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Economic Botany ISSN 0013-0001 Econ BotDOI 10.1007/s12231-014-9267-6

Comparing Medicinal Uses of Eggplant andRelated Solanaceae in China, India, andthe Philippines Suggests the IndependentDevelopment of Uses, Cultural Diffusion,and Recent Species SubstitutionsRachel S. Meyer, Maryam Bamshad,Dorian Q. Fuller & Amy Litt

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Comparing Medicinal Uses of Eggplant and Related Solanaceaein China, India, and the Philippines Suggests the IndependentDevelopment of Uses, Cultural Diffusion, and Recent SpeciesSubstitutions1

RACHEL S. MEYER*,2,3,6, MARYAM BAMSHAD4, DORIAN Q. FULLER5, AND

AMY LITT2

2Genomics Program, The New York Botanical Garden, Bronx, NY 10458, USA3The Graduate School and University Center, City University of New York, 365 Fifth Avenue, NewYork, NY 10016, USA

4Department of Biological Sciences, Lehman College and The Graduate Center, City University ofNew York, 250 Bedford Park Boulevard West, Bronx, NY 10468, USA

5Institute of Archaeology, University College London, 31-34 Gordon Square, London, WC1H 0PY,UK

6Department of Biology, New York University, 12 Waverly Place, New York, NY 10003, USA*Corresponding author; e-mail: rm181@nyu.edu

Comparing Medicinal Uses of Eggplant and Related Solanaceae in China, India, and thePhilippines Suggests the Independent Development of Uses, Cultural Diffusion, and RecentSpecies Substitutions. The ways in which geographically separate communities use cropsreflect the agricultural and cultural influences on each community. The eggplant (Solanummelongena L.; Solanaceae), which was domesticated in South and Southeast Asia, has longbeen used in a variety of medicinal and culinary preparations across many different Asianethnolinguistic groups. Here, we report the total uses for eggplant and sixteen related speciesin three regions, India, southern China, and Malesia, and conduct a comparative analysis inorder to form hypotheses about how influences on plant use in one region could haveaffected use and evolutionary trajectories in other regions. Results from literature review and101 interviews show a total of 77 medicinal attributes for eggplant, with few similar attributesmentioned in different regions, leading us to hypothesize that largely pristine (i.e., withoutinfluence from other regions) development of uses, which could serve as selection pressures,occurred for eggplant in India, southern China, and Malesia. Results also show that manySolanum species have been fluidly adopted into uses developed for other species in a singleregion.

Key Words: Philippine herbal medicine, Ayurveda, Traditional Chinese Medicine, cropdomestication, food, useful plants, nightshades, human migration, artificial selection, speciessubstitution, ethnobotany.

IntroductionCrop evolution under domestication provides a

fascinating representation of the co-evolutionaryprocesses that continue to shape plant genomesand human cultures. Human innovation, culturalexchange, adoption of practices, and many otherfactors have shaped the numerous ways in whichpeople apply selection pressures to crops, whichmay over time change their ability to be eaten,

1 Received 24 June 2013; accepted 2 April 2014;published online ___________

1Received 24 June 2013; accepted 17 January 2014;published online _______.

Electronic supplementary material The online ver-sion of this article (doi:10.1007/s12231-014-9267-6)contains supplementary material, which is available toauthorized users.

Economic Botany, XX(X), 2014, pp. 1–16© 2014, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

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cultivated, or used in applications such asmedicine. Knowledge of these potential selectiveforces can provide insight into key mechanismsand interactions that help explain how desirableplant traits evolved. Reciprocally, variations incrop or species use around the world reflectcultural influences throughout history, and canhelp guide research on the development ofpractices in a given area.There are three scenarios, not mutually exclu-

