Proc. of Second World
Avocado Congress 1992 pp. 191-198
Properties
of Persea indica, an Ornamental for Southern California
Ursula
K. Schuch, Rainer W. Scora, and Eugene A. Nothnagel
Department
of Botany and Plant Sciences, University of California, Riverside, CA 92521,
USA
Steven
D. Campbell
Department
of Plant Pathology, University of California, Riverside, CA 92521, USA
Abstract. Persea
indica L. is the only member
of the genus Persea surviving the Laurasian Mediterranean flora in
Europe and Africa. Its evergreen foliage and small fruit make P. indica an
attractive ornamental and a useful shade tree that improves the microclimate
throughout the year. The concentrations of macro- and micronutrients in leaf
tissue are comparable to the recommended levels for mature avocado trees.
Analysis of essential oils shows high amounts of sesquiterpenes, of which
caryophyllene and delta-cadinene are the major constituents. Chemical
components of the leaf oils of the commercial Hass and Fuerte cultivars show many
quantitative and qualitative differences compared to P. indica. Immature
fruit of P. indica can be used in a bioassay to detect Phytophthora
cinnamomi, P. citricola, and P. parasitica in soil.
Persea indica (L.) Spreng is
a living fossil, surviving the rich flora of the middle and late Pliocene
vegetation (Axelrod, 1978). As the only member of the genus Persea in
the once widespread Laurasian Mediterranean flora, P. indica currently
survives in the maritime climate of the Canary Islands, Madeira, and the
Azores. P. indica evolved in the African Gondwanaland flora and
dispersed to Europe where fossils were found in the Miocene flora of Southern
France and Spain (Axelrod, 1978). According to fossils identified in California
and dated back to the Miocene, this plant must have migrated west at least 50
million years ago.
Plants
of the Lauraceae are characterized by prominent oil cells in the leaves, wood
and fruit. These oils are mostly aromatic and find wide use as medicine
(sassafras tea, camphor), spices (cinnamon), and flavors (bay leaf) (Schroeder,
1976). P. indica is classified in the genus Persea, and the
subgenus Eriodaphne. Although P. indica is a relative of the
commercial avocado, P. americana, these two species are not graft
compatible (Schroeder and Frolich, 1955).
Ornamental properties. Plants
considered for landscape use enhance the environment in a variety of ways:
physical, aesthetic, economic, and psychological (Harris, 1983). Landscape
trees in populated areas are valued mainly for their improvement of the
microclimate. Shade from trees reduces solar radiation and reflection from
soil. Trees also modify wind patterns and reduce wind velocity. To a certain
extent, plants can remove pollutants from the air, and can absorb noise. Visual
benefits play an increasing role in densely populated areas, and in this regard
trees fulfill many different functions, such as providing color, form, texture,
and patterns in the landscape. Economic benefits of trees in the landscape
range from monetary value for the wood to an increase in real estate value. In
addition, trees can provide a potential wildlife habitat in urban settings.
Because
Persea indica exhibits many of these desired features as an ornamental
tree, it is frequently planted in Florida and Southern California. The smooth
and silver-colored bark provides a striking contrast to the shiny dark green
foliage. The tree grows with a straight trunk and has been used as a source of
timber in its native habitat. The leaves are oblong, glabrous, and 6 to 13 cm
long. Fruit are scarcely fleshy, grow to a length of under 2 cm and are borne
in clusters (Fig. 1). Immature fruit are green and turn black when ripening.
The evergreen foliage and the small fruit make P. indica an attractive
ornamental and a useful tree that ameliorates the microclimate throughout the
year.
Figure 1. Leaves and immature fruit of Persea indica.
Leaf nutrient analysis. The
nutritional status of mature P. indica leaves was determined. Samples
were collected in February from the distal end of branches, oven dried at 70C,
and analyzed for macro- and micronutrient concentrations.
Essential oils and alkanes in leaves. For essential oil analysis, 500 g of healthy leaves were randomly
harvested from a 32-year-old P. indica seedling tree at the University
of California Research and Extension Center, Irvine, California. The leaves
were macerated and steam distilled in a Clevenger apparatus for 2 h. The
resulting oil was kept under nitrogen until injected into a Shimadzu Mini-3 gas
chromatograph with a flame ionization detector and 50:1 split ratio. A J&W
DB-5 fused silica capillary column was used with 0.25 mm i.d., 60 m length and
0.25 micrometer coating thickness. The injector temperature was 200C. The oven
temperature was programmed to 65C for 30 min and then increased at a rate of
1C/min. to 200C, where it remained for 25 min. N-octane and n-eicosane were
added to some samples for standardization of retention times.
For
mass spectra identification a Hewlett-Packard 5890II gas chromatograph, a 5971A
mass selective detector and G1030A MS computer station were used. A capillary
direct interphase heated at 280C delivered column output into the quadruple
mass spectrometer with electrons of 70 eV energy for electron impact
ionization. The appropriately diluted oil in hexane (1 //L) was injected in
splitless mode. Injector temperature was 270C and helium flow was 22 cm/sec.
