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Issue
1, June 1999
Biological & Biomedical Sciences
Laticifers in Olona and Ulu: Biological Comparison and Ethnobotanical Significance
Jenny Harvey
Swarthmore College
Abstract
Olona, Touchardia
latifolia, and ulu, Artocarpus altilis, both contain
specialized cells called laticifers that have had significant and
useful roles in Hawaiian ethnobotany. The first part of this study
compared the origins, history, legends, and uses of these two plants.
Research revealed that olona laticifers were used for fishing lines
and other cordage purposes whereas ulu laticifers provided the Hawaiians
with glue and caulking material. In the second part of the study,
laticifers in olona and ulu were examined on a biological level.
Laticifers differed between olona and ulu on a microscopic scale
in distribution, size and shape, cell wall composition, contents,
and location, and differed on a macroscopic scale in latex concentration,
and extraction. This comparison of laticifer characteristics demonstrated
why these specialized cells were used for such different ethnobotanical
purposes in Hawaii.
Introduction
Olona, Touchardia
latifolia, is a woody shrub endemic to Hawaii. Though a member
of the Urticaceae, stinging nettle family, olona is free of stinging
hairs and only occasionally pubescent. Young stems and petioles
are light green and watery, breaking easily; mature stems have a
thin to rough bronze-brown bark. Olona stems grow upright and are
usually unbranched until they fall over. Patches growing wild often
originate from a single stem, which sends new roots into the loose
soil and sprouts branches on the upper side. Large ovate leaves
with three distinct veins grow near the top of the stems (Figure
1a). Leaf morphology such as leaf color, pubescence, and redness
of the midrib, varies from location to location. Plants are dioecious
and the flowers grow in the leaf axils in tight spherical clusters.
The five-parted male flowers have white stamen filaments and inconspicuous
petals. Male inflorescences grow from elongated, branching peduncles.
Female flowers are smaller, five-lobed, without conspicuous perianth,
and grow in smaller, more compact clusters appressed to shorter
peduncles than the male inflorescences. Fruits are bright to dull
orange at maturity, fleshy, and contain a flattened teardrop shaped
achene with a waxy, brittle seed coat and slightly winged margins
(Wagner et al., 1990).
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Figure
1a: Olona, Touchardia latifolia. Waiahole Valley,
Oahu.
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Handy and Handy
(1972) reported that olona grows in "boggy interior valleys" and
"upland areas" and Kamakau (1976) describes olona habitat as rainy,
marshy, mossy, in mountainous areas, often near banana trees. Funk
(1982), during sporadic field studies, confirmed the habit of olona
to grow in wet areas with deep soil and reported that although elevation
varied, olona preferred steep or disturbed terrain.
The ulu or breadfruit
tree, Artocarpus altilis, in the Moraceae (fig) family, grows
to be about 15-20 m, with smooth light colored bark and a hard yellow
wood. Leaves are large and leathery, usually glossy (Figure 1b).
Leaf morphology varies significantly in shape and depth of lobing.
Flowers are dioecious, with both male and female inflorescences
initiating from a terminal bud enclosed by two large stipules. Male
flowers are tightly packed into an elongated inflorescence that
emerges before the female inflorescence, which consists of 1500-2000
white flowers growing in a dense globular cluster terminating a
single, thick peduncle. The female perianths expand and fuse to
form the light green fruit, 12-20cm by 12cm large. The rind of the
fruit is made up of disks, originally the flower surfaces, and thus
ulu is a "multiple fruit". In varieties of Artocarpus found mainly
in the western Pacific Islands, seeds 1-2cm thick are imbedded in
the pulp of the fruit near the enlarged core (Ragone, 1997) but
the variety introduced in Hawaii was seedless. Ulu roots can send
up shoots when cut or damaged. Milky latex flows from wounds and
coagulates upon contact with air.
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Figure
1b: Ulu, Artocarpus altilis. University of Hawaii
at Manoa, Oahu.
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Though hardy, ulu
grows best in consistent rainfall on equatorial lowlands or coastal
plains. Hawaii is the northern limit of ulu, accounting for lower
production than in other Pacific Islands (Ragone, 1997). Ulu fruits
for 8months out of the year in Hawaii, primarily in January, June,
and August.