sive, that can explain the exploitation andinnovation of a given crop in disjunct regions(see Fuller 2011a Table 1): (1) crops may havebeen spread via human migration (“demic diffu-sion”) or trade from one region to another(“cultural diffusion”); (2) exploitation of a speciesin one region, where it is available wild, may havebeen encouraged by people from other regions(“stimulus diffusion”); (3) the crop may havebeen domesticated and diversified from local wildpopulations in different regions by unconnectedcultural groups (primary/pristine domestication).Understanding which of these scenarios occurredin crop histories contributes to our knowledge ofhuman history and addresses the question of whythere is such great varietal diversity in somedomesticated species. In addition, these scenariosare testable hypotheses for many crops that areimportant for archaeological, genetic, and linguis-tic research questions (e.g., Buckler and Stevens2005; Bustorf et al. 2003; de Candolle 1883,1886, 1967; Hutchinson 1974).Many crops have a complicated domestication

history, including multiple origins, and some-times have limited written records of crop uses.This presents a challenge because practices in oneregion could have been influenced by another,complicating our understanding of the history ofcrop evolution and human innovation. In thisstudy we collect and summarize data on the use ofa major global food crop with an unresolveddomestication history, the eggplant (Solanummelongena L.), to test hypotheses regardingcross-regional influences during the early historyof cultivation, drawing primarily on ethnobotan-ical data as well as more limited archaeologicaland historical evidence.The eggplant origin is in tropical Asia, where

fruit and vegetative plant parts have becomewidely used as both a food and a medicine.Although it has long been thought that eggplantwas domesticated on the Indian subcontinent(Lester and Hasan 1991; Polignano et al. 2010;

Sękara 2007; Weese and Bohs 2010) or Indo-Burma (Daunay and Janick 2007), recent popu-lation genetic studies and the presence of highvarietal diversity have led researchers to point toat least China as an additional center of domes-tication. Meyer et al. (2012) proposed threedomestication events, in India, southern China,and the Malay islands (i.e., Malesia, a term usedhere to include Indonesia, Malaysia, thePhilippines, and other countries in the MalayPeninsula and Malay Archipelago). Cericola et al.(2013) found Chinese and Indian eggplantaccessions were genetically divergent, supportinga multiple origin hypothesis. Likewise,Hurtado et al. (2012) supported this notionthrough findings that Sri Lankan and Chineseeggplants were highly divergent. Written recordsare limited, having traced eggplant use in Indiaand China back only 2,000 years, and lackingany evidence that eggplant cultivation haddiffused from one of these places to the other(summarized in Wang et al. 2008; Meyer et al.2012). Therefore, not only are the roles differentregions play in the development of eggplantcultivars unclear, but it is unknown whetherdemic, cultural, or stimulus diffusion affectedselection pressures that contributed to cropevolution.Because no broad surveys were available of

ethnobotanical uses of eggplant in differentregions of Asia, we performed an ethnographicsurvey focusing on three geographically andculturally distinct sites within the Tropical Asiacenter of origin: India, China, and thePhilippines. We began by conducting two litera-ture reviews. The first aimed to trace the earlyconnections (>2,000 years ago) among India,China, and the Philippines in terms of thediffusion of crop plant use through review ofarchaeobotanical and linguistic data, with the goalof forming hypotheses for the dominant direc-tions of influence. The second review was ofpreviously published Asian medicinal texts andwebsites to build a list of written documenteduses for eggplant and related Solanum species.Next, we conducted household interviews in thethree sites to obtain a better resolution of uses.Interview results formed a more complete set ofattributes (i.e., functions of a plant part) foreggplant and related species, and these were usedto test hypotheses on cross-cultural diffusionpatterns as well as to test species interchangeabil-ity in certain preparations.

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Table 1. SUMMARY OFMAJOR EVIDENCE OF ANCIENT HUMANMIGRATION AMONG INDIA, SOUTHERNCHINA, ANDMALESIA.INFORMATION IN ITALICS (OTHER THAN SCIENTIFIC NAME) IS CONTESTED OR BASED ON POORLY SUPPORTED EVIDENCE.

