INDUSTRIAL USES AND SUSTAINABLE SUPPLY OF

QUILLAJA SAPONARIA (ROSACEAE)SAPONINS l

RICARDO S A N M A R T I N AND R E I N A L D O B R I O N E S

San Martin, R., and R. Briones. (Department of Chemical and Bioprocess Engineering, Cath-
olic University, Av. Vicuna Mackenna 4860, Santiago--Chile). INDUSTRIALUSES ANDSUSTALN-
ABLESUPPLYOF QUtL~JA SAPONARIA (ROSACEAE)SAPONINS.Economic Botany 53(3):302-311,
1999. The bark of the tree Quillaja saponaria, indigenous to Chile, is one of the major sources
of industrially used triterpenoid saponins. For decades quillaja extracts have been used as
foaming agents in beverages, emulsifiers in foods, wetting agent in photography, etc. Overex-
ploitation of the bark has caused important ecological damage and a shortage of this resource.
However, this can still be remedied by using whole quillaja wood (and not just the bark), for
the production of saponins. This raw material can be obtained in large quantities from pruning
operations, reducing the need to fell trees. This review covers ecological aspects of quillaja
exploitation, as well as a discussion of its novel industrial applications.

USOS INDUSTRIALES Y ABASTECIMIENTO SUSTENABLE DE SAPONINA DE QUtLLAJA SAPONARIA. Za

corteza det drbol Quillaja saponaria, originario de Chile, es una de Ias principales fuentes
industriales de saponinas triterpgnicas. Durante d~cadas los extractos de quillay han sido
usados como espumante en bebidas, emulsificante en alimentos, agente humectante en fotogra-
f(a, etc. La sobre explotaci6n de la corteza ha causado un importante da~o ecol6gico y escasez
de este recurso. Esto adn puede ser remediado usando en forma integral toda la biomasa del
drbol y no s61o la corteza. Esta materia prima es abundante, y se obtiene del raleo de los
bosques existentes, sin necesidad de cortar drboles. Este trabajo cubre aspectos ecoldgicos de
la explotacidn del quillay, y novedosos usos industriales de sus saponinas.
Key Words: Quillaja saponaria; saponins; sustainable production.

Saponins are high molecular weight glyco-
sides, consisting of a sugar moiety linked to a
triterpene or steroid aglycone (Hostettmann and
Martson 1995). Interest in these compounds has
increased significantly in recent years because of
their properties as natural detergents and foam-
ing agents, as well as their applications in cho-
lesterol-reduction and flavor enhancement, their
effect on cardiac activity, hemolytic activity, im-
munostimulating activity and other uses (Waller
and Yamasaki 1996a,b). Among the most im-
portant botanical sources of industrial saponins
is the bark of the tree Quillaja saponaria Mo-
lina, indigenous to Chile. The bark contains
about 5% triterpenoid saponins (Kensil 1996),
which for decades have been used as a surfac-
tant in the production of photosensitized film, as
a foaming agent in beverages and shampoos, a
natural emulsifier in foods and similar uses. Its
Received 22 March 1997, resubmitted; accepted 18
April 1999.
use in foods has been facilitated by the fact that
quillaja extracts (QE) are approved for human
consumption by countries such as the U.S., EC,
and Japan. Other common names for Q. sapo-
naria and its bark are: quillaja tree, soap tree,
soap bark, soap tree bark and quillay (Chilean
common name). Also, because of the historical
trading routes, the bark is known as Panama
bark or Bois de Panama. Our review focuses on
the industrial applications of quillaja saponins,
as well as new process approaches which allow
a stable supply of saponins and a sustainable ex-
ploitation of the resource.