The purpose of
this study was two-fold. I wanted to first research the myths and
legends associated with olona and ulu, their origins and early cultivation
in Hawaii as well as harvesting techniques, and the ehtnobotanical
uses of each in ancient Hawaii. I also hoped to make a statement
about the current distribution and ecological status of olona and
ulu in Hawaii. Second, I undertook a biological investigation of
laticifers in both plants to help explain their ethnobotanical significance.
For this purpose, I compared microscopic laticifer anatomy, lataex
distibution, and ease of laticifer isolation.
Materials & Methods
My research took
place in two shifts, one spent in the library researching literature
on ethnobotanical aspects of olona and ulu, and a second spent collecting
material and conducting primary investigation. I began by finding
botanical descriptions of olona and ulu in flowering plant manuals
and special reports. I then turned to historical and legendary accounts
of the origins and cultivation of olona and ulu in Hawaii. My sources
included European explorer's journals and books of legends. Investigation
into the ethnobotanical uses of olona laticifers and ulu latex required
a synthesis of many sources, from books specializing in specific
areas of Hawaiian crafts, to personal communications. For current
distribution of these two plants in to personal communications.
For current distribution of these two plants in Hawaii, I relied
on recent field studies, scientific publications, and personal communications.
Background information on laticifers was found in a plant anatomy
textbook as well as scientific journal articles on laticifers in
other plants. All literature research took place in UH Manoa Hamilton
Library and the Bishop Museum library and Archives, as well as among
professors and colleagues in the UH Manoa Botany Department.
For biological
comparison of olona and ulu laticifers, fresh material was collected
from field sites (olona) and UH campus (ulu). Leaves, petioles,
peduncles, and stems of different ages from both plants were cut
into 1cm pieces and immersed in antifreeze fixative of 10% dimethyl
sulfoxide (DMSO), 1%Tween 20, 4%formaldehyde and 0.05M cacodylate
buffer, pH 7.4. Within 24 hours of immersion, plant material was
placed in a vacuum chamber for at least 24 hours. Material was stored
in airtight vials and refrigerated until used. Sections 35-40 micrometers
thick were made in a cryotome at -20° C. Laticifer distribution
in the tissue, size and shape, contents and staining response (cell
wall composition), and location throughout the plant was compared.
Because laticifers are often difficult to recognize in cross section,
I also made longitudinal sections of mature tissues to distinguish
laticifers from other tissue types and compare their length and
branching habits. Areas of high latex concentration were qualitatively
determined by puncturing live plants in different areas and comparing
latex flow.
Laticifers were
isolated from fresh olona stems as per literature information on
harvesting and retting. I chose stalk between 1.5 and 2 cm in diameter
from a male population on the Lower Manoa Cliffs Trail on Roundtop.
I cut the length of the straight stem with a knife, peeled the bast
layer with my fingers and rolled the strips inside out. I soaked
half of the strips in water for 24 hours and half for 72 hours.
I scraped the strips using a plastic hand paddle and a pocket knife
on a smooth stone surface and hung them to air dry.rips were hung
to air dry.
Strips of ulu were
soaked in water but could not be retted.
Legends and myths
Only one legend
was found involving olona: the story of the birth of Kamehameha.
The babe was in danger of being seized by marauders, so was taken
up into the hills of Kohala to be suckled by a woman who was scraping
olona bark. A pile of olona fibers on the floor of her shed concealed
Kamehameha from the enemy search party (Handy, 1972). One chant
was found referring to olona cordage (Abbott, 1992).
Numerous legends
are associated with ulu in Hawaii. To begin, the two morphologies
found in Hawaii are supposed to represent the two sexes: the upright
form, ulu-ku, symbolized the male, and the shrubby form, ulu-ha-papa,
the female (Beckwith, 1970). The best-known origin myth comes from
Tahiti. The god Ku feared his family would starve to death during
a famine, so he asked his wife to slay him and plant his body parts
in the front yard. The next day, a breadfruit tree stood in the
place of his head. After his family had eaten their fill, the breadfruit
continued to produce and feed the rest of the community. Another
origin myth involves two fishermen from Pu'uloa who were blown to
Kane-huna-moku, the "mythical land of Kane" (Handy, 1972), and returned
bearing a fruit tree (Beckwith, 1970) that propagated ai kameha'i,
by the will of the gods (vegetatively) (Handy, 1972). The more rational
story of ulu transport to Hawaii is that Kaha'i, son of Ho'okamali'i,
grandson of Moikeha, brought breadfruit from Upolo, the Hawaiian
word for a Tahitian valley and for Samoa. Kaha'i planted ulu on
the southern coast of Oahu, in the Ewa district near Pu'uloa. In
a later myth, which has many versions, the goddess Haumea rescues
her husband from which has many versions, the goddess Haumea rescues
her husband from kidnappers and passes through a breadfruit tree.