Direction anddate of movement Species From To Reference

Within China5,000 y.a. Rice (Oryza sativa) &

Foxtail millet(Setaria italica)

Middle YangtzeValley

Southern China:Lingnan

Castillo and Fuller2010; Fuller et al.2010; Fuller 2011b

2,400 y.a. Tea (Camellia sinensis) Xishuangbanna,Yunnan

Sichuan Province Keightley 2009

China → India4,000 y.a. Broomcorn millet

(Panicummilliaceum)

China Yemen via Iran/Pakistan/India

Fuller et al. 2011;Boivin et al. 2012;Frachetti et al. 2010

4,000 y.a Rice (Oryza sativa) China NW India viacentral Asia

Molina et al. 2011;Fuller et al. 2010;Fuller 2011a, 2011b

4,000 y.a. Peach (Prunus persica),apricot (Prunusarmeniaca)

China NW India viacentral Asia

Fuller 2011a, b;Boivin et al. 2012

4,000. y.a. Cannabis sativa China/ CentralAsia

Indus, Ganges Fuller 2011a, b;Boivin et al. 2012

China → MalesiaIndia → ChinaMalesia → China

None prior to2,000 y.a.

Malesia → India2 , 0 0 0 – 3 , 0 0 0y.a.

Areca palm (Areca catechu),betel (Piper betel)

Malesia India Fuller 2007; Fulleret al. 2011

~3,400 y.a. Citrus (Citrus spp.) SE Asia India Fuller et al. 2011;Asouti and Fuller 2008

~3,400 y.a. Sandalwood(Santalum album)

Indonesia India’s southernDeccan

Harbaugh and Baldwin2007; Fuller 2007

<4,000 y.a. Banana Malesia India’s Harappan1 Fuller and Madella 2009;Fuller et al. 2010

India → Malesia2,300 y.a. Cotton (Gossypium

arboreum)India Thailand, Malesia2, 3 Cameron 2010, 2011

2,400 y.a Mung bean (Vignaradiata)

India Malay Penninsula Castillo and Fuller 2010;Castillo 2011

2,400 y.a Horsegram (Macrotylomauniflorum)

India Malay Peninsula Castillo and Fuller 2010;Castillo 2011

Within India4 , 2 0 0 – 3 , 8 0 0y.a.

Pulses, wheat, barley SW Asia Southern Deccan andGanges valley

Fuller 2007, 2011a

3 , 9 0 0 – 3 , 5 0 0y.a.

Pigeonpea (Cajanuscajan)

East India andOrissa

Southern India Fuller 2007, 2011a

~3,300 y.a Cotton (Gossypiumarboretum)

Indus valleyregion

Southern India andGanges Valley

Fuller 2008

~3,000 y.a. Rice (Oryza sativa) Ganges South India Fuller 2007, 2011aWithin Malesia4,000 y.a.? Banana (Musa spp.) Philippines New Guinea (some

input local, somefrom outside)

Donohoe & Denham 2009;Kennedy 2009;Fuller et al. 2011

1 Indonesian boat building techniques were also found to be used in India (Fuller et al. 2010; Southworth 2005).2Cotton terminology is also clearly borrowed from Indian languages (Indo-Aryan) into Mon-Southeast Asian languages,especially Mon-Khmer languages (Southworth 2005; Fuller 2008).

3 Indian stone beads and iron work were found in Malay peninsula 2,400 y.a. (Bellina and Glover 2004; Bellina-Pryceand Silapanth 2008).

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MethodsREVIEW OF ARCHAEOBOTANICAL AND LINGUISTIC

LITERATUREBooks and research articles were selected for

r ev i ew by D.Q. Fu l l e r , P ro f e s so r o fArchaeobotany at University College London,who spec ia l i zes in Asian archaeology,paleolinguistics, and early world trade. We soughtto list the well-supported evidence of cropdiffusion between regions of focus prior to2,000 years ago. Ancient linguistic review hadpreviously been carried out by R.S. Meyer andpublished in Meyer et al. (2012) with the help ofexperts in Sanskrit and Chinese linguistics totranslate accurately.

