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Berry Productivity and Molluscicidal Saponin Yield of Phytolacca dodecandra (Phytolaccaceae) under Different Sunlight, Watering and Nutrient

Conditions Author(s): Jerikias Ndamba, Ian Robertson, Else Lemmich, Stephen K. Chandiwana, Peter Furu and Per Mlgaard Reviewed work(s): Source: Economic Botany, Vol. 50, No. 2 (Apr. - Jun., 1996), pp. 151-166 Published by: Springer on behalf of New York Botanical Garden Press Stable URL: http://www.jstor.org/stable/4255827 . Accessed: 03/05/2012 11:45
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BERRY PRODUCTIVITY AND MOLLUSCICIDAL SAPONIN YIELD OF PHYTOLACCA DODECANDRA (PHYTOLACCACEAE) UNDER DIFFERENT SUNLIGHT, WATERING AND NUTRIENT CONDITIONS'
JERIKIASNDAMBA, IAN ROBERTSON, ELSE LEMMICH, STEPHEN K. CHANDIWANA, PETER FURU, AND PER M0LGAARD

Ndamba, Jerikias. (Blair ResearchLaboratory,P.O. Box 8105, Causeway,Harare, Zimbabwe), Ian Robertson (Department of CropScience, University of Zimbabwe, P.O. Box MP167, MountPleasant,Harare,Zimbabwe), Else Lemmich (RoyalDanishSchool of Pharmacy,Universitetsparken 2, DK-2100 Copenhagen, Denmark),Stephen K. Chandiwana (BlairResearch P.O. Box 8105, Causeway,Harare, Zimbabwe), Laboratory, Peter Furu (Danish Bilharziasis Laboratory, JaegersborgAlle ID, DK-2920 Charlottenlund, Denmark),and Per M0lgaard* (Royal Danish School of Pharmacy,Universitetsparken 2, DK-2100 Copenhagen,Denmark).
BERRY PRODUCTIVITY ANDMOLLUSCICIDAL SAPONIN YIELDOF PHYTOLACCA DODECANDRA UNDER DIFFERENT ANDNUTRIENT SUNLIGHT, WATERING CONDITIONS. EconomicBotany50(2):151-166,

1996. The cultivationof Phytolaccadodecandrahas been studied in Zimbabwein order to


initiate local production of the berries from which molluscicidal saponins can be extracted and used in schistosomiasis control programs. The effect of shading, water and nutrient type on growth and berry yield of an imported Ethiopian and a Zimbabwean cultivar were monitored. The molluscicidal potency and saponin concentration of the aqueous extracts of berries, harvested at the full grown, unripe development stage, were determined. There was a marked seasonal variation in berry production with both cultivars being highly productive in the dry season. The best growth and the highest berry yield was obtained with plants grown in full sunlight, under irrigation and with application of cattle manure. The Ethiopian cultivar showed in general better growth and higher berry yield than the Zimbabwean cultivar. Cultivation in shade gave a substantially lower berry yield and saponin concentration in both cultivars, compared to full sunlight. Irrigation was beneficial to the berry yield in both cultivars but lowered the saponin concentration in berries of the Ethiopian cultivar. However, it did not counteract the advantage of irrigation on the total yield. Addition of manure did significantly increase the growth and the berry yield of both cultivars compared to fertilizer application and to the control. The saponin concentration was in general lower with addition of manure, with exception of irrigated Ethiopian plants. The combination of cattle manure and irrigation resulted in the highest saponin yield. Although the two cultivars had different saponin patterns, these patterns and the relative proportions of the saponins were constant, irrespective of the treatments. Production de baies et rendement de saponines de la molluscicide vegdtale Phytolacca dodecandra (Phytolaccaceae) influences de soleil, d'irrigation et d'engraissage. Un extrait aqueux des baies de Phytolacca dodecandra contient des saponines toxiques a 1'egard des mollusques, hotes intermediaires de la maladie tropique parasitaire schistosomiase. Cette plante a et selectionnee comme la meilleur pour produire des materiaux molluscicides dans l'utilisation de programmes sanitaires dans le but de controller la schistosomiase. La culture de deux cultivars de Phytolacca dodecandra a ete etudiee au Zimbabwe pour evaluer les meilleures circonstances de culture. Le plus grand rendement de baies et de saponines a ete obtenu d'une culture ensoleillee, d'irrigation et d'engraissage avec des engrais d'animaux domestiques.

Key Words: Phytolacca dodecandra; molluscicide, plant; schistosomiasis; monodesmosidic


saponins; berry yield; cultivation methods.

* Correspondence.

'Received 10 August 1995; accepted 3 October 1995. Economic Botany 50(2) pp. 151-166. 1996 ? 1996 by The New York Botanical Garden, Bronx, NY 10458 U.S.A.

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The use of snail killing compoundsof plant origin for the control of the tropical infection, schistosomiasis,is attractive because of the economic advantageof cultivatingthe plantslocally instead of importing synthetic molluscicides. The most widely studied plant for this purpose
is Phytolacca dodecandra L'Herit, because it is

fast growing and produceshighly molluscicidal saponinswhich can be extractedfromthe berries with water.P. dodecandrais consideredthe best candidatefor productionof a plant molluscicide to be used in integrated control programmes (WHO 1983); however,the use will requirecultivation in areasbeyond its naturalrange. Since the discovery of the molluscicidalpropertiesof the plant (Lemma 1965) only few and uncoordinated attempts have been made to relate growth and berry yield to the cultivationmethod, and molluscicidalpotency and saponincontent of the berrieshave not previouslybeen systematically studied related to the production method. In order to facilitate organizationof community based control programmes,including local productionof the molluscicide, it has been suggested that detailed agronomicdata be made available (Webbe 1987). Because of its low economic value, it is unlikely that P. dodecandrawill be considereda high prioritycrop by the farmersandplantedin the best production areas. Thus studies to assess the demandof the plant for light, water and nutrientsand their influence on growthperformance and yield are essential in orderto providecost effective production guidelines. It is well known that the production of secondary metabolites is influenced by the cultivation method. Thus, saponin production in Saponariaspp. is dependenton light and also on the availabilityof water soluble nutrientswhich may be leached from the roots induced by rainfalland related to fertilizerapplication (Bernath 1986; Fluck 1954). The composition and quantityof cardioactiveglycosides
of Digitalis purpurea L. from various parts of

