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    Industrial Uses and Sustainable Supply of Quillaja saponaria (Rosaceae) SaponinsAuthor(s): Ricardo San Martn and Reinaldo BrionesReviewed work(s):Source: Economic Botany, Vol. 53, No. 3 (Jul. - Sep., 1999), pp. 302-311Published by: Springer on behalf of New York Botanical Garden PressStable URL: http://www.jstor.org/stable/4256204 .

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    INDUSTRIAL USES AND SUSTAINABLE SUPPLY OFQUILLAJAAPONARIAROSACEAE) SAPONINS'RICARDO SAN MARTfN AND REINALDO BRIONES

    San Martin, R., and R. Briones. (Department f ChemicalandBioprocessEngineering,Cath-olic University, Av. Vicufia Mackenna 4860, Santiago-Chile). INDUSTRIALUSES AND SUSTAIN-ABLE SUPPLY OF QUILLAJA 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 sourcesof industrially used triterpenoid saponins. For decades quillaja extracts have been used asfoaming 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), forthe production of saponins. This raw material can be obtained in large quantities from pruningoperations, reducing the need to fell trees. This review covers ecological aspects of quillajaexploitation, as well as a discussion of its novel industrial applications.USOS INDUSTRIALESY ABASTECIMIENTO USTENABLEDE SAPONINA DE QUILLAJA APONARIA. acorteza del drbol Quillaja saponaria, originario de Chile, es una de las principales fuentesindustriales de saponinas triterpenicas. Durante deadas los extractos de quillay han sidousados como espumante en bebidas, emulsificante en alimentos, agente humectante en fotogra-fia, etc. La sobre explotacion de la corteza ha causado un importante danio ecol6gico y escasezde este recurso. Esto ain puede ser remediado usando en forma integral toda la biomasa deldrbol y no s6lo la corteza. Esta materia prima es abundante, y se obtiene del raleo de losbosques existentes, sin necesidad de cortar drboles. Este trabajo cubre aspectos ecol6gicos dela explotaci6n 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 atriterpene or steroid aglycone (Hostettmann andMartson 1995). Interest in these compounds hasincreased significantly in recent years because oftheir properties as natural detergents and foam-ing agents, as well as their applications in cho-lesterol-reduction and flavor enhancement, theireffect on cardiac activity, hemolytic activity, im-munostimulating activity and other uses (Wallerand Yamasaki 1996a,b). Among the most im-portant botanical sources of industrial saponinsis the bark of the tree Quillaja saponaria Mo-lina, indigenous to Chile. The bark containsabout 5% triterpenoid saponins (Kensil 1996),which for decades have been used as a surfac-tant in the production of photosensitized film, asa foaming agent in beverages and shampoos, anatural emulsifier in foods and similar uses. Its

    IReceived 22 March1997,resubmitted;ccepted18April 1999.

    use in foods has been facilitatedby the fact thatquillajaextracts (QE) are approvedfor humanconsumptionby countriessuch as the U.S., EC,and Japan.Othercommon names for Q. sapo-naria and its bark are: quillajatree, soap tree,soap bark, soap tree barkand quillay (Chileancommon name). Also, because of the historicaltrading routes, the bark is known as Panamabark or Bois de Panama.Our review focuses onthe industrialapplications of quillajasaponins,as well as new process approacheswhich allowa stablesupply of saponinsand a sustainable x-ploitationof the resource.

    BACKGROUNDFirstdescribedby Molina(1782), Quillajasa-ponaria trees grow naturally n the centralandsouthern egions of Chile(400 km northand500km south of Santiago).Figure 1 shows a pho-tographof a quillaja tree over 100 years old. Inthe past, extensive areaswere covered by quil-laja forests, but its continuousexploitationand

    Economic Botany 53(3) pp. 302-311. 1999?D1999 by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

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    1999] SAN MARTIN& BRIONES:QUILLAJA APONARIA 303

    Fig. 1. Adult quillaja tree (approximately100yearsold).