The pursuers try to cut down the ulu tree, but flying splinters
and the milky sap kill the woodcutters. After making sacrifice to
the ulu, the kidnappers are allowed to carve the ulu tree into a
goddess, whom Haumea names Kamehaikana (Westervelt, 1963). One other
ulu myth concerns Kaulula'au, son of Kaka'alaneo and Kanikani'ula
on Maui, who pulls up breadfruit as a prank. He is sent to a haunted
island by his father, but courageously tricks the spirits and saves
himself (Beckwith, 1970). There are a number of sayings also associated
with ulu, the most famous of which is "go for the breadfruit that
oozes gum" which tells a young girl to look for a rich husband (Handy,
1972).
Origins and cultivation
The first discovery
and use of olona by Hawaiians is unknown. Because, as Kamakau mentioned,
there are very few suitable places for large plantings of olona,
it was grown in small patches, though olona has been known to grow
in fields up to 2 acres large (Krauss, 1993). Different types of
patches had different names. A large cultivated patch was olona
kihapai; a garden patch was olona mala; and a wild clump
was opu olona (Summers, 1990). Stalks were encouraged to
grow straight and tall, "as alike as hairs on the head" (Kamakau,
1976), and close to one another to reduce branching (Krauss, 1993).
Lateral branches were removed from upright stems to reduce the number
of holes in the bast fiber (Summers, 1990). Old stalks or toppled
stalks were bent to the ground and covered with soil to promote
rooting and lateral branching to form new stalks. Finally, Hawaiians
weeded frequently, for olona has a tendency to get choked out (Krauss,
1993).
Handy (1972) states
that olona was "one of the few plants native to the islands that
were utilized by the Hawaiians". Actually, ancient Hawaiians used
many endemic, but Handy may have singled out olona because it became
prized as a trade commodity with European explorers. J. Smith (1902)
was excited by the prospects for a plant that produced, in his opinion,
a large fiber yield per plant and per area. Despite this potential,
extensive cultivation of olona ended probably around the time when
the prized cordage was replaced with nylon fishing line and other
synthetic materials.
Ulu originates
from a wild form domesticated in New Guinea and the Bismarck Archipelago
and spread throughout the Pacific Islands by the first Polynesians.
Fijians and Samoans cultivated the seedless variety, A. altilis,
popular in Remote Oceania. Since European discovery in the 1600's,
breadfruit has extended all the way to Africa and across South America
(Ragone, 1997). The seeds lose their viability quickly when dry,
so ulu was most likely transported as root cuttings. However, ulu
is not very resistant to salt water (Ragone, 1997) and requires
a ball of dirt and special care. Handy (1972) commented that the
introduction process couldn't have been easy. The importance of
ulu fruit as a food staple compensated for the trouble, however.
Ulu was most likely introduced to Hawaii from Tahiti (Handy, 1940),
although it may also have arrived with the Islands' first Polynesian
settlers who came from Marquesas. Unlike olona, ulu was never in
high demand in Hawaii, and breadfruit was of secondary food importance
to taro and sweet potato. Therefore, ulu was not incorporated intensively
into mixed cropping, and was found primarily in backyards, as it
is today. Like olona, ulu is cultivated via vegetative propagation.
An ulu tree begins fruiting after three to five years, and will
produce with little upkeep for over to 35 years (Abbott, 1992).