Europewas relatedto the methodof cultivation (Lugt 1976). In natureplants are found where a variety of conditions for growth and survival including their nutrient,light and water requirements are met. To grow P. dodecandrain a monoculture similar or better conditions must be provided. The variationin rainfallpatterns,soil types and fertility necessitates supplementarywater and nutrientswith potentialinfluence on berry sap-

onin contentand yield. Preliminary observations on the influence of sunlight and wateringon P. dodecandra berry yields of selected Ethiopian types (Lugt 1981) and the association between high content of organic matter in the soil and berry yield of P. dodecandra plants seen in Zambia (Chimbelu and Shehata 1987) clearly demonstratesthe need for more controlled nutritionalstudies. Biannualberry productivitywas seen among cultivatedplants in Ethiopiaand was attributed to the two annualrainy seasons in that country (Lugt 1981). However,this might be a genotypical phenomenonand the plants might produce berriestwice a year in areaswith only one rainy season. When conditions are conducive, Phytolacca plants growing naturally in Zimbabwe produce berries from July until October only, that is, from the winter until the beginning of the rainy season (Ndamba and Chandiwana 1988). The molluscicidalconstituentsof P. dodecandra aqueous berry extracts have been determined to be monodesmosidicsaponinsthatcontain the triterpene oleanolic acid as the aglycone and usually trisaccharides as the sugarmoiety of the compounds(Fig. 1) (Parkhurst et al. 1973a, b, 1974;Thiilborget al. 1993, 1994). These arise when the naturallyoccurringbidesmosidic saponins are enzymaticallyhydrolysedduringextractionwith water(Parkhurst et al. 1989). Only limited phytochemicaldatais availableon intrastrainvariationof saponincontentin relationto the agricultural productionmethods used when an Ethiopiancultivaris raisedunderforeignecological conditions.The view that P. dodecandra is availableonly for use in specific countriesbecause of scarcity and anticipateddifficulties of raising potent varieties in other schistosomiasis endemic localities (Marston and Hostettmann 1987) is probablydue to this paucityof scientific information. Our study was undertakento determinethe influence of cultivationmethods on the growth of P. dodecandraand on the content of monodesmosidic saponins and molluscicidalpotency of the aqueousextractsof its berries.The results presented here were obtained from an experimental plot established in Zimbabwe to assess productivitypatternand yields of an imported Ethiopianand a Zimbabweanvariety of P. dodecandra cultivated using different watering, fertilizerand light regimes. The saponincontent

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DODECANDRAMOLLUSCICIDAL SAPONIN NDAMBA ET AL.: PHYTOLACCA

153

HR1 C

OH5 H3

CH

ETEIIOPIAN CULTIVAR E44 1: R1=cz-L-Rhap-(1-42)-j3-D-G1up 2: R1=R3=13-D-Glup 3: R1=R4=H 4: R'=R3=R4=H 5: R1=R3=R4=H ZIMBABWEAN CULTVAR MS1 1: R1=H 2: RW==-D-Glup 3: RW=j-D-Glup 4= E44/2 5: RW=R3=H 6: R1=R3=H 7= E44/3 of the majormonodesmosidic Fig. 1. The chemicalstructures saponinsthathave been isolatedfromaqueous MS1 cultivarsof P. dodecandra(cf. Thiilborget al. berry extractsof the EthiopianE44 and the Zimbabwean 1993, 1994). R2=cx-L-Rhap-(1-*2)-1-D-Gap R2=P-D-Gapp R4H R4OH R5=CH3 R5=CH3
R2=3D-G

R2=R4=H R2=R4=H

R3=j3-D-Glup

R5=CH3 R5=CH3

R2=jD-GaOp

R3=S-D-Glup

R5=CH3 R5=CH3 R5=CH3

R2=ax-L-Rhap-(1--2)-fD-D-Ga1 R2=j-D-Xylp-(1-6)-R-D-Glup

R3=1-D-Glup R3=j-D-Glup R3=P-D-Glup

R4_ OH R5=CH23H R5=CH23H R4-OH R5=CH3

R2=R4=H R2=H G

and molluscicidal potency of the berries were Composite soil samples were taken from ten determinedand, for snail control,it is estimated randomlyselected spots at the plot afterploughthat at concentrations of 50 mg/I recommended ing and before planting. for field use (Ndamba,Chandiwana and Makaza PLANTING 1989), berriesfrom one plant would be enough to treata water volume of 30 m3. P. dodecandraplants,cultivar Sixty Ethiopian E44, were raised from soil-free rooted cuttings MATERIALSAND METHODS importedthroughplant quarantinefrom EthioSTUDY LOCALITY pia. The same number of plants were raised The plants were cultivated at the Kutsaga Re- from cuttings of the most potent Zimbabwean search Station situated 15 km east of Harare. cultivar,MS1 (Ndambaand Chandiwana1988). The station is located at 1484 m receiving 840 The plants were assigned a numberat planting mm of rainfallper year and with an annualmean before being divided into 12 groups of five of 21?C.A 1 ha plot was established plants each. Spacing of the holes was 2 m betemperature on virginland, clearedand deep ploughed(1 m). tween rows and 1 m within rows, shown from