BACKGROUND
First described by Molina (1782), Quillaja sa-
ponaria trees grow naturally in the central and
southern regions of Chile (400 km north and 500
km south of Santiago). Figure 1 shows a pho-
tograph of a quillaja tree over 100 years old. In
the past, extensive areas were covered by quil-
laja forests, but its continuous exploitation and

Economic Botany 53(3) pp. 302-311. 1999

9 1999 by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
1999] SAN MARTIN & BRIONES: QUILLAJA SAPONARIA
Fig. 1. Adult quillaja tree (approximately 100
years old).
the increase of agricultural areas have reduced
significantly its natural population. Virgin quil-
laja forests are now only found in more inac-
cessible areas, particularly in the Andes region,
up to 1500 m a.s.1. The tree grows well in areas
with little rainfall and can withstand long peri-
ods without water. Average densities are about
50 trees per hectare. Natural growth rates are
slow, with an average diameter increase of 0.6-
0.8 cm/year in the central part of Chile (Vita
1974). On good quality soils the tree may grow
much faster, however further research is needed
to determine optimal growing conditions. Repro-
duction is by seeds, but the trees also regenerate
vigorously from stumps that are left in the field
after the trees are felled for the extraction of
bark or when the above ground trunks are killed
by fire. At the present time major part of the
accessible quillaja forests consist of regenerated
trees which grow as shrubs with five or more
large branches per stump. A shrub-type tree
grows to about 10 m, while old trees grown from
seeds may be as tall as 30 m.
The foaming properties of the bark were first
303
recognized by the indigenous people of Chile
(Mapuches) who used aqueous bark extracts for
washing their hair and clothes. For this purpose
the bark was placed in boiling water and allowed
to settle all night before using the extract. In
fact, the local word quillay is derived from the
native Mapuche word "quillean" that means "to
wash." Another common local use is the pre-
vention of moth damage by suspending bark
slabs with clothes. Young pigs are fed with QE
to treat diarrhea.
The bark historically has been exported to the
U.S., Europe and Japan, with no processing
done in Chile. The present market size is about
180-200 tons of QE derived from 1000 tons of
bark exported by Chile each year. The price of
the extracts range from U.S. $12-18 per kg of
non-purified liquid concentrate (about 50% sol-
ids), to over U.S. $100 per kg for more purified
products. A QE market value of about U.S. $7
million per year is a conservative estimate. The
principal producers of QE from bark are Berg-
hausen and Bell Flavors in the U.S., Schmith-
man in Germany and Maruzen Pharmaceuticals
in Japan.
PRODUCTION OF QUILLAJA SAPONINS
FROM THE BARK
Although the wood contains significant quan-
tities, Quillaja saponins are found primarily in
the bark. Using hot water as a solvent, the bark
yields 20% extractives (on a w/w dry basis),
with a saponin content in the extracted solids of
about 20% (Kensil 1996). To obtain bark in eco-
nomical quantities, trees of about 30 to 50 years
old with a diameter of 20-50 cm are felled. To
satisfy present world demand, over 50 000 trees
are felled each year. This activity is regulated by
the Chilean National Forestry Corporation (CO-
NAF). The law dating from 1944, allows ex-
ploitation between March 31 and December 31,
provided a forestry management plan has been
approved by CONAE A maximum of 35% of
the basal tree area can be extracted every 5-10
years from the same site. In practice, exploita-
tion is carded out between September and De-
cember because the bark only peels off easily
during spring growth. Prior to debarking the ex-
ternal part of the bark is removed with knives.
The bark is air-dried in the shade and then it is
tied up in bundles ready for export. Economical
quantities of bark are obtained only from the
main trunk and bigger branches, leaving signif-
304 ECONOMIC BOTANY
glucuronlc qutllalcacid I
ackJ ~ fucose
['~l....J/-, "N , I
I I ~
O ~ ~ . ~ H O o H
9alaclose OH apiose ^Y"~~ rhamnose
Fig. 2. Structureof quillajasaponinQS-21(Recchiaet al. 1995).
icant quantities of biomass to rot in the field
(95% of the weight of the tree, 20 000 tons per
year). Despite increased governmental control
and more strict environmental regulations, the
population of old quillaja trees has decreased
dramatically as a result of continuous exploita-
tion. Because of this, the price of the bark has
increased more than four times in the last five
years.
QUILLAJA SAPONINS DERIVED FROM
W H O L E WOOD
The ecological damage caused by the exploi-
tation of the bark has stimulated research on the
use of the whole quillaja wood (wood with bark,
small branches), as a more stable supply of sa-
ponins. In our laboratory we have shown that
whole wood contains about 8% water soluble
compounds, with a saponin content in the solids
of 20% (determined by reverse-phase HPLC).