All other parameters were the same as for the Shimadzu gas chromatograph. Some
mass spectra were identified by the NIST/EPA/MSDC data base (49,000 spectra)
stored in the computer.
Indicator for Phvtophthora. Phytophthora root rot is a
serious disease causing substantial losses in avocado production (Zentmyer,
1980). In the seedling stage P. indica is very susceptible to Phytophthora
cinnamomi, the major pathogen of commercial avocado, and has been used to
test for the presence of the fungus (Zentmyer and Ohr, 1978). For this test,
young seedlings are placed in soil suspension, and within 3 to 5 days black
stripe cankers develop on the stem and root system. Other species of Phytophthora,
such as P. citricola Sawada and P. parasitica, are reported
to attack P. indica seedlings (Zentmyer, 1980). Another method to test
for the presence of P. cinnamomi uses whole firm fruit of Persea
americana that are placed in soil and flooded with water (Zentmyer et
al., 1960, 1967; Zentmyer and Ohr, 1978). Purplish brown spots develop at
the water line within 4 to 6 days. Recently, however, seedlings of P. indica
have been discovered in the Canary Islands that show resistance to Phytophthora
cinnamomi (Schroeder, 1989).
An
experiment was conducted to determine whether fruit of Persea indica can
be used to test for the presence of Phytophthora in soil. Green,
immature P. indica fruit were collected in February and March, 1991 from
a tree at the University of California Research and Extension Center, Irvine,
California. Field soil, known to be infested with Phytophthora cinnamomi and
P. parasitica, and autoclaved soil for control was placed in Styrofoam
cups and Petri dishes. The suspension in the cups consisted of 20 g soil with
200 mL of distilled water, while the slurry in the Petri dishes was prepared
with 10 g of soil and sufficient water to immerse half of the fruit. The root
system of ten-week-old P. indica seedlings was placed in the soil
suspension, and green, immature P. indica fruit, collected two days
before the experiment started, were placed in the soil slurry. The experiment
was conducted in a greenhouse at 19/29C average day/night temperatures and
natural photoperiod. Development of canker on stems, blackening of roots,
discoloration of fruit, and wilting symptoms of seedlings were monitored. Once
symptoms were visible, affected plant parts and comparable tissue of controls
were plated on selective PVP medium (Tsao and Ocana, 1969) to confirm the
presence of the Phytophthora fungi. A second experiment was conducted in
the same manner as described above, using autoclaved soil as control, P.
cinnamomi-infested field soil, and autoclaved soil that was inoculated with
1 mg per container Phytophthora citricola (P1273 strain), grown on
millet.
Results
Leaf nutrient analysis.
Concentrations of macronutrients in Persea indica leaves were mostly
within the ranges considered adequate for leaves of mature P. americana trees
(Table 1). Boron levels of P. indica were slightly low, and iron levels
were threefold above the highest recommended concentration for P. americana.
Leaves appeared healthy and showed no signs of mineral deficiencies or
toxicities.
Table 1. Nutrient concentration in mature Persea indica leaves and
adequate range of elements in Persea
americana leavesz .
|
|||
Element |
Unit |
P. indica |
P. americana |
Nitrogen |
% |
1.87 |
1.6 – 2.0 |
Phosphorus |
% |
0.14 |
0.08 – 0.25 |
Potassium |
% |
0.87 |
0.75 – 2.00 |
Calcium |
% |
1.35 |
1.00 – 3.00 |
Magnesium |
% |
0.47 |
0.25 – 0.80 |
Sulfur |
% |
0.25 |
0.20 - 0.60 |
Boron |
ppm |
40.00 |
50 – 100 |
Iron |
ppm |
671.00 |
50 – 200 |
Manganese |
ppm |
72.00 |
30 – 500 |
Zinc |
ppm |
41.00 |
30 – 150 |
Copper |
ppm |
13.00 |
5 – 15 |
Chloride |
% |
0.18 |
Excess: 0.25 – 0.50 |
Sodium |
% |
0.087 |
Excess: 0.25 – 0.50 |
z Range of elements for P. americana trees adapted from Leaflet
2024, Division of Agricultural Sciences, University of California - revised
May, 1978. |
Essential oils and alkanes in leaves. The occurrence of essential oils for P. indica and P.
americana cvs. Hass and Fuerte are shown in Figure 2. The ratio of
monoterpenes, sesquiterpenes, and sesquiterpene esters are shown in Table 2. Of
the monoterpenes, cis-ocimene was the most abundant with 1.1%, and of the
sesquiterpenes, beta-caryophyllene (24.0%) was followed by germacrene D (9.1%)
and delta cadinene (9.1%). Ratios were similar for P. indica and the
Hass cultivar, where the majority of oils were sesquiterpenes and sesquiterpene
esters. In contrast, oils from leaves of the cv. Fuerte were mainly comprised
of monoterpenes and contained no sesquiterpene esters at all.