Harvesting and
preparation techniques for olona The
age of the olona stalk made a difference in harvesting time. Plants
around 18 months were preferred for retting because the bast fiber
of older stalks is too knotty. The bast layer was easily stripped
off straight stalks with the fingers (Kamakau, 1976). Strips of
bark were carried over the shoulder or rolled with the bark inside
to flatten them out (Abbott, 1992). Most accounts report that the
strips of bark were softened in water, sometimes running water,
for a day or so, but natives interviewed by Dr. N. Russel (Smith,
1902) omit this step. Sheds were erected near olona patches for
scraping down the harvested bast fiber (Handy, 1972). Both sides
of the bast fiber layer were scraped down on long, thin hardwood
boards called la'au kahi olona or papa olona. The
scraper was usually a sea shell, though later turtle bone from the
costal plate of the shell was used. The scraper was sharpened frequently.
Russel claims that scraping was completed in 1-2 minutes, leading
him to advocate large scale production of the fiber-yielding plant.
However, Summers (1990) implies that practice was required to master
olona scraping: only experts could scrape a few hundred strips in
a day. The white ribbons of fibers (actually laticifers) remaining
were separated with the fingernail, dried, and twisted into cordage
by the women (Handy, 1972)
Uses of olona and
ulu Olona
cordage was prized because of its resistance to kinking and to many
other conditions (Handy, 1972). Fish lines could be few or many-ply,
and were often heavy, but always very strong. One fish line was
reported to be 250-295 meters long (Summers, 1990). Other types
of fish lines often used olona fishhook leaders. Olona cordage was
popular in making nets, including fishnets, 'upena, and carrying
nets, koko. Fine cordage used for the netted matrix in feathered
helmets and capes averaging 0.6mm in diameter. Olona was also used
to repair cracked gourds and canoe paddles, and attach two pieces
of a fishhook together or bind a stone adz to its handle. Other
uses included stretching drumskins over drums and tying the umbilical
cord (Summers, 1990).
The ulu fruit is
ready to eat when the sap oozes out over the skin. Hawaiians preferred
their ulu a little riper than other Polynesians. Hawaiians used
the fruit in a number of ways, though not with as much variety as
found elsewhere in the Pacific. The most common food preparations
included roasting, wrapping in ti leaves and baking in an imu
or earth oven, pounding into poi ulu, or mashing with coconut
cream. Ulu wood was used for house frames, shortboard surfboards,
and could be hollowed into drums (Abbott, 1992). Dried stipules
were used to polish kukui nuts and wooden bowls.
The ulu latex,
kepau, had a variety of uses in Hawaii. In some Pacific islands
at least, the trunk is cut in the morning and the sap collected
later in the day (Ragone, 1997), although Abbott (1992) says the
stem of the fruit was also a source of the milky sap. Kepau served
as a watertight caulking material for canoes, a glue for joining
double gourds used as drums in hula, gluing or coating pieces of
kapa, or setting shark's teeth in war clubs (Krauss, 1993), and
for repairing wooden objects (Abbott, 1992). Ulu latex was spread
on tree branches as birdlime to trap birds for feather gathering.
Children chewed the dried sap like gum. The latex also had medicinal
uses. For ulcers and skin sores, kepau was mixed with pounded, roasted
kukui nuts and pulverized lama wood and pressed into makaloa (Cyperus
laevigatus) fibers for a compress. 'Ea, or thrush, a
fungal disease of the mouth, was treated with a poultice of ulu
leaf buds and 'alaea, or red dirt loaded with ferrous oxide.
This salve was followed by eating a small green fish called aeaea.
Theobald (1976) believes the curative effect of ulu on 'ea is related
to the latex present in leaf buds and young growth.
Current distribution
and status Current
status of olona in Hawaii is unknown. No single researcher or crew
has catalogued wild sites on all islands, although Funk has informally
explored Oahu, Kauai, and Maui, and other scientists have located
specific sites on Oahu. When I explored the areas near Manoa and
Waiahole, I found fewer plants than reported in previous field excursions.
The plants grew tenuously in loose or disturbed soil, frequently
accessible only with great difficulty. Olona was often found growing
among an introduced Rubus species, which tends to outcrowd
less competative plants, a common downfall of endemic species. My
limited field experience leads me to believe that without human
intervention of some sort, olona may soon be rare.
Ulu is often found
in residential backyards, but also grows in arboreta throughout
Hawaii, and on UH Manoa campus. As an introduced species, 'ulu has
never grown wild; trees that appear feral are actually vegetative
ancestors or suckers from trees from an orchard garden agricultural
system. I do not fear for its longevity.