The quality of the products derived from whole
wood compares very well with commercial
products derived from bark. Using this new pro-
cess, fewer than 10 000 trees per year (or 4000
tons of whole wood) can satisfy the present
world demand for quillaja saponins. Moreover,
this wood can be obtained entirely from the
pruning of existing bush-type trees that regen-
erate after the exploitation of bark. This raw ma-
terial can be estimated at a minimum of 300 000
tons just in the central part of Chile (Duchens
1998). This new method therefore provides a
sustainable approach to quillaja exploitation,
[VOL. 53
I
tr rln'~ [ ,,u 3,S-dlhydroxy-6-methyl-
~ n octanolc acid
HO ~
OH
arablnose
that can withstand an increase in the world de-
mand without causing ecological damage. Based
on this novel process, a new company is now
operating in Chile (Natural Response S.A., Quil-
ptie), as a joint venture between the Catholic
University of Chile and Desert King Internation-
al, Chula Vista, CA.
CHEMICAL COMPOSITION OF QUILLAJA
SAPONINS
Quillaja saponins are bisdesmosides, that is,
they contain two sugar moieties linked to the
triterpene in positions 3 and 28. The triterpene
has been identified as quillaic acid (Higuchi, To-
kimutsu, and Komori 1988). Monomeric sapo-
nins have a molecular weight determined by size
exclusion HPLC in the range of 1800-2000
(Kensil et al. 1991). Above their critical micelle
concentration (300-800 mg/1), they aggregate in
micelles containing about 50 saponins per mi-
celle (Mitra and Dungan 1997). At least 20 dif-
ferent saponins have been identified from the
bark (van Setten and van de Werken 1996). Im-
portant structural features that distinguish Q. sa-
ponaria saponins from those of other plant spe-
cies are a fatty acid domain and a triterpene al-
dehyde group at position 4. Figure 2 shows the
structure of saponin QS-21 used as an adjuvant
in human vaccines (Recchia et al. 1995).
Though different techniques can be used for the
analysis of saponins, most companies use re-
verse phase HPLC (Kensil et al. 1991). Using
this technique it can be shown that both bark
1999] SAN MARTIN & BRIONES: QUILLAJASAPONAR1A
and wood commercial extracts contain 15-20%
saponins, though the distribution of individual
saponins is somewhat different.
PRODUCTION METHODS AND
COMMERCIAL PRODUCTS
QE are primarily commercialized with very
little purification. Standard liquid products are
prepared using water extraction after the raw
material has been adequately milled. Aqueous
methanol is used by Japanese companies to at-
tain higher extraction yields, although it is
claimed that aqueous extracts are cleaner (Hos-
tettmann and Martson 1995). Following extrac-
tion, the liquid is concentrated by evaporation to
attain the desired concentration of solids. In
some cases it is also necessary to purify the ex-
tract (e.g., contact with activated charcoal, filtra-
tion) to remove compounds that tend to precip-
itate during storage. The final products contain
saponins, protein, tannins, calcium oxalate and
sugars.
Non-refined liquid products contain about 550
g solids/1 (50~ and when diluted with water
they yield a reddish-orange color. Typical pre-
servatives are sodium benzoate (0.5-1 g/l) and
ethanol. Spray-dried products are prepared using
100% crude QE or by mixing the extract with
carriers such as maltodextrins and lactose. More
refined products involve the removal of non-sa-
ponin compounds such as calcium oxalate, sug-
ars, and tannins, which may interfere in terms
of color, chemical interactions, taste, and odor.
For this purpose, purification with polymeric
resins such as Diaion HP-20 and organic sol-
vents have been used (Ogawa and Yokota 1985;
Ogawa and Murakami 1987). An alternative ap-
proach employs the removal of low molecular
weight compounds using diafiltration with ultra-
filtration membranes with molecular cutoffs of
10-30 kDa that retain saponin micelles (Kensil
1991).
The quality of the final extracts is evaluated
in terms of clarity and color in solution, as well
as foaming properties. Traditionally, quality
standards have been developed by major con-
sumers according to their requirements. How-
ever, for novel applications there is a need to
standardize precise analytical procedures, partic-
ularly regarding the determination of saponin
content.