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previous studies to be ideal for P. dodecandra unripe berries were harvested, counted, and and Kloos 1989). numberrecorded.Immediatelyafter harvesting, cultivation (Wolde-Yohannes racemes from each plant were tied together,laThe study design was as follows: belled, air driedin the shade for two weeks, and Watering dry weight determined.Dried berriesfrom each All plants in the plot were planted on the plant were divided into two equal partsto facilsame day and wateredregularlyfor four weeks. itate determinationof the saponin content and Thereafter,watering varied depending on the the molluscicidalactivity. treatmentto which the plants were assigned. DETERMINATION OF MOLLUSCICIDAL Half the plants were flood-wateredat least two POTENCY (LC50mg/1) to three times a week, while the remaininghalf The molluscicidalactivity of the driedberries were not watered in order to facilitate assessment of the effects of wateringcombined with was assessed againstlaboratory bred and reared other types of treatment. Naturalrainfallpattern Bulinus globosus (Morelet 1866) snails accordwas monitoredfrom records of a nearby mete- ing to the WHO provisional technique (WHO 1965), but with only five snails for each conorological station. centration.The berries were crushed (particle Plant Nutrients size 0.42 mm) and a stock extractof 100 mg/l Plants were given three different nutrient prepared.The suspensionwas stirredovernight treatments: One level of commercialfertilizers, before serial dilutions of the extract with deone level of cattle manureand as a control no chlorinated water were prepared. The snails addedplant nutrients.The fertilizerswere given were exposed to concentrations rangingfrom 0 as superphosphate(80 g) addedand coveredby to 20 mg driedberries/I for 24 hoursandto fresh 10 cm of soil before planting.Immediatelyafter water for another24 hoursbefore mortalitywas planting, 80 g of ammonium nitrate fertilizer assessed. Potencytests were repeatedthreetimes was addedinto a trench 10 cm deep dug around for the same berry collection and the LC50 valthe top of the hole 60 cm in diameter.Ammo- ues per plant were calculated and group mean nium nitrate was applied once at planting and potenciesdetermined accordingto the methodof again in a similarmannerat the beginningof the Litchfield and Wilcoxon (1949) using a comrainy season. Cattlemanurefrom a nearbycom- puter program, Pharmacologic Calculations mercialdairy farmwas mixed with soil at a ratio (PCS) Version4 (Tallarida and Murray1981). of 1:2 (manure:soil). The mixturewas placed in QUANTITATIVE DETERMINATION OF the hole priorto planting.No analysisof nutrient MONODESMOSIDIC SAPONINS IN content in the manurewas attempted.
AQUEOUS BERRY EXTRACTS

Shading A 40% shade cloth of nylon was erected to cover half of the plants to simulatethe effect of shade trees.
PLANT GROWTH MONITORING

Two primary branches on each plant were identifiedand labelled using metal disks to ensure measurementof the same branchesduring the study. Plant height (length of shoot) and number of branches were recorded at planting and then weekly.
DETERMINATION OF BERRY PRODUCTIVITY PATYFERN AND YIELDS

The number of racemes with unripe berries were counted and recordedweekly from the labelled branches. All racemes with full grown,

The berries were ground (particle size 0.42 mm) and a stock extractwas prepared by mixing 100 mg berry powder with 50 ml water and shaking the suspension overnight. The sample was preparedby passing the extract througha Chromafilfilter (25 mm, pore width 0.45 pRm), washing the filter with an equal amount of 2propanolcontaining 50 [LI 0.1 M NH3/ml, and additionalwater until the stock extract was diluted five times. This sample solution was analyzed by HPLC (Ndamba,Lemmich and M01gaard 1994a) on a Spherisorb S5 C8 (4.6 X 250 mm) column with an eluent consisting of acetonitrile-0.05 M ammonium dihydrogenphosphatebuffer,pH 4.4 (42:58). The saponinswere monitoredat 203 nm and at a flow rate of 1.2 ml/min.The concentrations of the saponinswere determinedby means of externalstandardsolu-

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155

Ethiopian plant

Zimbabwean plant

UV abs
203 nm

UV abs
203 nm

1+2

6+7

12 mins

.l

12 mins

of aqueousberryextractsfromEthiopian E44 andZimbabwean Fig. 2. HPLCChromatograms MS1 cultivars of P. dodecandracultivatedunderanalogousconditions.

tions of pure saponins. Four concentrationsof mixed standardswere preparedfrom the saponins E44/1, E44/2, E44/3 and E44/5 from the

SOILCHEMISTRY

The soil was pale brown with mediumgrained sandy texture.Soil pH was 4.7 with no EthiopiancultivarE44. The saponinE44/4 was trace of free CO3.Available P205 (by resin exomitted because it occurred only in small amountsand was not fully separatedfrom E44/ tract)was 8 ppm and initialmineralN (NH4plus 3 and E44/5. Mixed standardsof the saponins NO3)before incubationwas 9 ppm and 17 ppm MS1/2, MS1/3, MS1/4 and MS1/7 from the after incubation.ExchangeableK was 0.07, Ca Zimbabwean cultivar MS1 were prepared as 0.84 and Mg was 0.30 mg/100g of soil. Total well. MSI/1 was analyzed jointly with MS1/2 exchangeablecations were 1.2 mg/lOOg.All the because the two compounds co-chromatograph results were expressed in terms of the air-dried under the circumstancesused. At a lower ace- soil samples passed througha 2-mm sieve. tonitril concentrationthey occur as separated peaks, but then retention times are very long. PLANT SURVIVAL AND However,it was shown that the two compounds GROWTH PERFORMANCE have identicalresponse factors and occur in almost equal amounts(Fig. 2). Five EthiopianE44 plantsdied duringthe first three month after planting but all the ZimbabRESULTS wean plants survived. Shaded Ethiopianplants CLIMATE had more shoot elongationcomparedto full sunThe experimentalplots received 910 mm of light (P < 0.001 by t-test). Cattle manurererainfall during the study. Temperatureswere sulted in the highest plant growth in full light, moderate with a mean annual temperatureof while fertilized plants were the fastest growing 21?C. Night temperatures were low with tem- under shading.In general,plant growthwas inbelow freezing in June throughto Au- fluencedby shadingand nutrient peratures type, while wagust. tering had no effect.

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Ethiopian cultivar
800 700 600 500 400 300 200 100 Harvested racemes Fertilizer Manure No nutrients Harvested racemes 800 700 C

600 500 400-

Fertilizer Manure No nutrients

Light

300 200 100

Shade

16 20

24 28 32 36 40 44 Time after planting (wks)

48

16 20

24 28 32 36 40 44 Time after planting (wks)

48

Zimbabwean cultivar
800Harvested racemes Fertilizer Manure No nutrients 800700600500400 Harvested racemes Fertilizer Manure No nutrients 700 600 500400-