Laticifers
Laticifers, though
sometimes confused with fibers, do not share all characteristics
with them. Whereas fibers are long tapering cells with thick walls
and pits and are often found in groups, laticifers have thin or
irregularly thickened primary cell walls, often contain latex, and
are found in groups or individually, often associated with vascular
tissues (Funk, 1979, Mahlberg, 1993). Laticifers may be articulated
(having sections) or unarticulated (entire), branched or unbranched.
They develop early in the embryo of a plant, and elongate by intrusive
growth that keeps pace with the surrounding tissue. Laticifers are
usually larger in diameter than neighboring cells, and may be pitted.
They are not found in primitive angiosperms and are harder to see
in the root than other parts of plants that contain them (Mahlberg,
1993). Laticifers in olona are unusual for their very thick cell
walls and great strength (Funk, 1979).
Anatomical comparison
Distribution
of laticifers in cross section
Olona laticifers
were found in the cortex of the stem usually clustered near the
phloem (Figure 2a). The laticifers are distributed evenly in the
middle of the cortex, sometimes up to five cells deep, although
rarely in dense clusters (Figure 2b). Laticifers were not present
in the pith.
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Figure
2: Cross section of olona stem showing distribution of laticifers.
a: laticifers (L) distributed in cortex between epidermis
(E) and phloem (P). Stained with toluidine blue; 40x.
b: laticifers (L) may be many cells deep. Viewed with polarizing
filters; 40x.
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Like olona, ulu
laticifers are present in the cortex (Figure 3a), but unlike olona,
they are also present in small numbers in the pith (Figure 3b),
and didn't seem as distinctly associated with vascular tissues.
In the cortex the laticifers are scattered throughout the collenchymous
area under the epidermis as well as in the region with larger-celled
parenchyma (Figure 3c). Fresh sections of ulu mature peduncle contained
large groups of latex-filled cells of various diameter (Figure3d),
indicating that the latex leached out of the preserved material,
making the laticifers hard to find
Size
and shape
Olona laticifers,
when developed, are large cells with unevenly thickened cell walls.
These laticifer cells are usually larger in diameter than their
surrounding cells, irregularly shaped, and have thick primary cell
walls. The cell wall thickens in sections, making the wall look
pitted (Figure 4a). Laticifers are more or less linear, following
the growth of the stalk, as seen in longitudinal section (Figure
4b). Laticifers are unbranched, unarticulated and have tapered ends.
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Figure
4: Size and shape of olona laticifers in stem.
a: cross section showing laticifers (L) with unevenly thickened
cell walls. Courtesy of Dr. Webb.
b: longitudinal section with unbranched, unarticulated laticifers
(arrows). Partial polarizing filter; 25x.
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Ulu laticifers
are not as distinctive as olona laticifers. The cell walls of ulu
laticifers are similar to those of the surrounding parenchyma (Figure
5a). They are unarticulated in the cortex and leaf mesophyll (Figure
5b), but may also be articulated in the pith (Figure 5c). The unarticulated
laticifer cells are generally circular in transverse sections because
they grow by intrusion between cells (Mauseth, 1997). Therefore,
in transverse section, they appear round and surrounded by slight
angular thickenings of the cell wall, perhaps to fill in between
them and neighbor cells (Figure 5d). Longitudinal sections revealed
that some of the laticifers have short branches, and meander through
the tissue rather than parallel to axis of growth (Figure 5e). Finally,
the laticifers are not as long as olona laticifers and have blunt
tips.
Contents
and composition
Olona laticifers
appear to have no specialized contents, although the entire section
was loaded with amyloplasts (Figure 6a). Olona laticifers stained
pink in toluidine blue, indicating pectins and cellulose. They are
also highly birefringent when viewed with polarizing filters (Figure
6b).
a. |
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Figure
6
a: laticifers (L) take up toluidine blue stain. Laticifers
appear empty but surrounding parenchyma is loaded with amyloplasts
(arrows). Partial polarizing filters; 63x.
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b. |
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b:
longitudinal section showing laticifers (arrows) birefringent
in polarizing filters. 25x.
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The contents of
ulu laticifers (unarticulated) and cell columns (articulated) were
primarily either a pale yellow grainy substance with occasional
larger globs or larger globules that stained very dark blue in toluidine
blue (Figure 7a and 5d). To differentiate between latex and other
cellular substances, I investigated the contents of fresh sections.