STUDY SITESThe three sites selected for fieldwork were the

Philippines, southern China including nearbycountries Taiwan and Hong Kong (grouped withChina for the remainder of this work), and India.Fieldwork was done during the winters of 2008and 2009. Sites were in tropical and subtropicalregions, and were selected to span a range inaltitude, ethnolinguistic populations, and geogra-phy.

China

Field sites in Taiwan included citiesPingTung, Shanhua, Taipei, Taitung, andTaizhong (Fig. 1). Subjects interviewed werea f f i l i a t e d w i th Ami , Han , o r Ruka iethnolinguistic groups. Fieldwork in HongKong was conducted only in Kowloon withoutregard to specific ethnolinguistic groups.Fieldwork in mainland China included areas inand around the cities of Chengdu, Guangzhou,Guiyang, Jinghong, Kaili, Kunming, and aroundthe countryside of Xishuangbanna, includingGanlanba, Mamushu, and Menglun. Subjectsinterviewed were of Bai, Dai, Han, Hani, or Yiethnolinguistic groups.

The Philippines

Field sites were all on the island of Luzon andincluded the towns and cities of Adams, Baguio,Batac, Dingras, Lagum, Laoag, Los Baños,Magulao, Manila, Peña Blanca, Paoay, andVillasis (Fig. 1). Interviewed subjects belongedto Eastern Ilocano, Pangasinense, Tagalog, orWestern Ilocano ethnolinguistic groups.

India

Field sites included areas within and surroundingtowns and cities of Bishnapur, Chennai, Conoor,Darjeeling, Durghapur, Jaipur, Kerala, Kochi,Kolkata, Mahabalipuram, New Delhi, Ooty,Pune, and Vellore (Fig. 1). Interviewed subjectswere of Bengali, Hindi, Malayam, Marathi,Rajasthani, or Tamil ethnolinguistic groups.

COLLECTION OF ETHNOBOTANICAL

INFORMATION

To obtain information on the contemporaryuses of eggplant, semistructured surveys wereconducted with adults at field sites. Surveys werecarried out by the researchers in Mandarin orEnglish, or translated into the local language byfield assistants. Surveys were designed to followthe ethical standards of the City University ofNew York Institutional Review Board (IRB # 08-12-1612). The number of survey responses wasn = 30 for China, n = 30 for India, and n = 41 forthe Philippines. Survey questions were abouteggplant diversity, culinary use, medicinal use,negative associations or warnings against use,folklore, and uses of other Solanum speciesincluding the wild ancestor of domesticatedeggplant, S. insanum (Knapp et al. 2013). Wecompiled the list of local Solanum species fromthe literature and from plants identified in thefield. Using the common name and photos of theplants, we asked interview subjects what theyknew about the plant and how it could be used.Literature review of published Solanum uses

included books and websites (Table 2 includesprimary sources). The major New World crops inthis genus, potato and tomato (Solanumtuberosum L. and S. lycopersicum L.), were omittedin this study. The following species were lumpedtogether because they were either treated togetherin the local classification system or were difficultto distinguish based on morphology, resulting inthe plants sharing the same use or in ambiguityover which species was intended for use: S.americanum Mill. with S. nigrum L. (referred toherein as S. americanum/nigrum), and S.lasiocarpum Dunal with S. ferox L. (S.lasiocarpum/ferox).Raw interview data and information from

reviewed literature were transformed into tables.The medicinal preparations (ways of using plantparts for a particular treatment) for eggplant andother Solanum species were classified into categories

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representing different non-exclusive attributes (ESM[Electronic Supplementary Material] Appendices 1–3). Information on the food and medicinal uses of

the other Solanum species was also left in narrativeformat to illustrate the types of descriptive dataprovided by informants (Appendix 1).

Table 2. LITERATURE AND WEBSITE SOURCES FOR LOCAL USES OF EGGPLANT AND RELATED SPECIES IN INDIA, CHINA,AND THE PHILIPPINES. TOP SECTION = PRINTED SOURCES, DATES OF CONSULTED TEXT (DATE AND TITLE OF PRIMARY

EARLY WORK. BOTTOM SECTION = WEBSITES.