    the increase of agricultural areas have reducedsignificantly 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.l. The tree grows well in areaswith little rainfall and can withstand long peri-ods without water. Average densities are about50 trees per hectare. Natural growth rates areslow, with an average diameter increase of 0.6-0.8 cm/year in the central part of Chile (Vita1974). On good quality soils the tree may growmuch faster, however further research is neededto determine optimal growing conditions. Repro-duction is by seeds, but the trees also regeneratevigorously from stumps that are left in the fieldafter the trees are felled for the extraction ofbark or when the above ground trunks are killedby fire. At the present time major part of theaccessible quillaja forests consist of regeneratedtrees which grow as shrubs with five or morelarge branches per stump. A shrub-type treegrows to about 10 m, while old trees grown fromseeds may be as tall as 30 m.The foaming properties of the bark were first

    recognized by the indigenous people of Chile(Mapuches)who used aqueous bark extractsforwashing their hair andclothes. For this purposethe barkwas placedin boilingwaterand allowedto settle all night before using the extract. Infact, the local word quillay is derivedfrom thenativeMapucheword "quillean" hatmeans "towash." Another common local use is the pre-vention of moth damage by suspending barkslabs with clothes. Young pigs are fed with QEto treat diarrhea.The barkhistoricallyhas been exportedto theU.S., Europe and Japan, with no processingdone in Chile. The presentmarketsize is about180-200 tons of QE derived from 1000 tons ofbarkexported by Chile each year.The price ofthe extracts range from U.S. $12-18 per kg ofnon-purified iquid concentrate about 50% sol-ids), to over U.S. $100 per kg for morepurifiedproducts.A QE market value of about U.S. $7million per year is a conservative estimate. Theprincipalproducersof QE from bark are Berg-hausen and Bell Flavors in the U.S., Schmith-man in Germanyand MaruzenPharmaceuticalsin Japan.

    PRODUCTION OF QUILLAJA SAPONINSFROM THE BARKAlthoughthe wood containssignificantquan-tities, Quillaja saponins are found primarily nthe bark.Using hot water as a solvent, the barkyields 20% extractives (on a w/w dry basis),with a saponincontent in the extractedsolids ofabout20%(Kensil 1996). To obtainbark n eco-nomicalquantities, rees of about 30 to 50 yearsold with a diameterof 20-50 cm are felled. Tosatisfy presentworlddemand,over 50 000 trees

    are felled each year.This activityis regulatedbythe Chilean NationalForestryCorporation CO-NAF). The law dating from 1944, allows ex-ploitationbetween March31 and December31,provideda forestry managementplan has beenapproved by CONAE A maximum of 35% ofthe basal tree area can be extractedevery 5-10years from the same site. In practice, exploita-tion is carried out between Septemberand De-cember because the bark only peels off easilyduring spring growth.Priorto debarking he ex-ternalpartof the bark is removed with knives.The bark is air-dried n the shade and thenit istied up in bundlesreadyfor export.Economicalquantities of bark are obtained only from themain trunk and bigger branches, eaving signif-

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    304 ECONOMICBOTANY [VOL. 53

    glucuronic quillak-cidacld; lucosexylose OHHylosemH > H 3,5-dlhydroxy-6-methyl-octanolc acid

    H ~ ~ ~ ~ ~ H mH ~ ~ ~ ~ ~ ~ ~ ~~~~Oalaclose OH apOse yls

    arabinoseFig. 2. Structure f quillajasaponinQS-21 (Recchia et al. 1995).

