
total
number
of
glands
on
five
ma-
ture
and
fresh
whole
leaves
from
several
plants
of
each
seed
lot
(an
average
of
50
leaves
per
seed
lot)
both
on
the
abaxial
and
adaxial
sur-
faces,
and
the
gland
density
was
as-
sessed.
The
glands
on
the
stems
and
flowers
were
also
investigated
and
described.
To
count
the
glands,
a
very
fine
needle
(microlepidoptera
mounting
pin)
mounted
onto
a
wooden
cock-
tail
stick,
was
used
as
a
microprobe,
each
gland
being
stabbed
and
burst
to
eliminate
double
counting.
The
leaf
area
was
calculated
by
photo-
graphing
the
leaf
with
a
millimetre
scale
and
by
transferring
the
image
to
the
image-editing
programme
on
the
computer.
A
1-mm
square
grid
98
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Fig.
7
Secretory
glands
on
the
surface
of
the
stem
of
a
single
floret
of
Salvia
officinalis
[x2501.
Fig.
8
Secretory
glands
on
the
ray
floret
of
Achillea
millefolium,
with
a
pollen
grain.
(Table
2).
The
highest
content
ap-
peared
to
be
during
the
period
of
full
flowering.
The
implication
of
these
find-
ings
is
very
important,
especially
for
the
large-scale
cultivation
and
pro-
duction
of
volatile
oil.
A
threefold
higher
volatile
oil
yield
represents
a
far
greater
financial
return
for
the
grower;
a
volatile
oil
yield
of
0.9%
at
1.2
t/ha
produces
11.2
litres
per
hectare,
but
3.2%
at
1.2
t/ha
pro-
duces
33.6
litres
per
hectare.
overlay
over
the
image
projected
the
leaf
area.
From
this
data
and
the
se-
cretory
structures
count,
the
gland
density
of
the
leaf
was
calculated.
RESULTS
Fresh
and
dry
weight
Detailed
results
for
fresh
weight
and
dry
weight
yield
were
recorded
for
each
variety.
There
were
no
signifi-
cant
differences
between
the
varieties
(Table
1).
Considering
that
dry
leaves
represent
the
final
commercial
product
and
taking
into
account
that
the
leaves
constitute
approximately
60%
of
the
whole
plant,
the
final
commercial
yield
for
all
individual
varieties
would
be
around
0.7
t/ha
of
dried
leaves.
Volatile
oil
yield
Investigation
of
volatile
oil
yield
showed
clear
differences
between
seed
lots.
It
ranged
from
0.9%
for
the
classical
type
of
savory,
to
3.5%
for
the
high
oil
yielding
varieties
(Aro-
mata,
Saturn
and
Compacta)
Volatile
oil
composition
A
detailed
analysis
of
the
volatile
oil
composition
of
individual
varieties
identified
the
major
components
as
carvacrol,
y-terpinene,
para-cymene;
and
the
minor
components
under
5%
as
oc-terpinene,
myrcene,
camph-
ene
and
oc-pinene
(Table
3).
There
were
no
statistical
differences
in
oil
composition
between
the
varieties.
The
full
flowering
period
represents
the
optimum
stage
for
harvesting,
particularly
with
regard
to
oil
com-
position,
where
a
high
carvacrol
content
is
required
by
the
food
and
cosmetic
industries.
Observation
of
secretory
glands
Volatile
oil
glands
of
the
sessile
pel-
tate
type
(Fig.
9)
were
distributed
on
the
leaf
surfaces,
sometimes
in
the
epidermal
depression
(Fig.
10).
The
glands
were
counted
on
each
variety,
on
five
young
leaves
(fully
expanded
and
mature
on
the
upper
part
of
the
plant)
and
on
five
older
leaves
Table
1
DRY
WEIGHT
(9)
OF
FOUR
DIFFERENT
VARIETIES
OF
SAVORY
(EARLY
FLOWERING
STAGE)
Stems
Leaves
%
Dry
wt./fresh
wt.
Classic
Aromata
Saturn
Compacta
3.7
3.9
2.4
3.4
4.9
5.6
4.2
5.1
21.9
22
17.3
18.9
Numbers
given
are
for
the
mean
figures
for
10
plants.