The
order of elution of MS-identified monoterpenes was dimethyl benzene,
alpha-pinene, beta-myrcene, decane, beta-phellandrene, cis-ocimene and
trans-ocimene. That of sesquiterpenes was delta-elemene, alpha-cubebene,
alpha-copaene, beta-bourbonene, beta-caryophyllene, beta-gurjunene, alpha
cadinene, gamma gurjunene, alpha-caryophyllene, gamma-muurolene, germacrene D,
valencene, alpha-muurole, gamma-cadinene, delta-cadinene, beta-calacorene,
alpha-cadinol, and santalol. In 'Hass' and 'Fuerte', beta-pinene, +-limonene,
1,8-cineole, estragole and anethole were additionally identified with the MS.
Hass spectra of alpha-cadinene, caryophyllene, sesquiterpene esters, and
germacrene D that were isolated from essential oils of P. indica are
shown in Figure 3.
Alkanes
of Persea have been analyzed earlier (Scora et a/., 1975) and are
shown in Table 3. P. indica has an alkane composition characteristic of
the subgenus Eriodaphne with high levels of C33 and low
levels of C29. Some of the cultivars of the subgenus Persea that
are hybrids between guatemalensis and drymifolia are
characterized by leaf alkanes comprised of one third of C29 and
nearly one third of C33.
Table 2. Component
percentages of essential oils from leaves of Persea indica and P.
americana cultivars. |
|||
|
Monoterpenes |
Sesquiterpenes (%) |
Sesquiterpene esters (%) |
P. indica |
1.9 |
72.5 |
25.6 |
P. americana cv. Hass |
9.9 |
66.2 |
23.8 |
P. americana cv. Fuerte |
88.1 |
11.9 |
0.0 |
Table
3. Percent alkane composition in leaves of Persea indica and P.
americana cultivars (from Scora et al., 1975). |
|||
Alkane |
P. indica |
P. americana |
P. americana |
C23H48 |
0.5 |
0.0 |
0.1 |
C24H48 |
0.5 |
0.0 |
0.1 |
C25H52 |
1.0 |
0.7 |
0.7 |
C26H54 |
0.9 |
0.6 |
0.4 |
C27H56 |
2.5 |
12.1 |
13.3 |
C28H58 |
1.9 |
1.3 |
1.5 |
C29H60 |
3.1 |
33.6 |
33.3 |
C30H62 |
7.5 |
3.8 |
4.1 |
C31H64 |
3.7 |
15.2 |
12.5 |
C32H66 |
3.8 |
0.3 |
0.1 |
C33H68 |
64.2 |
29.8 |
29.5 |
C34H70 |
5.2 |
0.6 |
1.1 |
C35H72 |
4.9 |
2.0 |
3.4 |
Table
4. Number of Persea indica seedlings and immature fruit that developed
pathogenic symptoms in a soil suspension or slurry with Phytophthora infested
soil, or autoclaved soil as control. |
|||||||
|
Seedlings |
|
Fruit |
||||
Treatment |
Total |
Stem canker |
Black root tips |
Wilting |
|
Total |
Discoloration |
Exp. 1: |
|
|
|
|
|
|
|
Control |
25 |
0 |
0 |
0 |
|
40 |
0 |
P. cinnamomi |
20 |
18 |
20 |
14 |
|
30 |
27 |
P. parasitica |
20 |
17 |
17 |
0 |
|
30 |
12 |
Exp.
II: |
|
|
|
|
|
|
|
Control |
18 |
0 |
0 |
0 |
|
36 |
0 |
P. cinnamomi |
18 |
16 |
18 |
15 |
|
36 |
28 |
P. citricola |
18 |
17 |
18 |
17 |
|
36 |
22 |
Support for purchase of the mass selective detector used in this study
was provided by National Science Foundation grant DIR-8914574.
Literature cited
Axelrod D.I. 1973. History of the Mediterranean
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Castri and H.A. Mooney (eds.) Springer Verlag, New York.
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Scora, R.W., B.O. Bergh, and J.A. Hopfinger. 1975. Leaf
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Schroeder, C.A. and E.F. Frolich. 1955. Avocado
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Schroeder, C.A. 1976. Some useful plants of the
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Schroeder, C.A. 1989. The Laurel or Bay Forest of the
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Zentmyer, G.A., J.D. Gilpatrick, and W.A. Thorn.
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host tissue (Abstr.). Phytopathology 50:87.
Zentmyer, G.A. and H.D. Ohr. 1978. Avocado root rot.
Univ. Calif. Div. Agric. Sci. Leafl. 2440. 15pp.
Zentmyer, G.A., A.O. Paulus, and R.M. Burns. 1967.
Avocado root rot. Calif. Agr. Exp. Stn. Ext. Serv. Circ. 511. Revised. 16pp.
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Figure 2. Gas chromatograph
analysis of essential oils from leaves of Persea indica and Persea americana
cultivars Hass and Fuerte.
Figure 3. Mass spectrum of compounds isolated from essential
oils of Persea indica leaves.