The white substance oozing from freshly cut ulu was composed of
small uniform grains or globs that sometimes coagulated into larger
globs. The liquid matrix of the exudate coalesced under the microscope
into a netlike mass with a stringy consistency. Only the elongated
cells contained this substance. I concluded that the latex is a
grainy substance with a matrix of rubber components (Mauseth, 1997).
The grainy globs are not birefringent through a polarizing filter
(Figure 7b) and did not take up stain, indicating a substance other
than starch and crystals. Some cell contents in the leaf midrib
stained golden with IKI. Those contents and the dark-staining contents
of cell columns in the vascular system (Figure 7c) may be other
secondary metabolites such as tannins (Mauseth, 1997). Ulu laticifer
cell walls did not take up stains preferentially.
a. |
b. |
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Figure
7: Contents of ulu laticifers.
a: laticifers (arrows) contain uniform grainy substance. Cortex
and vascular tissue of young petiole stained with toluidine
blue; 25x.
b: crystals are birefringent with polarizing filters, but
laticifer contents are not. Section as in 7.a.
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c. |
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c:
articulated laticifers and columns of cells containing dark-staining
substance in pith of woody stem. Stained with toluidine blue;
25x.
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Laticifer
distribution throughout the plant
The young petiole
of olona contained no obvious laticifers. Aside from excessive collenchyma
just under the epidermis and the vascular bundles, all cell walls
are about the same thickness. In the young petiole from ulu, the
laticifers are scattered throughout the section (see Figure 7).
In a cross section
of the young olona stem 1cm in diameter laticifers were easily distinguished
in the cortex between the epidermis and the phloem. A longitudinal
section of this stem tissue revealed long laticifer cells in different
stages of maturity. A young woody ulu stem 3/4cm diameter contained
the laticifers observed in the petiole, but the pith also displayed
other cells with contents mentioned above.
Olona appeared
to have no laticifers in the midrib or blade of the mature leaf.
Ulu, however, had a complex distribution of laticifers in these
organs. As in the mature stem, latex-containing cells were slightly
more frequent near vascular tissues of the leaf midrib, i.e. near
the phloem fibers if in the cortex, and near the xylem if in the
pith. Elongated, branched laticifers were present in the mesophyll
of the leaf blade (Figure 8a). Some large latex-containing cells
varying in size and shape were hanging suspended from the mesophyll
the large air spaces between vascular bundles (Figure 8b).
a. |
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Figure
8: Distribution of laticifers throughout the ulu plant.
a: unarticulated, meandering and branched laticifers (arrows)
in mature leaf. 32x.
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b. |
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b:
latex-containing cells (arrows) suspended in air spaces of
mature leaf. 63x.
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In very mature
ulu tissues, e.g. the outer layers of a branch 7.5cm in diameter,
the heavy distribution of laticifers was especially evident in a
thick section (Figure 8c). Though not examined in this investigation,
the fruit should also contain laticifers because the rind oozes
latex during the ripening process.
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Figure
8c: frequent distribution of laticifers (circled) in very
woody stem. Thick section.
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Latex distribution
within plant
Qualitative comparison
of relative latex concentration among different plant parts within
ulu as conducted between 7:30 and 9:00 am in full sun in mid-October
on two plants on UH Manoa campus. When I broke a mature leaf off
near the stem, the wound oozed 24 drops of a whitish fluid (Figure
9). Slicing off the stem apical meristem (immature stipules and
terminal bud), produced 22 drops in the same amount of time. When
examined later, the white substance had congealed on the cut surface
to the consistency of rubber cement. I then made a slice 2cm long
into a trunk 17cm in diameter 1m above the ground. The cut immediately
oozed the whitish fluid. As it dribbled down the trunk, a thick
opaque white component separated out from the milky liquid part
and stuck to the bark in clumps. For some time one corner of the
cut continued to ooze a thick white sap that emerged in spurts like
bubbles and congealed upon contact with air. I scraped my knife
tip across the length of the cut occasionally, removing a tan-white
film with a consistency resembling rubber cement. I set it aside
as I continued to remove it; the glob remained workable and sticky,
like chewing gum. I concluded that younger tissue has higher liquid:congealant
ratio in its latex, although even the rather clear liquid component
was tacky.