    icant quantities of biomass to rot in the field(95% of the weight of the tree, 20 000 tons peryear). Despite increased governmentalcontroland more strict environmentalregulations,thepopulation of old quillaja trees has decreaseddramaticallyas a result of continuousexploita-tion. Because of this, the price of the barkhasincreased more than four times in the last fiveyears.QUILLAJA SAPONINS DERIVED FROMWHOLE WOODThe ecological damagecaused by the exploi-tation of the barkhas stimulated esearchon theuse of the whole quillajawood (wood withbark,small branches),as a more stable supply of sa-ponins. In our laboratorywe have shown thatwhole wood contains about 8% water solublecompounds,with a saponincontent n the solidsof 20% (determinedby reverse-phaseHPLC).The qualityof the productsderivedfromwholewood compares very well with commercialproductsderivedfrombark.Using this new pro-cess, fewer than 10 000 trees per year (or 4000tons of whole wood) can satisfy the presentworld demandfor quillajasaponins.Moreover,this wood can be obtained entirely from the

    pruningof existing bush-typetrees that regen-erateafterthe exploitationof bark.Thisrawma-terialcan be estimatedat a minimumof 300 000tons just in the centralpart of Chile (Duchens1998). This new method thereforeprovides asustainable approach to quillaja exploitation,

    that can withstandan increase in the world de-mandwithoutcausingecological damage.Basedon this novel process, a new company is nowoperating n Chile (NaturalResponseS.A., Quil-pue), as a joint venture between the CatholicUniversityof Chile and DesertKingInternation-al, ChulaVista, CA.CHEMICAL COMPOSITION OF QUILLAJASAPONINS

    Quillaja saponins are bisdesmosides, that is,they contain two sugar moieties linked to thetriterpene n positions 3 and 28. The triterpenehas been identifiedas quillaic acid (Higuchi,To-kimutsu, and Komori 1988). Monomericsapo-nins have a molecularweightdeterminedby sizeexclusion HPLC in the range of 1800-2000(Kensil et al. 1991). Above theircriticalmicelleconcentration300-800 mg/l), they aggregate nmicelles containing about 50 saponinsper mi-celle (MitraandDungan 1997). At least 20 dif-ferent saponins have been identified from thebark(van Settenand van de Werken1996). Im-portant tructuraleatures hatdistinguishQ. sa-ponaria saponinsfromthose of otherplantspe-cies are a fatty acid domainand a triterpeneal-dehyde groupat position 4. Figure2 shows thestructureof saponinQS-21 used as an adjuvantin human vaccines (Recchia et al. 1995).Thoughdifferenttechniquescan be used for theanalysis of saponins, most companies use re-verse phase HPLC (Kensil et al. 1991). Usingthis techniqueit can be shown that both bark

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    1999] SAN MARTiN & BRIONES: QUILLAJASAPONARIA 305

    and wood commercialextractscontain 15-20%saponins, though the distributionof individualsaponinsis somewhatdifferent.PRODUCTION METHODS ANDCOMMERCIAL PRODUCTS

    QE are primarilycommercialized with verylittle purification.Standard iquid productsarepreparedusing water extraction after the rawmaterial has been adequatelymilled. Aqueousmethanolis used by Japanesecompaniesto at-tain higher extraction yields, although it isclaimed thataqueousextracts are cleaner(Hos-tettmannand Martson1995). Following extrac-tion, the liquidis concentrated y evaporationoattain the desired concentration of solids. Insome cases it is also necessaryto purifythe ex-tract(e.g., contactwith activatedcharcoal, iltra-tion) to remove compoundsthat tend to precip-itate duringstorage.The final productscontainsaponins,protein,tannins,calcium oxalate andsugars.Non-refined iquidproductscontainabout550g solidsfA50?Brix)and when diluted with waterthey yield a reddish-orange olor. Typical pre-servatives are sodium benzoate (0.5-1 g/l) andethanol.Spray-dried roductsarepreparedusing100% crude QE or by mixing the extractwithcarriers uch as maltodextrins ndlactose.Morerefinedproducts nvolve the removal of non-sa-ponin compounds such as calciumoxalate, sug-ars, and tannins, which may interfere in termsof color, chemical interactions, aste, and odor.For this purpose, purificationwith polymericresins such as Diaion HP-20 and organic sol-vents have been used (OgawaandYokota1985;OgawaandMurakami1987). An alternativeap-proach employs the removal of low molecularweight compoundsusing diafiltrationwith ultra-filtrationmembraneswith molecularcutoffs of10-30 kDa that retainsaponinmicelles (Kensil1991).The quality of the final extracts is evaluatedin terms of clarityandcolor in solution,as wellas foaming properties. Traditionally, qualitystandardshave been developed by major con-sumers according to their requirements.How-ever, for novel applications there is a need tostandardize reciseanalyticalprocedures,partic-ularly regarding the determinationof saponincontent.