305
REGULATORY ASPECTS OF QUILLAJA
SAPONINS
The toxicology of QE have been studied ex-
tensively in rats (Gaunt, Grasso, and Gangolli
1974; Phillips et al. 1979) and no toxic effects
have been observed even at intake rates as high
as 1500 mg/kg/day (Drake et al. 1982). Based
on these studies, QE are approved as additives
in foods in the U.S. by the FDA under 21 CFR
172.510, FEMA GRAS number 2973. Its current
CAS number is 977002-27-9. In the European
Union they are approved in water-based non-al-
coholic drinks under code E 999. In Japan they
are allowed for human consumption and cos-
metics. The World Health Organization recom-
mends a maximum dally intake of 5 mg of QE/
day-kg human weight.
INDUSTRIAL APPLICATIONS
The scientific and patent literature shows
about 200 references detailing different uses of
QE. What follows is a review of the most rele-
vant scientific articles and patents, with empha-
sis in novel industrial applications.
FOOD AND BEVERAGES
Foaming Agent in Beverages
The most important use of QE is as a natural
foaming agent in beverages (70% of world con-
sumption), such as root beer and soft drinks
(Mukal, Matsuda, and Kanazawa 1993) at con-
centrations of 40-100 ppm. QE are used also in
slush type drinks to improve the ice crystal
structure/morphology and the refreshment value
of the product (Nayyar et al. 1998).
Emulsifier in Food and Beverages
Extensive work performed in Japan has
shown that purified QE (e.g., product Quillaya-
nin C-100, Maruzen Pharmaceuticals, Japan),
are effective emulsifiers in food and beverages.
Oil-in-water emulsions are similar to those ob-
tained with sugar fatty acid esters, with an HLB
value above 12 (Chino and Wako 1992; Mura-
kami 1988, 1996). They can be used also in the
production of microemulsions, an emulsification
technique that minimizes power requirements
(Kudo and Nishi 1992). Some industrial appli-
cations include the production of mayonnaise
(Maeda et al. 1989), enhancement of oil-soluble
flavors for candies (Toya et al. 1994), dissolving
of propolis (Kawal, Hirashita, and Kanae 1994)
306 ECONOMIC BOTANY
and red coloring material (Oono and Higashi-
mura 1995), for soy sauce (Murakami and Wa-
tanabe 1988a) and whipping cream (Murakami
and Watanabe 1988b). Other functional uses are
as antioxidants (Hisayuki and Takashi 1987;
Kooryama and Chiba 1996), leavening agents
(Watanabe et al. 1989) and in health foods (Tas-
hiro 1990).
Production of Low Cholesterol Foods
Quillaja saponins can be used for the produc-
tion of low-cholesterol dairy food products
(Richardson and Jimenez-Flores 1991; Sundfeld,
Krochta, and Richardson 1994), based on the
ability of saponin micelles to form insoluble ag-
gregates with cholesterol, which can be easily
removed by filtration. A recent report details
physicochemical properties of QE that can be
used to optimize this process (Mitra and Dungan
1997). Other patents are related to feeding
chickens with QE, for the production of low-
cholesterol eggs (Yamamoto and Kameda 1993).
NON-FOOD APPLICATIONS
Additives for Animal Feed
Saponin-rich extracts, particularly those de-
rived from Yucca spp, are used extensively as
feed additives to reduce atmospheric ammonia
levels in confinement livestock facilities such as
pigs, broilers (Tymczyma 1996) and cows (Wal-
lace, Arthaud, and Newbold 1994), and to ame-
liorate canine and feline fecal aroma (Lowe and
Kershaw 1997). Recent reports indicate that QE
have similar properties (Ikeda and Suzuki 1995),
and because of their lack of toxicity, their en-
hancement of the synthesis of microbial protein
and their lack of effect on digestibility, they can
be considered as potential feed additives (Mak-
kar and Becker 1996).