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Figure 9: ulu latex
dripping from broken petiole. |
A cut 2m above
ground in an older tree 116cm in diameter oozed about the same amount
of sap as the smaller trunk, but the oozing did not persist. The
amount of white component was extruded in higher proportion than
in the younger trunk, however. Simply inserting my knife 1/2-1cm
into the trunk yielded much more latex than the shallow cuts. Letting
the latex run down the bark or constantly scraping it off seems
an inefficient means of collecting the sap; I wondered what was
the traditional collection method. I made a makeshift collection
method by inserting a strip of palm frond in the cut. The sap quickly
flowed down and off the frond and could be easily collected. Latex
from the deep cuts was very runny and didn't congeal to the same
thickness as that from the younger trunk, but was still very tacky.
Old wounds in the
mature tree had oozed latex some time earlier and dried into brittle
grayish globs. I scraped off the thin gray bark and thin red underlayer
from the older tree to reveal the pulpy orange wood. A thin sheen
of latex slowly collected on the scraped area. When I cut 1/2cm
further into the wood the latex dripped out quickly. I concluded
that the sap-containing cells or ducts are located in the mature
woody tissues under the bark.
Finally, I observed
the flow of latex from a mature peduncle at 4:00pm in full sun on
November 13. When I cut it near the top of the fruit, the latex
spurted out with more force than any other organ I had tested.
Acetone was the
best solvent for cleaning hands and instruments after dealing with
ulu and its latex.
Olona tissues exude
a clear watery liquid when wounded although the flow is not maintained
after the initial few drops. Field observations indicate that young
and herbaceous tissues such as petioles and leaves contain more
fluid than woody tissues in the middle and mature stem region. The
exudate is not sticky and does not harden or congeal upon exposure
to air, but rather resembles contents of vascular tissues in other
plants.
Isolation of
laticifers
Although the bast
layer was easy to remove from olona (Figure 10a), preliminary extraction
of laticifer strands yielded disappointing results. During scraping,
I alternately used the plastic paddle angled toward me, and the
knife, angled away from me (Figure 10b). I found that it helped
to keep the strip and area very wet with water. The leaf scars were
difficult to scrape over, and created curves and holes in otherwise
straight grain of the bast fibers. I was left with a greenish brown
band of fibers that dried to a hard, curly, inseparable strip.
> a. |
b. |
|
Figure 10: Isolation
of laticifers from olona.
a: stripping the bast layer.
b: scraping the bast layer. |
The strips that
had soaked for 72 hours, however, were more easily retted. The strips
looked and smelled a little fermented when I removed them from the
water, and the outer bark and inner pithy layer slid off easily
using only the plastic paddle. Much of the fibrous material came
off along with the bark and pith, so that what fibrous material
remained was a very thin layer only about 1/2 the original length
of the strips. The fibrous material was pure white as mentioned
in the literature, and could be separated into individual strands
or cells using only my fingernail. The strands were very strong,
resisted tearing and breaking, and clung to one another as if their
surfaces were rough. Once isolated, the strands hung straight and
dried almost instantly without shrinking. After several days of
air drying, the strands, which were finer than human hair, were
still pliable, separable, and lightweight, and did feel slightly
rough when pulled between two fingers. Of these strands, which seemed
more suitable for cordage than those from my first attempt, the
average length was 27cm, but some individual strands were over 45cm
long.
Ulu outer layer
was not easily distinguishable or separable from the rest of the
woody growth in the trunk. Soaking in water did not soften the material
at all: after three days the bast layer was still tough and cohesive.
Scraping merely broke the woody fibrous tissue or pulled it apart
in large clumps. The outer bark did not come off easily and the
layer was so thick (0.5cm) that no laticifer layer could be discerned
nor individual strands isolated.
Discussion
Results of this investigation, summarized in Table 1, leave little
question about why the laticifers of ulu and olona were used for
such vastly different purposes by ancient Hawaiians. Olona laticifers
are long and flexible with thick cell walls, giving them strength
and versatility for the uses mentioned above. The strands are somewhat
rough due to irregularly thickened cell walls visible under the
microscope, making them ideal for twisting together. Ulu laticifers
are much shorter and have cell walls no thicker or stronger than
regular parenchyma cells. They might have made a crude tapa well
enough, but would never have withstood tensions like those applied
to olona nets and fish lines. The olona laticifers are conveniently
located in the outer layer of the stem. Tissue layers between the
phloem fibers and the epidermis are often collectively called the
bast fibers. In olona the bast fibers are actually the laticifers.