    REGULATORY ASPECTS OF QUILLAJASAPONINSThe toxicology of QE have been studiedex-tensively in rats (Gaunt,Grasso, and Gangolli

    1974; Phillips et al. 1979) and no toxic effectshave been observedeven at intakeratesas highas 1500 mg/kg/day (Drake et al. 1982). Basedon these studies, QE are approvedas additivesin foods in the U.S. by the FDA under 21 CFR172.510, FEMA GRAS number2973. ItscurrentCAS number is 977002-27-9. In the EuropeanUnion they are approved n water-basednon-al-coholic drinks under code E 999. In Japantheyare allowed for human consumptionand cos-metics. The WorldHealth Organization ecom-mends a maximumdaily intakeof 5 mg of QE/day-kg humanweight.INDUSTRIAL APPLICATIONS

    The scientific and patent literature showsabout 200 referencesdetailingdifferent uses ofQE. Whatfollows is a review of the most rele-vant scientific articlesandpatents, with empha-sis in novel industrialapplications.FOOD AND BEVERAGESFoaming Agent in Beverages

    The most importantuse of QE is as a naturalfoamingagentin beverages (70%of worldcon-sumption), such as root beer and soft drinks(Mukai,Matsuda,and Kanazawa1993) at con-centrationsof 40-100 ppm. QE areused also inslush type drinks to improve the ice crystalstructure/morphologynd the refreshment alueof the product(Nayyaret al. 1998).Emulsifier in Food and Beverages

    Extensive work performed in Japan hasshown thatpurifiedQE (e.g., productQuillaya-nin C-100, Maruzen Pharmaceuticals,Japan),are effective emulsifiers n food andbeverages.Oil-in-wateremulsions are similarto those ob-tained with sugarfatty acid esters,with an HLBvalue above 12 (Chino and Wako 1992; Mura-kami 1988, 1996). They can be used also in theproductionof microemulsions,an emulsificationtechnique that minimizes power requirements(Kudo and Nishi 1992). Some industrialappli-cations include the production of mayonnaise(Maedaet al. 1989), enhancementof oil-solubleflavorsfor candies (Toyaet al. 1994),dissolvingof propolis(Kawai,Hirashita,and Kanae 1994)

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    306 ECONOMICBOTANY [VOL. 53and red coloring material (Oono and Higashi-mura 1995), for soy sauce (Murakamiand Wa-tanabe 1988a) and whippingcream (MurakamiandWatanabe1988b). Otherfunctionaluses areas antioxidants (Hisayuki and Takashi 1987;Kooryama and Chiba 1996), leavening agents(Watanabe t al. 1989) andin health foods (Tas-hiro 1990).

    Production of Low Cholesterol FoodsQuillajasaponinscan be used for the produc-tion of low-cholesterol dairy food products(Richardson nd Jimenez-Flores1991;Sundfeld,Krochta,and Richardson 1994), based on theability of saponinmicelles to form insolubleag-

    gregates with cholesterol, which can be easilyremoved by filtration. A recent report detailsphysicochemicalpropertiesof QE that can beusedto optimizethisprocess(MitraandDungan1997). Other patents are related to feedingchickens with QE, for the productionof low-cholesteroleggs (YamamotoandKameda1993).NON-FOOD APPLICATIONS