Perhaps the most exciting potential for quil-
laja in feeds is as a natural supplement to in-
crease immunoprotection of animals with the re-
sulting reduction in the use of antibiotics. These
immunological properties are unique to quillaja
saponins. Oral administration of QE to mice in-
creases the resistance to rabies infection (Cha-
vali, Barton, and Campbell 1988; Chavali and
Campbell 1987a,b; Maharaj, Froh, and Camp-
bell 1986). In combination with bile powder,
garlic powder, and bacterial cells containing
peptidoglycan, quillaja saponins increase pseu-
dorabies resistance in livestock, poultry and
nursery fish (Yabiki et al. 1993). In fish and
[VOL. 53
shellfish, the oral administration of small doses
of quillaja saponins improves resistance towards
pathogens (Kikuchi et al. 1996; Mabe, Okubo,
and Takahashi 1996; Ninomiya et al. 1995a,b;
Ninomiya et al. 1996). Quillaja saponins have
also been used as delivery systems to enhance
enteric uptake of human gamma globulin in the
tilapian Oreochromis mossambicus (Jenkins,
Harris, and Pulsford 199l) and as a means for
potentiating the immune response elicited by im-
mersion vaccination with commercial enteric
redmouth vaccine (Grayson et al. 1987). It must
be noted that at high doses, toxic effects have
been observed in salmon (Bureau, Harris, and
Young Cho 1998) and shrimp (Chen, Chen, and
Chen 1996).
Agricultural Applications
QE have many potential applications in agri-
culture such as wetting agents, biopesticides and
plant growth stimulants. As wetting agents, their
capacity to reduce surface tension of water is
comparable to that of chemical products (35-40
dyn/cm). This is particularly important for or-
ganic agriculture, where saponins are among the
few products allowed as wetting agents (Organic
Crop Improvement Association 1994). QE can
also be used to increase percolation of hydro-
phobic soils (e.g., product Oasys, Soil Technol-
ogies Corp., Iowa). As biopesticides, saponin-
rich extracts can be used to control nematodes
(Emerson and Crandall 1997), and phytopatho-
genic fungi such as powdery mildew (Emerson
and Crandall 1997). In combination with capsi-
cum oleoresin, they have been formulated as a
safe bio-repellent to control a variety of pests,
such as birds and insects, and fire ants (Wiersma
and Wiersma 1997).
QE also have beneficial effects on seed ger-
mination and plant growth. Research performed
55 years ago in France (Balansard, Pellisier, and
Conil t945), showed that soaking corn seeds in
QE at a concentration of 20 ppm, increased seed
germination and stimulated plant growth. Ben-
eficial effects have also been reported for radish
roots (Fragstein and Buchloh 1981).
Cosmetics
There has long been interest in the use of QE
in the formulation of natural shampoos since this
was its original use by indigenous Chilean peo-
ple. For this purpose, light-colored, low tannin
preparations are preferred to reduce any aller-
1999] SAN MARTIN & BRIONES: QUILLAJASAPONARIA
genic reactions (Poisson 1972, 1974), though
some commercial formulations use non-refined
(more natural) extracts, e.g., Phytologie, France.
Also, in some formulations it is used in combi-
nation with other natural compounds such as
Aloe vera, e.g., Pharmos Aloe Vera, Germany.
In Chile it is extensively used in many shampoos
as a secondary surfactant. A recent work shows
the compatibility of purified QE with chemicals
commonly used in shampoo formulations such
as sodium lauryl sulfate, cocoamidopropyl be-
taine, methylparaben, propylparaben and gluca-
mate (Ceppi 1998). Other applications include
the solubilization of vitamin E (Shirikawa et al.
1994) and essential oils (Murakami 1986; Mu-
ramatsu and Suzuki 1993), hair tonic to reduce
hair loss (D'Aquino Silva 1994) and the pro-
duction of hand creams (Kinekawa et al. 1992).
Photography
QE have been used for decades in the pro-
duction of films as wetting agents to obtain a
uniform distribution of silver halide microcrys-
tals dispersed in gelatin (Ito et al. 1993; Kato et
al. 1994). For this application major photograph-
ic companies (e.g., Konica, Agfa) use purified
QE, with special emphasis in low-tannin, low
color properties.