India China Philippines

Kiritikar and Basu 1918Raghunatha 1956 (17th century,

Bhojanakutuhala)Sastri 1972Lad 1984Muthaliar 1988Ramalingayya and Sastry Vaidya

Yoga Ratnavali 1994Kumari 2004Vardhana 2006Rao 2011

Lan 1940 (1436, Dian Nan BenCao)Li 1982 (1596, Bencao gangmu)Zhao 1969 (1765, Bencao gangmu shiyi)Duke and Ayensu 1985Henry 1986 (1893, Notes on the

economic botany of China)Lu 1986Wu 2006Yang and Gao 2011

Quisumbing 1951Co 1989

lifestyle.iloveindia.com/lounge/benefits-of-brinjal-2032.html (Anonymous2012a)

ayurveda.astrologytutorials.com/vegetables-remedies/eggplant-vartaka/(Anonymous 2012b)

www.ayushveda.com/ayurveda-articles/menstruation.htm (Ayushveda 2012)

www.stuartxchange.org/Talong.html(Umali-Stuart 2012a)

www.chinesemedicinedoc.com/boulder-acupuncture/articles-and-handouts/diet-chinese-medicine/ (Blalack and Blalack2012)

infopedia.nl.sg/articles/SIP_171_2004-12-15.html (Thulaja 2012)

tcmhealthtalk.com/eggplant (Tan 2012)live.shanghaidaily.com/

column_detail.asp?type=column&id=73(Zhang 2009)

www.pingminghealth.com/article/550/eggplant-vegetable-for-protecting-the-large-intestine/ (Anonymous 2009)

www.stuartxchange.org/Talong.html

www.philippineherbs.com/ph i l i pp inehe rb index .h tm(Anonymous 2012c)

www.medicalhealthguide.com/herb/talong.htm (Anonymous2012d)

www.stuartxchange.org/Talong.html (Umali-Stuart2012a)

www.stuartxchange.org/Talong-pipit.html (Umali-Stu-art 2012b)

Chengdu

Kaili

GuiyangKunming

Xishuangbannaloca�ons

Guangdong

Kowloon

Taipei

TaitungPing Tung

ShanhuaTaichung

AdamsTaitung

ManilaVillasis

LaoagBatac

PaoayDingras

Los Baños

Baguio

Darjeeling

DurghapurBishnupur

Kolkata

New Delhi

Jaipur

Pune

ChennaiMahabalipuram

Vellore

Coonoor

CoimbatoreKochi

Ooty

Fig. 1. Map of field sites where interviews were conducted on eggplant and local Solanum use.

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In all three regions eggplant fruits were usedfor food in a variety of ways including both rawand cooked; these data were not included in this

analysis because the variation in cooking styleswas too extensive. Descriptions of medicinal useswere sorted into 77 attributes, and a spreadsheet

Cardiovascular System (8)Positive/Neutral: anti-cholesterol, heart

attack protection, reduces blood pressure,

atherosclerosis, stops bleeding, treats

enlarged spleen, improves circulation,

relieves hemorrhoids

Gastrointestinal System (9) Positive/Neutral: calcium source, fiber source,

increase appetite, good for digestion, meat

replacement, easy to digest, removes food

stagnation, reduces liver problems.

Negative: heavy to digest

Urinary System (4)Positive/Neutral: reduces blood pressure,

diuretic, cleans kidneys

Negative: avoid if you have kidney stones

Reproductive System (6)Positive/Neutral: miscarriage recovery-uterus stabilization, good

for fetus development, good nutrition for lactating mothers, affects

fertility

Negative: ingestion can abort a fetus; avoid if menstruating

Skeletomuscular System (2)Positive/Neutral: heal broken bones

Negative: avoid arthritis

Integumentary System (10)Positive/Neutral: freckle reduction, age spot

reduction, skin wash, replaces bandage.