    Additives for Animal FeedSaponin-richextracts, particularly hose de-rived from Yucca spp, are used extensively asfeed additives to reduce atmosphericammonialevels in confinement ivestock facilities such aspigs, broilers(Tymczyma1996) and cows (Wal-lace, Arthaud,and Newbold 1994), and to ame-lioratecanine and feline fecal aroma(Lowe andKershaw1997). Recentreports ndicatethatQEhave similarproperties IkedaandSuzuki1995),and because of their lack of toxicity, their en-hancementof the synthesis of microbialprotein

    andtheirlack of effect on digestibility, hey canbe considered as potential feed additives(Mak-kar andBecker 1996).Perhapsthe most exciting potentialfor quil-laja in feeds is as a naturalsupplementto in-creaseimmunoprotection f animalswith there-sultingreduction n the use of antibiotics.Theseimmunologicalpropertiesare unique to quillajasaponins. Oraladministration f QE to mice in-creases the resistance to rabies infection (Cha-vali, Barton, and Campbell 1988; Chavali andCampbell 1987a,b; Maharaj,Froh, and Camp-bell 1986). In combination with bile powder,garlic powder, and bacterial cells containingpeptidoglycan,quillaja saponins increase pseu-dorabies resistance in livestock, poultry andnursery fish (Yabiki et al. 1993). In fish and

    shellfish, the oral administration f small dosesof quillajasaponins mprovesresistance owardspathogens(Kikuchi et al. 1996; Mabe, Okubo,and Takahashi1996; Ninomiya et al. 1995a,b;Ninomiya et al. 1996). Quillaja saponins havealso been used as delivery systems to enhanceentericuptakeof humangammaglobulin in thetilapian Oreochromis mossambicus (Jenkins,Harris,and Pulsford 1991) and as a means forpotentiating heimmuneresponseelicitedby im-mersion vaccination with commercial entericredmouthvaccine (Graysonet al. 1987). It mustbe noted that at high doses, toxic effects havebeen observed in salmon (Bureau,Harris,andYoungCho 1998) and shrimp Chen,Chen,andChen 1996).

    Agricultural ApplicationsQE have many potentialapplications n agri-culturesuch as wettingagents,biopesticidesandplantgrowthstimulants.As wettingagents,theircapacity to reduce surface tension of water iscomparable o that of chemicalproducts 35-40dyn/cm). This is particularly mportant or or-ganic agriculture,wheresaponinsareamongthefew productsallowed as wetting agents(OrganicCrop ImprovementAssociation 1994). QE canalso be used to increase percolation of hydro-phobic soils (e.g., productOasys, Soil Technol-ogies Corp., Iowa). As biopesticides, saponin-rich extracts can be used to control nematodes(Emersonand Crandall1997), and phytopatho-genic fungi such as powdery mildew (Emersonand Crandall1997). In combinationwith capsi-cum oleoresin, they have been formulatedas asafe bio-repellent to control a variety of pests,

    such as birdsandinsects, andfireants(WiersmaandWiersma1997).QE also have beneficialeffects on seed ger-minationandplantgrowth.Researchperformed55 yearsago in France(Balansard,Pellisier,andConil 1945), showed thatsoaking corn seeds inQE at a concentration f 20 ppm,increasedseedgerminationand stimulatedplant growth. Ben-eficial effects have also been reported or radishroots (Fragsteinand Buchloh 1981).CosmeticsTherehas long been interest n the use of QEin theformulation f natural hampoossince thiswas its original use by indigenousChileanpeo-ple. For this purpose, light-colored, ow tanninpreparationsare preferred o reduce any aller-

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    1999] SAN MARTIN& BRIONES:QUILLAJAAPONARIA 307genic reactions (Poisson 1972, 1974), thoughsome commercial formulationsuse non-refined(morenatural)extracts,e.g., Phytologie,France.Also, in some formulations t is used in combi-nation with other naturalcompounds such asAloe vera, e.g., Pharmos Aloe Vera, Germany.In Chileit is extensivelyused in manyshampoosas a secondarysurfactant.A recent work showsthe compatibilityof purifiedQE with chemicalscommonly used in shampoo formulationssuchas sodium lauryl sulfate, cocoamidopropylbe-taine, methylparaben, ropylparaben nd gluca-mate (Ceppi 1998). Other applications ncludethe solubilizationof vitamin E (Shirikawaet al.1994) and essential oils (Murakami1986; Mu-ramatsuand Suzuki 1993), hair tonic to reducehair loss (D'Aquino Silva 1994) and the pro-duction of hand creams(Kinekawaet al. 1992).