Wastewater Treatment
QE are used in waste treatment in Japan. It is
claimed that the addition of QE in the treatment
of waste water enhances oxygen transfer and the
degradation of fats and decreases the amount of
sludge and the load of sludge dewatering (Tas-
hiro 1991, 1992, 1995, 1996). Specific applica-
tions include waste water treatment of marine
products and edible meat industries (Nagasaka
1995a,b; Nagasaka and Tashiro 1995).
Miscellaneous Non-Food Applications
Some special non-food applications of quil-
laja saponins are in the preparation of light-
weight composites for building materials (Ton-
yan and Gibson 1993), high-solids aqueous dis-
persions of hydrophobizing agents (Craig 1995),
reduction of sulfuric acid mist in zinc electro-
winning (Mackinnon 1990, 1994), and aerobic
ex situ bioremediation of a chronically polych-
lorobiphenyl (PCB)-contaminated soil (Fava and
Di Gioia 1998).
307
PHARMACEUTICAL USES
Adjuvants for Animal and Human
Vaccines
The use of saponins as adjuvants in animal
vaccines was first tested against foot and mouth
disease (Espinet et al. 1951). It was then shown
that only quillaja saponins are effective adju-
vants, and a fraction that minimized unwanted
side effects was purified (Dalsgaard 1974). This
product, known as Quil-A, is produced by the
Danish firm Superfos and is used commercially
in many animal vaccines. Later it was shown
that Quil-A is a complex mixture of four major
saponins with varying degrees of toxicity, that
can be separated using RP-HPLC (Kensil et al.
1991). Some of the individual saponins, notably
saponin QS-21, showed such low toxicity that
potentially it could be used as adjuvant in the
preparation of human vaccines (Kensil et al.
1991; Kensil 1996; Kensil, Wu, and Soltysik
1995). Experimental trials include the testing of
an HIV-1 vaccine in non-human primates (New-
man et al. 1992), melanoma (Helling et al. 1995)
and Plasmodium falciparum malaria (Stoute et
al. 1997). QS-21 also augments both antibody
and cell-mediated immune responses, suggesting
that this adjuvant could be a valuable component
in subunit vaccines (Wu et al. 1992). Another
important application is in the preparation of im-
munostimulating complexes, ISCOMS (Morein
et al. 1990). ISCOMS are used to present viral
membrane proteins in a multimeric form and are
produced by complexing cholesterol, saponin
and another lipid (Morein et al. 1984). The po-
tential of ISCOMS has been explored in numer-
ous articles and patents and some commercial
vaccines are already produced by this method.
Miscellaneous Pharmaceutical
Applications
Purified quillaja saponins are used to lyse se-
lectively red cells from whole blood prepara-
tions, allowing for leukocyte differential analy-
sis (Ledis et al. 1985), and the preparation of
nucleic acid from white blood cells (Ryder and
Kacian 1993). Modified quillaja saponins such
as saponin DS-1 derived from saponin QS-21
have been used to enhance nasal and ocular de-
livery of insulin (Kensil, Soltysik, and Marciani
1993; Pillion, Amsden, and Kensil 1996) and as
a permeation enhancer for mucosal delivery of
the aminoglycosides, an important class of an-
308 ECONOMIC BOTANY
tibacterial agents used for the treatment of seri-
ous gram-negative infections such as pneumonia
and peritonitis (Recchia et al. 1995). Also an
enteral formulation containing purified quillaja
saponins and oils rich in omega 3 polyunsatu-
rated fatty acids, has been used for treatment of
infection and inflammation (Forse and Chavali
1995).
CONCLUSIONS
Quillaja saponins have a wide range of indus-
trial applications that require an adequate supply
of raw materials. The use of bark as the sole
source of quillaja saponins is no longer viable,
because over-exploitation of natural forests has
caused important ecological damage. The devel-
opment of a new process based on whole quil-
laja wood, as well as plantations, can solve this
problem, allowing the use of quillaja saponins
in novel and interesting industrial applications.
ACKNOWLEDGMENTS
This work was supported by the Chilean Research and Scientific Coun-
cil (FONDEE Project #I 2010). DISCLOSURE: Dr. Ricardo San Martin
is Professor at the Faculty of Engineering of the Catholic University of
Chile, and is President of the Natural Response S.A., a private company
partly owned by the Catholic University.