reduces swelling, improves cell elasticity,

treats skin rashes and irritation

Negative: avoid if skin is sensitive, avoid

because it can cause an allergic reaction

Respiratory System (5)Positive/Neutral: cures respiratory problems,

cures throat problems and cough, reduces

phlegm, increases phlegm

Negative: makes you short of breath

Immune System (13)Positive/Neutral: heal wounds/cuts, treats

skin rashes/irritation, reduces swelling, cures

throat problems/cough, eating reduces fever,

compress reduces fever, compress heals burn,

relieves edema, relieves enlarged spleen,

reduces infection

Negative: can cause allergic reaction, avoid if

bitten by dog, makes you short of breath

Homeostasis (5)Positive/Neutral: healthy, high in nutrients,

reduces heat, gains heat

Negative: avoid if weak

Nervous System (17) Positive/Neutral: weight loss, weight gain, heat gain,

heat loss, narcotic or prevents breakdown of

narcotics, seeds used for toothache, reduces blood

pressure, reduces pain, stomach pain reducers, cure

headache/migraine, protects memory, tranquilizer,

insomnia treatment, eating reduces fever, compress

reduces fever

Negative: makes you short of breath, induces

headache

Body System

Categories and Attributes

Toxicity (8)Positive/Neutral: reduces liver problems, narcotic or

prevents breakdown of narcotics

Negative: avoid combining with other medicines,

avoid-adverse reaction of nausea, avoid if bitten by

dog, avoid eating too much, avoid if weak, avoid if

skin is sensitive

Metabolism (9) Positive/Neutral: weight loss, weight gain, heat gain,

heat loss, diabetes, anti- cholesterol, high nutrients,

atherosclerosis, meat replacement/mimicker

Cancer (3)Positive/Neutral: skin cancer treatment,

treatment for gastrointestinal cancers,

general anticarcinogenic

Heal (9) Positive/Neutral: general topical medicine, heals

wounds/cuts, compress heals burn, compress

cooked, compress raw, heals broken bones,

reduces swelling, replaces bandage, reduces

infection

Endocrine System (9) Positive/Neutral: weight loss, weight gain, heat

gain, heat loss, reduces blood pressure,

miscarriage recovery/uterus stabilization, good

for fetus development, affects fertility

Negative: can induce abortion, can give the fetus

a rash

Fig. 2. Descriptions of health-related attributes given to eggplants. There are 77 health-related attributesidentified from interviews and literature. These were sorted into 15 non-exclusive categories based on body systemor broader health-related characteristic.

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was created categorizing these according to plantpart and fruit color and size: all fruits; purple,green, or white fruits; big or small fruits;v e g e t a t i v e a e r i a l o r g a n s ; a n d r o o t s(Appendices 1–3). Interviewed subjects oftenprovided descriptions of multiple uses.Attributes were classified as beneficial, neutral,or negative depending on whether the useproduced positive effects (e.g., relieving a symp-tom), neutral effects (i.e., a function that can begood or bad depending on the situation, such as adiuretic), or negative effects (e.g., caused gastro-intestinal distress or toxicity). Beneficial/neutralattributes and negative attributes were coded asseparate even if the affected body part was thesame because the functions were different. Forinstance, “cures respiratory problems” was codedas a separate attribute from “makes you short ofbreath” (Fig. 2). If an attribute was onlymentioned in a single interview or literaturesource, it was coded in red (Appendices 2–3).The 77 attributes were assigned to 15 categoriesof body system or body function; assignments ofa single use to multiple categories were made

where appropriate (Fig. 2). To compare usesamong China, India, and the Philippines, Venndiagrams were made to display the number ofuses per category that were unique to each regionand the number that regions had in common(Fig. 3).