Ulu laticifers are distributed throughout the stem; more material
would be wasted trying to isolate the laticifers than would be yielded
in strands. Olona laticifers contain no obvious latex, posing little
problems when extracting the strands, whereas ulu latex makes the
bast fiber region too sticky for easy extraction. The latex would
work as excellent birdlime, however, and after trying to wash the
latex from my hands, I understand why it was used as a waterproof
caulking.
From a biological
perspective, the distribution of laticifers in these plants may have
some significance. Olona laticifers take the place of phloem fibers
found in many other plants, yet they do not offer as much support
as lignified fibers. Perhaps the laticifers merely protect the vascular
tissues from wounding. Another function could be to maintain flexibility
as the stems fall over before sending down new roots. If the stems
were brittle or lignified, they would crack or break when the loose
soil in their habitats slumps. Ulu laticifers are not strong enough
to offer support or protection, but their latex may have evolved as
a wound sealant, and may also have a role in herbivore deterrence.
Olona
cordage has been replaced by nylon and other synthetics in modern
marine technology; but its incredible strength has not lessened but
by comparison. Olona could still be used in communities or cultures
that produce their own cordage, for it is superior to hemp and agave
(Webb, pers. comm). The strands are also rather soft and one can imagine
their use in clothing. Olona has been forgotten, however, except in
places remote from modern society where it is still cultivated on
a small scale (Yoshinaga, 1997). Minor but constant weeding and clearing
would probably increase the longevity of olona patches currently growing
wild. If more wild patches are located and tended, or plants are reared
in nurseries and outplanted, olona could become a significant source
of fiber products in the future. Ulu also has potential for resin
products on the natural product market. I don't know if any steps
are being taken in these directions; perhaps the replacement of synthetics
with organic products is a possibility for future investigations.
| |
Olona
|
Ulu
|
| Major
uses |
Laticifer
cells: fishing line, matrix for feather capes, nets,
cordage |
Latex
only: birdlime, caulking |
| Distribution |
In
cortex near phloem |
Throughout
cortex and pith |
| Size
and shape |
Cross
section: larger than surrounding cells; irregularly
shaped
Longitudinal:
unarticulated, parallel to axis, unbranched, tapered
ends, long |
Cross
section: same size as surrounding cells, circular
Longitudinal:
unarticulated, tubular, meandering, sometimes branched,
with blunt ends; in pith occasionally articulated
(columns of cells) |
| Contents
|
Clear
liquid to none |
Yellowish
grainy substance in watery medium; very tacky, congeals
upon air exposure |
| Cell
wall composition |
Cell
wall thickened irregularly; pectin and cellulose
stains pink in toluidine blue |
Cell
wall no more thickened than surrounding parenchyma;
no stain preference |
| Location
in plant |
Young
to mature and woody stem |
Petiole,
stems of all ages, leaves, peduncle, fruit, woody
growth |
| Latex
concentration |
Mostly
in young and herbaceous tissues |
Peduncle
> mature branch > young branch > woody
tissues, leaves, fruit etc. |
| Isolation |
Easy
to strip bast layer; easily softened in water; quickly
scraped, dried and separated |
Difficult
to isolate bast layer and individual laticifers;
does not soften in water |
|
|
Table
1:
Comparison of ethnobotanical uses and biological characteristics
of olona and ulu laticifers. |
Acknowledgements
I would like to thank Dr.
David Webb for his extensive help in all aspects of field and laboratory
work and the use of his microscopy and computer equipment. I thank
Dr. Isabella Abbott for her direction and understanding over the course
of the research and writing process. I also thank graduate students
Michelle Stephens and Nathan Nishimura for their advice and support.
Thanks to the staff of the Hawaiian/Pacific Collection at University
of Hawaii Hamilton Library for their help and patience; Dr. Paul Mahlberg
of Indiana University for laticifer insight; Judy Harvey and Darren
Sandvig for specimen collection assistance.
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Copyright © 1999 by Jenny Harvey and JYI. All rights reserved.
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