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, L. D. Barton, and J. B. Campbell. 1988.
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[VOL. 53
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BOOK REVIEW
Wild Plants and Native Peoples of the Four Cor-
ners. William W. Dunmire and Gail Tierney. 1997.
Museum of New Mexico Press, P.O. Box 2087,
Santa Fe, New Mexico 87504. 313 pp. (paperback).
$22.50. ISBN 0-89013-319-0.
It is rare to encounter an ethnobotanical book of any
technical nature successfully written for a general au-
dience without formal training in anthropology and/or
botany. Rarer yet is to find a book written with those
intentions that is also credible and useful to practicing
anthropologists and botanists, especially those inter-
ested in the other's discipline. Dunmire and Tierney
have compiled such a book about plants in the Four
Comers Region of the American Southwest by draw-
ing on an impressive review of primary and technical
literature, interviews, personal observations, and an
excellent collection of illustrations and color photo-
graphs.
Many books on economic plants are structured as
inventories of plants and their uses that are either in-
dependent of cultural affiliation or independent of eco-
logical assemblages. The first chapter introduces the
subject by discussing the concept of place and uses
this as the organizing principle in the book. The phys-
iography and the kind of vegetative communities that
can currently be supported on the land in this region
are described. The following chapters introduce the
cultures inhabiting this area through time (Ancestral
Puebloans, Hopi, Navajo, Utes, and Jicarilla Apache),
and how they used and managed plants. A brief history
of the people is given, their culture is described, and
the kinds of settlements they occupied are discussed
in the context of plant procurement for human use.
Pictures of artifacts utilizing plant materials are inter-
311
laja saponin and carrageenan for cake manufacture.
JP 01 168,227.
Wiersma, C., and J. Wiersma. 1997. Non lethal bio-
repellent compositions. U.S. patent 5,698,191.
Wu, J-Y., B. Gardner, C. Murphy, J. Seals, C. Ken-
sil, and J. Reeehia. 1992. Saponin adjuvant en-
hancement of antigen-specific immune responses to
an experimental HIV-1 vaccine. Journal of Immu-
nology 148: 1519-1525.
Yabiki, T., T. Narahashi, T. Okamoto, and K. ma-
tsuo. 1993. Feed and feed additive for livestock,
poultry and nursery fishes for increasing disease,
pseudorabies resistance. U.S. patent 5,268,357.
Yamamoto, M., and S. Kameda. 1993. Feed addi-
tives containing quillaja saponins and/or yucca sa-
ponins for poultry, egg production with the sapo-
nins, and low cholesterol eggs produced by the
poultry. JP 05 07,462.
spersed throughout the chapters to help illustrate how
plants are used. For each time period, the authors pro-
vide an idea of how environmental change in the re-
gion may have influenced the available plant resourc-
es, and how this may differ from the present "place"
described in the first chapter of the book.
Though hundreds of plants are discussed in general
terms throughout the text, a select group of approxi-
mately 50 plant species common to this place are given
two to four page descriptions in the final chapter of
the book. After a brief introduction to ecozones--re-
inforcing the role environment plays in plant distri-
bution the habitat is mentioned where the selected
plants are most common, the cultures historically uti-
lizing the plant are described as are the useful com-
ponents of the plant and the modes of preparation of
those components. The plants in this chapter are beau-
tifully illustrated and well photographed, but I found
the identification descriptors sparse and potentially dif-
ficult to use. One needs to already know the name of
the plant or flip through nearly 150 pages to find it,
which may not be appropriate for a general audience.
Perhaps this section of the book would be easier to use
if there were a very simplistic key based on leaf type
or flower shape, for example.
One of the highlights of this book is a chart of over
300 plant species known to have been used by the
people occupying the region. The uses, cultural groups
known to use them and references are all in one place.
This book will be a useful tool for ethnobotanists and
those with a general interest of the economic uses of
plants alike.
ROBIN C. D. CURREY
SCHOOL OF NATURAL RESOURCES AND ENVIRONMENT
UNIVERSITY OF MICHIGAN