300200100l

Light
;200

300-

Shade

~~~~~~~~~10024 32 36 28 20 Time after planting (wks) 16 32 36 28 24 20 Time after planting (wks)

16

Fig. 3. Number of racemes harvested from Ethiopian E44 and Zimbabwean MS 1 plants cultivated under similar watering, sunlight, and nutrient levels.

BERRY PRODUCTIVITY AND YIELD

Number of Racemes Harvested Dry Berry Yield Shade, water, and nutrienttype had a highly significantinfluenceon the numberof harvested There was a significant(P < 0.001) influence racemes. More racemes were harvested from of shade, water and nutrienttype on dry berry

plants cultivatedin full light, watered and provided with cattle manure(Fig. 3).

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157

Ethiopiancultivar
Berry yield (g/plant)

Zimbabweancultivar
1600 Berry yield (g/plant)

1600

Water [
1200| 1200 No water 8sticde BShade water

no water

830021

eL

800.

4001

Bt400

Fertilizr
candra.

Manure

Nothing

Fertilizer

Manure

Nothing

Fig. 4. Dry berry yield of groups of similarlycultivatedEthiopianand Zimbabweancultivarsof P. dode-

yield of Ethiopian and Zimbabwean cultivars (Fig. 4). No significanteffect of the shade/water and water/nutrients interactionon berry yield of the Ethiopian cultivar was apparent(Table 1). Cultivationwith cattle manuregave consistently high berry yields with both light and shaded plants. The average yield was 1443 g/plant for the full light, manured,and watered Ethiopian
TABLE 1. DRY BERRY YIELDa OF THE ETHIOPIAN E44

plants, while similarly cultivated Zimbabwean plants produced816 g/plant (Table2). Berry Yield and Productivity in Relation to Growth Performance Regression analysis indicated significant linear relationships between plant height (shoot length) and berry productivity among plants
VARIETY IN RELATION TO CULTIVATION METHODS. Nutrients

Treatment

Fertilizer

Manure

None

Mean

Water No Water Mean


Maineffects

No shade Shade No shade Shade

567 530 335 294 431


P-value

+ 87
? 49

? 13 ? 57 ? 136

1443 ? 808 ? 1141 ? 417 ? 981 ?

96 70 121 31 389

549 ? 328 ? 314 ? 189 ? 367 ?

27 32 57 21 133

853 557 638 317 600

? 438 ? 217

? 421 ? 99 ? 375

Analysisof variance Interactions P-value

Shade Water Nutrients

0.001 *** 0.001 *** 0.001 ***

Shade/water Shade/nutrients Water/nutrients

0.057 n.s. 0.001 *** 0.166 n.s.

Height(covariate)

0.004

**

Shade/water/nutrients

0.022

' Meanof five plants,g/plant,? s.d. Levelof significance: n.s. = not significant. *, **, and *** indicateP < 0.05, P < 0.01, andP < 0.001, respectively;

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TABLE 2. ODS.

MS 1 VARIETY IN RELATION TO CULTIVATION METHDRY BERRYYIELDa OF THE ZIMBABWEAN

Nutrients Treatment Fertilizer Manure None Mean

Water No Water Mean


Mean

No shade Shade No shade Shade

377 + 299 + 180 ? 205 ? 271 +


P-value

95 124 60 127 125

816 ? 426 ? 650 ? 256 + 537 +

63 149 65 76 235

230 ? 138 ? 167 ? 153 ? 175 ?

56 73 19 46 62

474 287 357 213 339

? 267 ? 167 ? 246 ? 93 ? 225

Analysis of variance Interactions P-value

Shade Water Nutrients Height (covariate)


a Mean

0.001 0.001 0.001 0.988

*** *** ***

n.s.

Shade/water Shade/nutrients Water/nutrients Shade/water/nutrients

0.443 0.011 0.125 0.314

n.s. * n.s. n.s.

of five plants, g/plant, ? s.d. Level of significance: *, **, and *** indicate P < 0.05, P < 0.01, and P < 0.001, respectively; n.s. = not significant.

raised in full sunlight (P < 0.01). Negative coefficients were recordedin the shade, but these were not significant.This was interpretedas a reducedproductivityamong high growthplants. Seasonal Variation in Berry Productivity and Yield There was marked seasonal variationin the berry productivityand yield of both cultivars. Berry productionin Ethiopianplants cultivated in full light began four months after planting reaching a peak in October (Fig. 5). The Zimbabwean variety was also productive four months after planting, but with berry yield reaching a peak in September(Fig. 5). In both cases, shade lowered yields and delayed initiation of fruit setting by at least a month.
SAPONINCONTENT

potency of the Ethiopianvariety was water/nutrient (Table 3). For the Zimbabweancultivar, similaranalysis showed markedinfluenceof all parameters,shade, water and nutrienttype on saponin content (Table 4) and on the molluscicidal potency of the berries(Fig. 6). Saponin Contribution to Berry Molluscicidal Potency Regression analysis showed a strongcorrelation between berry potency and extractedmonodesmosidic saponinconcentration (Fig. 7). Although the proportionsof the individual saponins in berry extracts harvested from watered and droughttreatedplants were consistent, the quantitywas nutrientdependent(Fig. 8). There was a markedgenotypicaldifferencein the number and identity of monodesmosidic saponins extractedfromthe two cultivarsas seen fromthe peaks in the HPLCchromatograms of the aqueous berry extracts(Fig. 2). Dry Berry and Saponin Yield Ethiopianplants fertilizedwith cattle manure and irrigatedhad the highest saponinyield, 291 g/plant(Table5). Plantsthat were fertilized,but not irrigated,had the lowest saponin yield (74 g/plant)due to a low berry yield and in spite of a high saponin concentration.Plants cultivated underdroughtconditionsand providedwith cattle manure were the second most productive group (199 g/plant). The Zimbabweanvariety reactedsimilarlywith the highest saponinyields

Saponin Concentration and Molluscicidal Potency Berries harvestedfrom the Ethiopiancultivar had higher saponin content and molluscicidal potency compared to those from the Zimbabwean variety(Tables3 and 4, respectively).The highest berry saponin concentrationfrom the Ethiopiancultivar was 222 mg/g, estimated to be 22% of berry weight. Analysis of variance showed that only water level had a significant effect on saponincontentof berry extractsfrom the Ethiopiancultivarwith significantinteraction with nutrient type (Table3). The only interaction that significantlyinfluencedsaponincontentand

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159

Ethiopian cultivar
Berry yield (kg/month)
-E

Zimbabwean cultivar
Berry yield (kg/month)
--3

Rainfall (mm) 300

Rainfall (mm) 300

Fertilizer
Manure / - 20

Fertilizer
Manure
0

3-

3D

3 - 200
...Total

Nothing Total

No nutrients

- 200

2E]Rainfall

15 100

IlRainfall15
-100

1 0 M A M J J A S 0 N D J F Month of the year

?2SFs 1

~~~~~~~~~~~50
MA M J J A S O N D J F Month of the year

50

Fig. 5. Seasonal variationin berry yield (kg/month)of EthiopianE44 and ZimbabweanMS1 cultivarsof P. dodecandra.

of 127 g/plantobtainedunderirrigationand cat- A high saponin yield was primarily based on a tle manurefertilization(Table5). Althoughsap- high berry yield. was relatedto the cultivation onin concentration DISCUSSION methods (Tables 3, 4), the total saponin yield PLANT GROWTH AND PERFORMANCE was to a much higher degree related to berry yield, which was more affected by cultivation One of the major criteria for a potential plant (Fig. 9). molluscicide is its suitability for cropping (Boymethodsthanthe saponinconcentration

TABLE 3.