    PhotographyQE have been used for decades in the pro-duction of films as wetting agents to obtain auniformdistributionof silver halide microcrys-tals dispersed n gelatin(Ito et al. 1993; Katoetal. 1994).For this applicationmajorphotograph-

    ic companies (e.g., Konica, Agfa) use purifiedQE, with special emphasis in low-tannin, lowcolor properties.Wastewater Treatment

    QE areused in waste treatment n Japan.It isclaimedthat the additionof QE in the treatmentof waste waterenhancesoxygen transferandthedegradationof fats anddecreasesthe amountofsludge and the load of sludge dewatering(Tas-hiro 1991, 1992, 1995, 1996). Specific applica-tions include waste water treatmentof marineproducts and edible meat industries(Nagasaka1995a,b;NagasakaandTashiro1995).

    Miscellaneous Non-Food ApplicationsSome special non-food applicationsof quil-laja saponins are in the preparationof light-weight composites for building materials(Ton-yan and Gibson 1993), high-solidsaqueousdis-

    persionsof hydrophobizing gents(Craig1995),reductionof sulfuric acid mist in zinc electro-winning (Mackinnon1990, 1994), and aerobicex situ bioremediationof a chronicallypolych-lorobiphenyl PCB)-contaminatedoil (FavaandDi Gioia 1998).

    PHARMACEUTICAL USESAdjuvants for Animal and HumanVaccines

    The use of saponins as adjuvants n animalvaccines was first tested againstfoot andmouthdisease (Espinetet al. 1951). It was then shownthat only quillaja saponins are effective adju-vants, and a fraction that minimizedunwantedside effects was purified Dalsgaard1974). Thisproduct,known as Quil-A, is producedby theDanish firmSuperfos and is used commerciallyin many animal vaccines. Later it was shownthatQuil-A is a complex mixture of four majorsaponinswith varying degrees of toxicity, thatcan be separatedusing RP-HPLC(Kensil et al.1991). Some of the individualsaponins,notablysaponin QS-21, showed such low toxicity thatpotentially it could be used as adjuvant n thepreparationof human vaccines (Kensil et al.1991; Kensil 1996; Kensil, Wu, and Soltysik1995). Experimentalrialsinclude the testingofan HIV-1vaccine in non-humanprimates New-man et al. 1992), melanoma Hellinget al. 1995)and Plasmodium alciparum malaria(Stoute etal. 1997). QS-21 also augments both antibodyandcell-mediated mmuneresponses,suggestingthatthis adjuvant ouldbe a valuablecomponentin subunitvaccines (Wu et al. 1992). Anotherimportant pplication s in thepreparation f im-munostimulating omplexes, ISCOMS (Moreinet al. 1990). ISCOMS are used to presentviralmembraneproteins n a multimeric ormand areproduced by complexing cholesterol, saponinand another ipid (Moreinet al. 1984). The po-tentialof ISCOMS has been exploredin numer-ous articles and patents and some commercialvaccines are alreadyproducedby this method.

    Miscellaneous PharmaceuticalApplicationsPurifiedquillajasaponinsareused to lyse se-lectively red cells from whole blood prepara-tions, allowing for leukocyte differentialanaly-sis (Ledis et al. 1985), and the preparationofnucleic acid from white blood cells (RyderandKacian 1993). Modified quillajasaponins such

    as saponin DS-1 derived from saponin QS-21have been used to enhancenasal and ocularde-livery of insulin (Kensil, Soltysik, andMarciani1993; Pillion, Amsden,andKensil 1996) andasa permeationenhancerfor mucosal delivery ofthe aminoglycosides,an importantclass of an-

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    308 ECONOMICBOTANY [VOL. 53tibacterialagentsused for the treatmentof seri-ous gram-negativenfectionssuch as pneumoniaand peritonitis (Recchia et al. 1995). Also anenteral formulation containingpurifiedquillajasaponins and oils rich in omega 3 polyunsatu-ratedfatty acids, has been used for treatmentofinfection and inflammation Forse and Chavali1995).