Results and DiscussionREVIEW OF HISTORICAL CROSS-CULTURAL

CROP TRADE

Written records from India and China demon-strating that eggplants were being cultivated 2,000years ago (y.a.) in the Charaka and SushrutaSamhitas (Sushruta 600 B.C.E; Charaka 200B.C.E); [Appendix 1; recent translations byBhishagratna 1907, Agnivesa et al., 1984 (1st ed1941), and Sharma and Dash 1983; Swarup 1995;Wang et al. 2008], although evidence of Solanumuse in emergent Neolithic zones in India suggests apossibility of much earlier domestication of egg-plant in central India, such as Rajasthan (Kashyap2007; Kashyap and Weber 2010). There are manypieces of evidence suggesting that Chinese crops

Fig. 3. Venn diagrams of overlapping health-related attributes of eggplant in India, China, and the Philippines.White circle = China; light gray circle = Philippines; dark gray circle = India. Each diagram portrays the attributesin a category of body part or process, e.g., healing, or cancer. See Fig. 2 for breakdown of attributes within eachbroader system or characteristic. Numbers in each section refer to fruit use/ vegetative aerial part use/ root use.

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from temperate regions were adopted in Indiamore than 3,000 y.a. (Table 1; Boivin et al. 2012);however, tropical China, a candidate domestica-tion center, remained relatively disconnected fromtrade with India until more recently, as there is noevidence that Indian crops or language had enteredYunnan (Fuller 2011b). Evidence suggests that it isonly within the past 2,000 years that theXishuangbanna Dai Autonomous Prefecture,which was the proposed Chinese area of domesti-cation in Meyer et al. (2012), joined the SouthAsian-East Asian trade network, through whichagricultural practices could have spread betweenthese putative centers (Beckwith 2009; Fuller2011b). Therefore, it is possible that domesticationof eggplant as a food began independently in Indiaand tropical China, but when these regions becamenetworked ca. 2,000 y.a., information abouteggplant use could have diffused.Evidence reviewed from other domesticated

species in South Asia (including India) and EastAsia (including China) showed that multiplepristine/independent domestications of the samespecies were common. Examples from livestockinclude chicken (Gallus gallus L.; Kanginakudruet al. 2008; Mwacharo et al. 2011) and waterbuffalo (Bubalus bubalis L.; see Fuller [2007] andreferences therein). Rice (Oryza sativa L.) is anexample of a plant crop domesticated in bothIndia and China; although the lineage domesti-cated in China was widely disseminated and is amajor global staple today, it is generally accepted

that rice was independently domesticated in Indiaas a minor crop that later was replaced by Chineserice (Fuller 2011a, 2011b; Glover and Higham1996; Molina et al. 2011). Although many cropswere domesticated in India, most ancient evi-dence supports the subsequent adoption of manyChinese agricultural practices into India and notin the reverse direction (Bellwood 2005; Fuller2007; Glover and Higham 1996; Higham 2003;Table 1). These patterns led us to hypothesizethat if there was stimulus or cultural diffusion ofeggplant cultivation and uses, the direction wouldmore likely have been from China to India(Diffusion hypothesis 1 [Hyp 1]).The history of eggplant in Malesia and the

connectedness of this region with other putativedomestication locales is poorly understood com-pared to India and China. There are no writtenrecords from antiquity regarding eggplant orother crops in Austronesian languages (Malesianlanguages); knowledge was largely passed downorally. The earliest documented Malay andTagalog writings only date to the 7th centuryC.E. (Blust 1990). Therefore we could not lookto ancient written records to investigate thepossibility of ancient eggplant exploitation inMalesia.In addition to there being no ancient record for

eggplant in Malesia, there is limited evidence thatcrops and agricultural practices from other regionswere adopted into Malesian culture prior to about2,400 y.a. During the Neolithic, cereal cultiva-

Table 3. NATIVE HABITATS OF 16 SPECIES CLOSELY RELATED TO EGGPLANTS, AND WHETHER THEY ARE INVASIVE,CULTIVATED, OR ORNAMENTAL (NOT EXCLUSIVE CATEGORIES).

Species Native range Invasive? Cultivated? Ornamental?