THE EFFECT OF CULTIVATION METHODS ON THE SAPON1INCONCENTRATIONaOF P. DODECANDRA

AQUEOUS EXTRACTS OF BERRIES HARVESTED FROM THE ETHIOPIAN VARIETY E44. Nutrients Treatment Fertilizer Manure None Mean

Water No Water Mean


Main effects

No shade Shade No shade Shade

168 + 196 ? 222 ? 220 203


P-value

20 17 23 31 28

202 + 187 + 174 + 177 181

7 23 13 23 20

187 ? 188 ? 212 ? 193 ? 189 +

19 42 14 9 27

185 ? 190 ? 201 ? 197 ? 193 ?


P-value

21 28 27 30 27

Analysis of variance Interactions

Shade Water Nutrients Berrywt (covariate)


a

0.364 n.s. 0.038 * 0.064 n.s. 0.122 n.s.

Shade/water Shade/nutrients Water/nutrients Shade/water/nutrients


=

0.391 n.s. 0.407 n.s. 0.004 ** 0.319 n.s.


P < 0.05; ** = P < 0.01.

Mean of samples from five plants, mg/g dry berries, ? s.d. Level of significance: n.s. = not significant;

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TABLE

4.

THE EFFECT OF CULTIVATION METHODS ON THE SAPONIN CONCENTRATIONa OF P. DODECANDRA

AQUEOUS EXTRACTS OF BERRIES HARVESTED FROM THE ZIMBABWEAN VARIETY MS 1.


Nutrients Treatment Fertilizer Manure None Mean

Water No Water Mean


Main effects

No shade Shade No shade Shade

188 170 207 150

? ? ? ?

30 10 24 12

181 ? 27
P-value

156 ? 28 135 8 161 ? 12 5 126 21 146


Analysis of variance Interactions

? ? ? ? 180 ? 192 166 190 166

25 5 18 4 20

178 ? 30 158 ? 18 186 ? 24 146 19 168


P-value

28

Shade Water Nutrients Berry wt (covariate)

0.001 0.045 0.010 0.020

*** * ** *

Shade/water Shade/nutrients Water/nutrients Shade/water/nutrients

0.805 0.658 0.079 0.549

n.s. n.s. n.s. n.s.


=

' Mean of samples from five plants, mg/g dry berries, + s.d. Level of significance: n.s. = not significant; * = P < 0.05; ** = P < 0.01; * < 0.001.

er 1982). The importanceof evaluating simple for domestication culturalcroppingrequirements of a plant molluscicide has been highlighted (Lugt 1987; Webbe 1987). In this project we have investigated factors influencing dry berry yield, saponin content and molluscicidalpotency, and the correlationbetween these parameters and the general plant growth.

The high allocation of assimilates to stem growth may coincide with the period of maximum growth of the fruits, resulting in an increased competitionfor nutrientsbetween vegetative and reproductive components,therebydeand growth(Donald creasingracemeproductivity and Hamblin 1976). This was seen among the shadecultivatedplants.High growthbut low ber-

Ethiopiancultivar
20-LCSO(mg/L) Water

Zimbabweancultivar
20
LC50 (mg/L)

I
1 -2

No water Shade, water


Shade, no w.