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2
VOLATILE
OIL
YIELD
(%
WW)
OF
FOUR
DIFFERENT
VARIETIES
OF
SAVORY
(FULL
FLOWERING
PERIOD)
Whole
plant
Flowers
Young
leaves
Older
leaves
Classic
0.9
Aromata
2.6
Saturn
3.1
Compacta
2.8
0.8
3.1
4.2
2.6
0.9
2.4
2.8
2.5
0.3
2
3.6
1.9
Numbers
given
are
the
mean
of
several
distillations.
Table
3
GC
ANALYSIS
OF
OIL
DISTILLED
FROM
PLANT
MATERIAL
Compound
Classic
Aromata
Saturn
Compacta
oc-Pinene
0.5
1
1.3
0.9
Camphene
0.8
1.3
2.1
1.6
Myrcene
1.6
2
2.9
2.3
oc-Terpinene
2.1
4.1
5.3
4.4
x-Terpinene
37.9
36.8
47.4
41.8
p-Cymene
6
1.6
2.1
1.9
Carvacrol
46.1
39.9
35.6
42.9
Total
%
95
86.7
96.7
95.8
Total
number
of
peaks
65
43
41
41
40um
›-
-
Fig.
9
Sessile
peltate
glands
on
savory
(Satureja
hortensis)
on
the
exterior
surface
of
corolla
[x5301.
Fig.
10
Sessile
recessed
gland
on
lower
swface
of
the
leaf
of
savory
(Satureja
hortensis)
[x6301.
(larger
and
from
the
lower
part
of
the
plant).
Total
leaf
area,
total
gland
count
on
the
lower
surface
and
the
number
of
glands
per
cm
2
were
calculated.
The
results
are
summarised
in
Table
4.
In
addition
five
leaves
from
each
plant
variety
were
taken
at
random.
The
glands
were
observed
and
screened
for
the
total
gland
count
on
both
surfaces
of
the
same
leaf
and
the
ratio
was
calculated
(Table
5).
On
the
lower
surface
the
glands
were
randomly
distributed
over
the
whole
leaf
including
the
main
vein.
On
the
upper
surface
the
glands
were
absent
from
the
middle
third
of
the
leaf
running
parallel
to
the
main
vein
and
had
a
preference
for
the
outer
edge
(see
Fig.
11).
In
addition,
the
glands
were
counted
on
the
calyces.
The
total
calyx
area
measured
was
approxi-
mately
0.8
cm
2
,
with
approximately
200
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Fig.
11
Sessile
secretory
cell
of
savory
(Satureja
hortensis)
on
the
exterior
surface
of
ca
ly
x,
showing
the
raptured
cuticle
and
twelve
secretory
cells
inside
[x640.1.
Table
4
TOTAL
VOLATILE
OIL
GLAND
COUNT
ON
THE
LOWER
SURFACES
OF
FOUR
VARIETIES
OF
S.
HORTENSIS
Variety
Leaf
age
Total
leaf
area
Total
gland
count
Glands
per
cm
2
Classic
Young
39.1
36.47
93
Old
71.1
5078
71
Aromata
Young
39.3
3699
94
Old
80.1
4808
60
Saturn
Young
43.7
4338
99
Old
75.4
5445
72
Compacta
Young
16.7
1647
99
Old
33.1
1965
59
Table
5
THE
RATIO
OF
GLAND
DENSITY
BETWEEN
ABAXIAL
AND
ADAXIAL
SURFACES
OF
LEAVES
OF
FOUR
VARIETIES
OF
S.
HORTENSIS
Variety
Total
glands
no.
on
upper
surface
Total
glands
no.
on
lower
surface
Ratio
upper/lower
Classic
Aromata
Saturn
Compacta
69
67
60
68
178
192
174
200
2.57
2.83
2.9
2.94
The
numbers
represent
the
total
number
of
glands
of
five
leaves.
(Mean
of
five
leaves.)
650
glands
per
cm
of
calyx.
Visually,
the
density
of
each
variety
was
very
similar.