    CONCLUSIONSQuillajasaponinshave a wide rangeof indus-trialapplications hatrequirean adequate upplyof raw materials.The use of bark as the solesource of quillajasaponinsis no longer viable,because over-exploitationof natural orests has

    causedimportant cological damage.The devel-opmentof a new process based on whole quil-laja wood, as well as plantations,can solve thisproblem,allowing the use of quillaja saponinsin novel and interesting ndustrialapplications.ACKNOWLEDGMENTS

    This work was supported by the Chilean Research and Scientific Coun-cil (FONDEF, Project #1 2010). DISCLOSURE: Dr. Ricardo San Martinis Professor at the Faculty of Engineering of the Catholic University ofChile, and is President of the Natural Response S.A., a private companypartly owned by the Catholic University.

    LITERATURE CITEDBalansard,J., F. Pellisier,and S. Conil. 1945.Actionde saponitesde Quillayaet de Sapindus ur le pou-voir absorbant, a germinationet la croissance deZea mais L. Proceedings Societe de Biologie deMarseille,MeetingNovember28. 140-142.Bureau, D. P., A. Harris, and C. Young Cho. 1998.The effects of purifiedalcohol extractsfrom soyaproduct on feed intake and growth of chinooksalmon (Oncorhynchusshawytscha)and rainbowtrout(Oncorhyncusmykiss). Aquaculture161:27-

    43.Ceppi, P. 1998. Uso de las saponinasen cosmeticos.B.Sc. thesis, Facultyof Engineering,CatholicUni-versityof Chile, Santiago,Chile.Craig, D. 1995. High solids aqueousdispersionsofhydrophobizing gents. U.S. patent5,403,392.Chavali, S. R., and J. B. Campbell. 1987a. Immu-nomodulatory ffects of orally-administeredapo-nins and nonspecific resistanceagainst rabies in-fection.International rchivesAllergy ApplicationImmunology84:129-34., and . 1987b.Adjuvant ffects of oral-ly administered aponinson humoraland cellularimmuneresponses in mice. Immunobiology174:347-59., L. D. Barton, and J. B. Campbell. 1988.Immunopotentiationy orally-administereduillajasaponins:effects in mice vaccinated ntraperitone-

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    Nagasaka, M. 1995a.Effect of dosing quillaiasaponinon wastewater rom marineproducts ndustry. ba-rakiDaigakuKogakubuKenykyuShuho43:91-96.. 1995b. Effect of dosing quillaiasaponinonwastewater romediblemeatindustry. barakiDai-gakuKogakubuKenykyuShuho43:97-103., and E. Tashiro. 1995. Oily wastewater reat-mentdosingquillaja aponin.Wastewaterreatmentplant n a marineproductsndustry.Shigen KankyoTaisaku31:1221-1228.Nayyar, D., W. Schulok, F. Saleeb, and E. Modesta.1998.Dry mix forproducinga slushbeverage.U.S.patent5,853,785.Newman, M., J-Y. Wu, R. Coughlin, C. Murphy, J.Seals, M. Wyand, and C. Kensil. 1992. Immu-nogenicityand toxicity testingof an experimentalHIV-1vaccine in non-humanprimates.AIDS Re-search HumanRetroviruses :1413-1418.Ninomiya, M., M. Matsumoto, K. Mabe, T. Okubo,B. Kin, and R. Kusuda. 1996. Compositionforfeed. JP 813 1087 A., H. Hatta, M. Fujiki, M. Kim, T. Yama-moto, and R. Kusuda. 1995a. Enhancementofchemotacticactivityof yellowtail(Seriolaquinque-radiata) eucocytes by oraladmistration f quillajasaponin.Fish and Shell Immunology5:325-328., M. Matsumoto, K. Busaku, and K. Riichi.1995b. Oral immunopotentiatoror aquatic ani-mals. JP 7107922 A.Ogawa, S., and T. Yokota. 1985. Surfactants.JP60,190,224., and F. Murakami. 1987. Saponinpurifica-tion. JP 62, 10,628.Oono, T., and Y. Higashimura. 1995. Method formanufacturing f redcoloringmaterial. P9590188A2.Organic Crop Improvement Association. 1994. In-ternationalCertification tandards.Phillips, J. C., K. R. Butterworth, I. F. Gaunt, J. G.Evans, and P. Grasso. 1979. FoodCosmeticsTox-icology 17:23-27.Pillion, D., J. Amsden, and C. Kensil. 1996. Struc-ture-function elationshipamongquillajasaponinsservingas excipients for nasal andoculardeliveryof insulin. Journalof Pharmaceutical cience 85:518-524.Poisson, J. 1972. Purified Panama wood extract.Frenchpatent2,126,523.1974. Tannin-free, urifiedextractof quillajarind and its uses in shampoos.German Offenle-gungsschrift2,421,129.Recchia, J., M. H. Lurantos, J. A. Amsden, J. Sto-rey, and C. R. Kensil. 1995. A semisyntheticQuil-