15

10- -

10

5--

5-

0Fertilizer Manure No nutrients

~~~~~~~~~0
Fertilizer Manure No nutrients

of aqueousextracts levels on molluscicidalpotency(LC50) Fig. 6. The influenceof shade,water,and nutrient cultivarsgrown in a commonenvironment. of berriesharvestedfrom the EthiopianE44 and Zimbabwean MSI1

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SAPONIN DODECANDRAMOLLUSCICIDAL NDAMBA ET AL.: PHYTOLACCA

161

in raceme formationand peak berry production the height of the dry season (Fig. 5). in October, At the beginningandduringthe rainyseason,priority is given to vegetative growth,and the nutrientresourcesare allocatedaccordingly (Ndam15 ba 1993). Such seasonal changes appearto be inducedbecausethe two producenvironmentally tive periods previously reportedfor Ethiopian r -0.93 in Zimbabweduring cultivarswere not apparent 10 r2- 0.87 the present year long investigation (Lugt 1981). p ( 0.001 The net effect of irrigation was to increaseplant growth,but not berryproduction. Transmission of schistosomiasisin Zimbabwe 5has been shown to be markedlyseasonal,reaching a peak in the monthof Octoberand coinciding with the maximumin berry yield (Fig. 5). in July is most preHowever,winterapplication 250 100 150 200 ferred for maximumreductionin snail populaSaponin content (mg/g) tions (Chandiwana,Christensen and Frandsen Fig. 7. The relationshipbetween berry saponin 1987; Shiff and Clarke 1967). In spite of P. doin winter,it is decandraberryproduction starting contentand molluscicidalpotency. wouldbe harvested unlikelythatenoughmaterial duringthe same season to meet the demandsfor It wouldbe essentialto devote ry yield of P. dodecandra plants has been re- a controlprogram. corded in Ethiopia (Wolde-Yohannes 1983). In the first year of a controlprogramonly to proaddition, the negative regression coefficients of duction of berries, which could be dried and plant height and productivity clearly demonstrates storedfor use duringthe following transmission the disadvantages of shading, at least in the way season. increased it was used in this study. The artificial shade was Artificial fertilizers plantgrowthboth created by use of a shade cloth supported by long in shadeand sunlight(Ndamba1993),but did not (Figs. 3, 4). Increased poles. Although this would not be recommended increaseberry production for practical use, it was intended to mimic the biomass yield coupled with low harvestindices effect of shade trees. Considering the costs and after applicationof nitrogenfertilizershas been complexity of erecting shading, such agricultural documentedwith other medicinal plants (Lugt methods are difficult to justify unless there are 1987). This has been attributed to a high evapoother added benefits. Although shading might be transpiration rateenhancing vegetativegrowthreuseful in areas with high daytime temperatures to sultingin a higherleaf areaindex, which leads to conserve the moisture, this would be deemed im- a net decreasein availablesoil water.However, practical under local farming practices. artificialfertilizersare expensive and would not Berry productivity and mature raceme forma- be ideal for a village based cultivationprogram tion are directly related to plant growth perfor- for plantmolluscicides.The use of cattlemanure mance (Lugt 1981, 1987). Berry yield is a func- increased with unfertilized berryyields compared tion of the number and size of racemes, size and plots in Ethiopia (Wolde-Yohannes and Kloos quantity of berries on the raceme, moisture con- 1989). Most farmersin rural areas alreadyuse tent and vegetative growth, not necessarily an in- cattle manureto fertilize their fields. If they redicator of the productive potential of the plant alize that they could cultivateP. dodecandraas (McDermit and Loomis 1981). The likelihood a crop without additional inputs, their profits that a high number of racemes with small berries might equal what they get for a food crop.
20 of low dry matter content can be harvested from high performing plants is well recognized (Donald and Hamblin 1976). During the dry season, it appears that most of the available plant nutrients are used for reproductive purposes, resulting SAPONINYIELD

LC50 (mg/L)

The finding that plant saponin content responds to new environmentalconditions imposed by the domesticationprocess highlights

162

BOTANY ECONOMIC

[VOL. 50

Ethiopian cultivar
140 Saponin content (mg/g) 120 100= ~~Fertilizer Manure No nutrients 140 Saponin content (mg/g)

Water

12 100N-

IIFertilizer LII No nutrients

Manure

40

0;6

|S4

E44/1 80 -8060 -60

E44/2

E44/3

E44/5

E44/1

E44/2

E44/3

E44/5

Zimbalbweancultivar
140 Saponin content (mg/g)

40 -4 140 Saponin content (mg/g) 20 -2

Water Fertilizer
i Manure

120-~~I
0 100

120- 0 t 100

Fertilizer
Manure No nutrients

o water

ae

LI] No nutrients

[III

MS1/2

MS1/3

MS1/4

MS1/7

MS1/2

MS1/3

MS134

MS1/7

Fig. 8. Saponincontentof aqueousextractsof berriesharvestedfrom individualsof the EthiopianE44 and the Zimbabwean MS1 cultivarsin relationto cultivationmethods (mean of five plants)

the need to studycultivationmethodsto increase yields of potent berries (Small, Jurzysta, and Nozzolillo 1990). In field studies, the complex biological processesinvolved in secondaryplant

metabolism make it difficult to attributesingle factor influences. But, based on our cultivation of two clonal varieties of P. dodecandraunder analogous soil and climatic conditions, intra-

1996]
TABLE 5.

SAPONIN DODECANDRAMOLLUSCICIDAL NDAMBA ET AL.: PHYTOLACCA

163

BERRY YIELD, SAPONIN CONCENTRATION AND YIELDa OF PLANTS CULTIVATED IN FULL SUNLIGHT

WITH DIFFERENT LEVELS OF NUTRIENTS AND IRRIGATION.


Nutrients Treatment Productivity Fertilizer Manure None

Ethiopianplants Water

No water

Berry yield (g/plant) Saponincontent(mg/g) Saponinyield (g/plant) Berry yield (g/plant) Saponincontent(mg/g) Saponinyield (g/plant) Berry yield (g/plant) Saponincontent(mg/g) Saponinyield (g/plant) Berry yield (g/plant) Saponincontent(mg/g) Saponinyield (g/plant)

567 168 95 335 222 74 377 188 71 180 207 37

1443 202 291 1141 174 199 816 156 127 650 161 105

549 187 103 314 212 67 230 192 44 167 190 32

plants Zimbabwean Water

No water

aBerry

saponin yield was calculated from dry berry weight (g/plant) X berry saponin concentration (mg/g).

strainberry saponin content and potency variations could largely be ascribedto the cultivation methods used. The results indicate that cultivation methods,environmental and genotypicfactors, play critical roles in determiningthe sapo-

nin content of the berries of P. dodecandraas in other species (Aubert,Daunay, and Pochard 1989; Kawanishiet al. 1983; VetterandHaraszri 1987). The saponin content of 22% is in agreement

Ethiopian cultivar
300 Berry saponin yield (g/plant)

Zimbabwean cultivar
300 Berry saponin yield (g/plant)

250 200 -

E Water 'KW
INo w
2.

250 200 -

Water
No water

150 -

150 -

100 50 -

dZ

100

50-

0-

'N

v~*

0-

Fertilizer

Manure

Nothing

Fertilizer

Manure

Nothing

Fig. 9. Saponinyield calculatedfrom dry berry weight and saponincontentof the EthiopianE44 and the Zimbabwean MS1 cultivarscultivatedin full sunlight.