On
all
calyces
of
all
varieties,
the
glands
were
mainly
on
the
sur-
face,
not
in
depressions.
There
were
very
few
volatile
oil
glands
on
the
petals.
These
were
mainly
on
the
surface,
approximately
5-10
per
corolla,
evenly
distributed
and
arranged
in
small
groups.
On
the
stems
there
were
very
few
volatile
oil
glands,
approximately
5
or
6
per
cm
length
for
all
varieties.
The
structure
and
size
of
all
the
volatile
oil
glands
was
similar
on
all
surfaces
and
in
all
varieties.
Each
volatile
oil
gland
contained
12
se-
cretory
cells
at
the
base
(Fig.
10)
with
four
cells
approximately
centrally
positioned
forming
a
rough
square,
around
which
there
was
a
circle
of
eight
cells.
The
diameter
of
20
glands
from
each
plant
variety
was
measured
(mean
mesure):
Classic
123
Aromata
123
Saturn
124
µm
and
Compacta
130
CONCLUSIONS
From
the
results
it
is
obvious
that
fresh
and
dry
matter
yield
are
not
factors
responsible
for
the
differ-
ences
between
low
and
high
oil
yielding
varieties.
Having
eliminated
this
factor,
the
further
hypothesis
was
that
the
low
oil
yielding
varieties
would
have
a
lower
amount
of
glands
per
cm
2
,
would
be
of
smaller
size,
or
different
structure
which
would
re-
flect
the
low
oil
yield
production.
However,
this
was
not
the
case
and
it
is
necessary
to
look
for
another
explanation.
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The
explanation
could
lie
in
in-
creased
synthesis
of
oil
from
the
biosynthetic
pathways
which
lead
to
monoterpene
production.
These
pathways
are
facilitated
by
a
number
of
enzymes,
which
are
genetically
programmed.
Their
function
may
be
enhanced
in
high
yielding
varieties.
To
elucidate
this
hypothesis
it
would
be
necessary
to
conduct
a
DNA
analysis
of
the
given
type
of
savory,
map
the
genes
and
study
the
ex-
pression
of
individual
enzymes.
Until
recently,
terpenoids
(the
main
components
of
volatile
oils)
were
assumed
to
be
formed
exclu-
sively
by
the
mevalonate
pathway.
It
has
been
now
proved
that
there
is
an
alternate
metabolic
route
leading
to
their
production,
both
in
plants
and
in
the
majority
of
bacteria.
This
path-
way
is
called
the
1-deoxy-D-xylulose-5-
phosphate
pathway,
and
only
during
the
last
year
were
many
missing
links
and
enzymes
identified,
with
the
use
of
isotopic
labelling
and
genetic
in-
vestigation
using
microbial
mutants
of
various
gene
functions.
The
regula-
tion
of
plant
metabolism,
based
on
genetic
and
biochemical
knowl-
edge,
provides
considerable
insight
into
the
biosynthesis
of
secondary
met-abolites,
including
volatile
oils.
Characterisation
of
enzymes
and
cor-
responding
genes
will,
in
the
near
fu-
ture,
answer
the
questions
posed
in
this
study
and
with
high
probability
will
also
influence
the
quality
and
quantity
of
oil
obtained
from
plants
through
the
use
of
biotechnology.
ACKNOWLEDGEMENTS
SAC
receives
financial
support
from
the
SEERAD
and
this
investigation
was
a
part
of
a
core
project
registered
as
Developing
Bioactive
Crop
Com-
ponents
with
Antioxidant
and
Anti-
microbial
Properties.
Andrew
Syred
carried
out
the
SEM
photographs
(Microscopix
Photolibrary,
Middle
Travelly,
Beguildy
near
Knighton,
Powys,
Wales
LD7
1UW,
UK)
and
Professor
Y.
Asakawa
verified
the
GC
analysis
of
volatile
oils
by
GCMS
(Faculty
of
Pharmaceutical
Sciences,
Tokushima
Bunri
University,
Yamas-
hiro-cho,
Tokushima
770,
Japan).
We
would
like
to
acknowledge
technical
assistance
of
V.
McFarlane,
J.
Morton
and
T.
Warrie,
SAC.
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