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    BOOK REVIEWWild 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 anytechnical nature successfully written for a general au-dience without formal training in anthropology and/orbotany. Rarer yet is to find a book written with thoseintentions that is also credible and useful to practicinganthropologists and botanists, especially those inter-ested in the other's discipline. Dunmire and Tierneyhave compiled such a book about plants in the FourCorners Region of the American Southwest by draw-ing on an impressive review of primary and technicalliterature, interviews, personal observations, and anexcellent collection of illustrations and color photo-graphs.Many books on economic plants are structured asinventories of plants and their uses that are either in-dependent of cultural affiliation or independent of eco-logical assemblages. The first chapter introduces thesubject by discussing the concept of place and usesthis as the organizing principle in the book. The phys-iography and the kind of vegetative communities thatcan currently be supported on the land in this regionare described. The following chapters introduce thecultures inhabiting this area through time (Ancestral

    Puebloans, Hopi, Navajo, Utes, and Jicarilla Apache),and how they used and managed plants. A brief historyof the people is given, their culture is described, andthe kinds of settlements they occupied are discussedin the context of plant procurement for human use.Pictures of artifacts utilizing plant materials are inter-

    spersedthroughout he chapters o help illustratehowplants areused. Foreach timeperiod,the authorspro-vide an idea of how environmental hange in the re-gion may have influenced he availableplantresourc-es, andhow this may differfromthe present"place"described n the firstchapterof the book.Thoughhundredsof plants arediscussed n generaltermsthroughout he text, a select groupof approxi-mately50 plantspeciescommon o thisplacearegiventwo to four page descriptions n the final chapterofthe book. After a brief introductiono ecozones-re-inforcingthe role environmentplays in plant distri-bution-the habitat is mentionedwhere the selectedplantsare most common,the cultureshistoricallyuti-lizing the plant are described as are the useful com-ponents of the plantand the modes of preparation fthosecomponents.The plants n this chapterarebeau-tifully illustratedand well photographed, ut I foundthe identification escriptorsparseandpotentiallydif-ficult to use. One needs to alreadyknow the nameofthe plant or flip throughnearly 150 pages to find it,which may not be appropriateor a generalaudience.Perhaps his sectionof thebookwouldbe easierto useif therewere a very simplistickey basedon leaf typeor flowershape, for example.One of the highlightsof thisbook is a chartof over300 plant species known to have been used by thepeopleoccupyingtheregion.Theuses, culturalgroupsknownto use themandreferencesareall in one place.This book will be a useful tool for ethnobotanists ndthose with a generalinterestof the economicuses ofplantsalike.

    ROBIN C. D. CURREYSCHOOLOF NATURALRESOURCESAND ENVIRONMENTUNIVERSITYOF MICHIGAN