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BOTANY ECONOMIC

[VOL. 50

LITERATURE CITED Aubert, S., M. C. Daunay, and E. Pochard. 1989. Steroidicsaponinsfrom eggplantSolanummelongema L II. Effects of culturalconditionsgenotypes and parthenocarpy. AgronomieParis9:752-758. Bernath, J. 1986. Productionecology of secondary plantproducts.Pages 185-234 in L. E. Craker and J. E. Simon, eds., Herbs spices and medicinal plants.Recent advancesin botany,horticulture and pharmacology. Volume 1. Oryx Press. Boyer, J. S. 1982. Plant productivityand environment. Science 218:443-448. Chandiwana, S. K., N. 0. Christensen, and F. Frandsen. 1987. Seasonal patternsin the transmission of Schistosomahaematobium, S. matthei, and S. mansoni in the highveld region of Zimbabwe. Acta Tropica44:433-444. Chimbelu, E. G., and M. A. Shehata. 1987. A review of researchin Zambiaon Phytolacca dodecandra (Endod). Pages 109-126 in L. Makhubu, CONCLUSION A. Lemma,andD. Heyneman,eds., EndodII (Phytolacca dodecandra);Reportof the Second InterOne of the major objectives of using plant nationalWorkshop on Endod,Phytolaccadodecanmolluscicides in schistosomiasiscontrol is that dra, Mbabane,Swaziland,April, 1986. Councilon the production methodsshouldbe cheap and less International and Public Affairs,New York. labour intensive. The decision on which agri- Defago, G. 1977. The functionof saponinsin the reculturalproductionmethodsare used will be desistance of plantsto parasiticfungi. Bulletin de la terminedby the financialresourcesavailableand Societe BotaniqueSuisse 87:79-132. an assessment of the balance between the cost Donald, C. M., and J. Hamblin. 1976. The biolog-

with earlierestimates of 25% saponins (WoldeYohannesand Kloos 1989). Since the ratioof all the saponins analyzed was almost always the same in berries harvested from both varieties andM0lgaard1994a),it can (Ndamba,Lemmnich be concludedthat cultivationaffected total sapof the inonin contentmore than the proportion dividual components of the mixture. Thus, seof the lection of cultivarswith high proportions most potent saponins would be beneficial because this characterwould not be influencedby the productionmethod. The dependenceof plant saponin content on light, temperature, day length and the cultivation methods has been established (Defago 1977). The accumulationof terpenoids in Helianthus parthenium (Gershenzon 1984) has been described as a response to droughtconditions, in the same way as the haemolyticactivityof Medicago spp. increased under water and nutrient deficits (Vetterand Haraszri1987). Saponinproduction is enhanced in Saponaria (Gershenzon 1984) grown on nitrogen rich soils and under droughtconditions which appearsto be an amplificationof the water deficit response through an increase in evapotranspiration. Similarly,an expected increase in saponin content in P. dodecandraplantsgrownunderdroughtconditions was not seen in plants suppliedwith manurebecause of the water holding capacity of organic matter. Althoughshadinginfluencedsaponincontent, the influence was not statisticallysignificant(P > 0.05). The finding that artificiallyfertilized, watered, and shaded plants producedberries of high saponincontentwas interpreted to be a water stress dependentphenomenon,ratherthan a light intensity response (Tables 3, 4). Given its previouslydiscussedundesirable effects on plant growth and productivity (Ndamba, Lemmich and M0lgaard1994a),coupledwith similarfindings from other independent studies (Lugt 1981), it is unlikely that shading would play a majorrole in P. dodecandracultivation.

to benefit ratio for net saponinyield. Productivity of berrieswith high saponincontent are important,but judging from the influenceof berry yield on net saponin yield, efforts should be made to maximize berry yields as these will inevitably have significanteffect on saponinyield (Table 5, Fig. 9). Cultivationof P. dodecandra with manure and water is desirable since this resulted in the highest saponin yield of 291 g/ plant (Table 5). Since cattle manureis readily available in most ruralareas of Zimbabwe,our resultssuggeststhis as the most ideal cultivation strategy.Althoughmanureis beneficialalso for the waterretainingcapacityof the soil, irrigation is recommendedwhen water is available.
ACKNOWLEDGMENTS
The authors are grateful to the DanishInternational Development Assistance (DANIDA) for financialsupport.The technicalassistanceof NormanMakaza,Kizito Kaondera, CharlesAnderson and VivianChadukura of the BlairResearch is greatlyappreciated. The coLaboratory and assistancerendered operation by the field staff at the KutsagaResearchStation,Dr Rydingand the farmmanager, Mr Zengeya,will alwaysbe remembered. Specialthanks aredueto LegesseWolde-Yohannes for providing rootedcuttingsso crucialto the successof this study.The authors aregrateful to Dr.HenryMadsen, in data DBL,for his assistance assistance of Bibi SpliidandAnnieJ0rgensen for analysis.The technical chemicalanalysesis greatlyappreciated. Thisworkis published withthe kind permission of the Secretary for Health,Zimbabwe.

1996]

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165

vestigationof the diurnal,ontogeneticand seasonal ical yield andharvestindex of cerealsas agronomic variationin the molluscicidal saponin content of and plantbreedingcriteria.Advancesin Agronomy Phytolacca dodecandra aqueous berry extracts. 28:361-405. 35:95-99. Phytochemistry Fluck, H. 1954. The influenceof the soil on the contentof activeprinciplesin medicinalplants.Journal Parkhurst, R. M., B. M. Mthupha, Y. S. Liang, J. I. Bruce, J. D. H. Lambert, T. L. Collier, J. W. and Pharmacology 6:153-163. of Pharmacy ApSimon, L. W. Yohannes, G. E. Heath, W. 0. Gershenzon, J. 1984. Changesin the levels of plant Jones, J. K. Stobaeus, and L. P. Makhubu. 1989. secondary metabolites under water and nutrient The molluscicidalactivityof Phytolaccadodecanstress. Recent Advances in Phytochemistry 18: dra. 1. Location of the activating esterase. Bio273-320. chemical and Biophysical Research CommunicaKawanishi, F., Y. Nagaoka, H. Watanabe,H. Oshio, tion 158:436-439. and K. Nakamoto. 1983. Cultivationof Bupleu, D. W. Thomas, W. A. Skinner, and L. W. of the TakedaResearchLabrum falcatum.Journal Cary. 1973a. Molluscicidalsaponinsof Phytolacoratories42:57-63. IndianJournalof ca dodecandra:Lemmatoxin-C. Lemma, A. 1965. A preliminary reporton the molluscicidal property of endod (Phytolacca dodecanChemistry11:1192-1195. . 1973b. Mol, , , and dra). EthiopianMedicalJournal3:187-190. luscicidal saponins of Phytolacca dodecandra: Litchfield, J. T. (Jr.), and F. Wilcoxon. 1949. A Olenaoglycotoxin-A. Phytochemistry 12:1437simplifiedmethodof evaluatingdose-effectexper1442. iments.Journal of Pharmacology andExperimental . 1974. MolluTherapeutics 96:99-113. , ,and 9 " sicidal saponins of Phytolacca dodecandra: LemLugt, C. B. 1976. The cardiac-glycoside composition matoxin. CanadianJournalof Chemistry52:702of differentDigitalis purpureapopulations.Phar111:441-445. 705. macologicalWeekbulletin . 1981. Phytolacca dodecandra berries as a Shiff, C. J., and V. de Clarke. 1967. The effect of snails. snail surveillance in natural waterways on the means of controllingbilharziatransmitting Litho Printers, Addis Ababa. transmission of Schistosoma haematobium in Rho. 1987. Feasabilityof growth and production desia. CentralAfricanJournal of Medicine13:133of molluscicidal plants. Pages 231-244 in K. E. 137. Mott,ed., PlantMolluscicides.UNDP/World Bank/ Small, E., M. Jurzysta, and C. Nozzolillo. 1990. The WHO Special Programfor Researchand Training evaluation of hemolytic saponin content in wild in TropicalDiseases. JohnWiley & Sons Ltd. and cultivatedalfalfa (Medicagosativa, Fabaceae). EconomicBotany 44:226-235. Marston, A., and K. Hostettmann. 1987. Antifungal, molluscicidal and cytotoxic compounds Tallarida, R. J., and R. B. Murray. 1981. Manual from plants used in traditionalmedicine. Page 27 of pharmacologic calculationswith computerproin K. Hostettmannand P. J. Lea, eds., ProceedNew York. grams.Springer-Verlag, ings of the Phytochemical Society of Europe, Thiilborg, S. T., S. B. Christensen, C. Cornett, C. No. 27: Biologically active naturalproducts.OxE. Olsen, and E. Lemmich. 1993. Molluscicidal ford Science publications, ClarendonPress, Oxsaponins from Phytolacca dodecandra. Phytoford. chemistry32:1167-1171. . 1994. McDermitt, D. K., and D. K. Loomis. 1981. Ele, , , , and mental compositionof biomass and its relationto Molluscicidalsaponinsfrom a Zimbabwean strain of Phytolacca dodecandra. Phytochemistry 36: energycontent,growthefficiencyandgrowthyield. Annalesof Botany 48:275-290. 753-759. Ndamba, J. 1993. Agronomicand other factorsthat Vetter, J., and E. Haraszri. 1987. Haemolyticsapinfluencethe yield, molluscicidalpotencyand saponin contentof herbage.Acta BotanicaHungarica onin contentof the berriesof Phytolaccadodecan33:431-436. dra. Ph.D. thesis. Royal Danish School of Phar- Webbe, G., ed. 1987. The toxicology of molluscicides. International macy, Copenhagen,113 pp. of Pharmacology Encyclopedia and TherapeuticsSection 125. Pergamon Press, , and S. K. Chandiwana. 1988. The geographicalvariationin the molluscicidalpotency of Oxford. Phytolacca dodecandra in Zimbabwe. Tropical and WHO 1965. Molluscicide screeningand evaluation. Medicine40:34-38. Geographical Bulletinof the WorldHealthOrganization 33:567581. , and N. Makaza. 1989. The use of 9 Phytolacca dodecandra berries in the control of . 1983. The controlof schistosomiasis. Report trematode-transmitting snails in Zimbabwe. Acta of a WHO ExpertCommittee. WHOTechnicalReTropica46:303-309. port Series No. 728. WHO Geneva. , E. Lemmich, and P. M0lgaard. 1994. In- Wolde-Yohannes, L. 1983. Past and ongoing agro-

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botanical studies of Phytolacca dodecandra(Endod) in Ethiopia.Pages 125-132 in A. Lemma,D. Heyneman,and S. M. Silangwa, eds., Phytolacca dodecandra (Endod). Towards controlling transmission of schistosomiasis with the use of a natural product.Final Report of the International Work-

shop, Lusaka,Zambia.Tycooly International Publishing Ltd., Dublin. , and H. Kloos. 1989. Agronomicand molluscicidal characteristics of three types of endod (Phytolaccadodecandra). Ethiopian Journal of Agricultural Science 11:25-33.

BOOK REVIEW
Environmental and Economic Dilemmas of Developing Countries. Africa in the Twenty-firstCentury. ValentineUdoh James (ed.). 1994. Praeger Publishers, 88 Post Road West, PO. Box 5007, Westport,CT 06881. xix + 264 pp. (hardcover). $59.95. ISBN 0-275-94666-5. This publicationdiscusses the assumptionthat the challengefacing Africa is a choice betweeneconomic developmentat the expense of the environment,or maintainingthe environment(if possible) at the expense of economicdevelopment. The introduction discusses the need for the assessmentof development projects that have economic and environmental impacts affecting societies. In some countriesrules and regulations for making impact assessmentsare often ambiguous, while in other countriesthere are no rules. Even in the places where there are rules the authors conclude that "the benefitsof the [development] proposals are usually economic and the costs environmental." PartI considersthe negative environmental and social impactsof economic programssupported by nonAfrican institutions.Examples are given of projects thatcould yield positive resultsand thatdo and do not need environmental impactassessmentsuch as renewable resourceuse, industrial activities,and agricultural activitiesand technologytransfer. This sectionemphasizes that the problemsof today result from policies chosen in the last decade.These old policies encourage trans-national companiesfrom industrialized countries to exploit Africa withoutregardfor the environment. This is a degradingapproach. PartII considersthe linkage betweenpoverty,population growth, and degradationof the environment. The authorsrecommenda global partnership between rich nations and African countriesto reduce poverty associatedwith populationgrowth.The conclusion is drawnthat failureto form this partnership will aggravate environmental in Africa and the redegradation mainderof the globe. PartIII detailsthe economicimpacts of resourcedecline. Part IV addressesthe political implicationsof developmentprocesses.Africancountriestypicallywish to develop in a sustainableway without unduly affecting their naturalenvironment. Such development, usually done in one or a few sectors in a country,is futile. The need for cooperation betweengovernmental, non-governmental and the publicis adinstitutions, vocated.Emphasisis placed on the need to recognize traditionalpolitical institutionsin development and conservation.Political instabilityand ethnic regionalism (that counteractsnationalism),both resulting in carelessness about the conservation of natural resources,are otheraspectsof Africa. PartV discusses women as custodiansof the environment.Past ruraldevelopment programs intendedto alleviate povertyhave not given enough assistanceto women. Indeed, some programshave added to women's burden.Severalsocial factorshave contributed to this situation,including:1) lack of female involvement in the planningand design of developmentprograms; 2) lack of understanding thatwomenarethe producers; 3) not understanding that men dominatethe decision making process in rural households, and 4) lack of infrastructure andcreditfacilitiesfor women.The need to involve women, who are the principleactorsin Africanfamily life, in all the planninganddecisionmaking process is summedup by the following statement: "No human group today can progresseconomically, politically and socially without the contributionsof women, who constitutemore than half of the world's population." AlthoughI would have like to have seen more pertinent examples dealing with these issues from other parts of Africa, along with those cited from west Africa, I sincerely recommendthe book to those interested in the Africanenvironment and development.
SEBSEBEDEMISSEW SCIENCEFACULTY ADDIs ABEBA UNIVERSITY

P.O. Box 3434


ADDIs ABEBA, ETHIOPIA

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