S. aethiopicum Tropical Africa + +S. americanum/nigrum Tropical and subtropical world +S. capsicoides Tropical and subtropical Americas +S. dulcamara Europe and Asia +S. erianthum Tropical and subtropical Americas + +S. insanum Tropical and subtropical AsiaS. lasiocarpum/ferox Tropical India to Pacific Islands + +S. lyratum Parts of East and Southeast AsiaS. macrocarpon Tropical Africa + +S. pseudocapsicum Central and South America + +S. spirale Paleotropical +S. torvum Caribbean + +S. trilobatum IndiaS. viarum South America +S. violaceum Tropical Asia + +S. virginianum Africa, SW and SE Asia, Pacific Islands

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tion (Oryza sativa and Setaria italic [L.] P.Beauvois.) spread overland and possibly via thecoast from China through mainland SoutheastAsia (Thailand and Vietnam) (Castillo 2011;Weber 1991), and there may have been someminor adoption of rice within MaritimeSoutheast Asia (e.g., Bellwood 2011; Fuller et al.2010). Nevertheless, clear evidence for theadoption of other crops and for trade with otherregions dated to less than 2,500 y.a. This includesevidence for trade between southeast China,Taiwan, Vietnam, and Malesia seen in Nephriteornaments (Hung et al. 2007), and probablemovement of textile spinning technology andfiber crops of Chinese origin (Cameron 2011). Inaddition, contacts with India were indicated byceramics and beads (Bellina and Silapanth 2008),cotton textiles (Cameron 2010, 2011), and arange of Indian crops, especially pulses such asmungbean (Vigna radiate [L.] R. Wilczek), horsegram (Macrotyloma uniflorum [Lam.] Verdc.), andpigeon pea (Cajanus cajan [L.] Millsp.) (Castilloand Fuller 2010). This same period (2,500 y.a.)provided suggestive evidence that Malesian cropswere becoming well known in south India basedon linguistic evidence for coconut and arecapalms (Cocos nucifera L., Areca catechu L.)(Fuller 2007). Even earlier, perhaps by 3,300y.a., sandalwood (Santalum album L.) of Malesianorigin first found its way to South India, based ondated charcoal remains (Asouti and Fuller 2008;Fuller et al. 2011). There is no evidence as towhether eggplants were also known or exchangedat this period. However, evidence from trade ofother items and species suggests cultural diffusionwould have proceeded in the direction fromMalesia into India (Hyp 2).

INTERVIEW AND LITERATURE-BASED SURVEY OF

EGGPLANT AND SOLANUM USE

The data gathered from 101 interviewedindividuals, 19 books and print publications,and 12 websites, included information about theuses of eggplant and related Solanum speciesincluding the wild progenitor, S. insanum. A totalof 200 medicinal preparations were recorded foreggplant from all three regions and including allplant parts and fruit colors and sizes(Appendix 2). Of these, 82 were mentioned inthe literature (Appendix 3; Table 2), and 196 ininterviews, with 78 preparations in common.Only four preparations were found only in the

literature: avoidance during menstruation (India),topical use of boiled leaves for hemorrhoids(China) or to reduce swelling (Philippines), andingestion of leaves to reduce fever (Philippines)(Appendix 3). These 200 preparations wereclassified into 77 categories of medicinal attri-butes (Appendix 2; Fig. 2). For instance, inChina, white, green, and small eggplants wereeaten to cause weight loss; therefore these threeapplications were listed under the attribute“weight loss.” The eggplant medicinal attributesbased on total evidence were compared amongthe three regions by clustering attributes intobroader health categories (Fig. 3).

MEDICINAL AND CULINARY USES OF WILD

SPECIES IN INDIA, CHINA, AND THE PHILIPPINES

In the three regions included in this study,sixteen Solanum species (Table 3 lists origins) orspecies groups in addition to S. melongena wereidentified as food or medicine (Appendix 1).Some Solanum species were used in the same waysin all regions, while other species were not used inall regions. The vegetative aerial parts of S.americanum/nigrum were used everywhere underthe attribute of skin problem treatment, such asfor sores. Other attributes of S. americanum/nigrum in China and India were the same,including as a purgative, to ease urination, andto increase sex drive. S. insanum uses were varied,but topical skin wash was common in thePhilippines and India. Solanum lasiocarpum/feroxwas used as a food in China and the Philippines,though it was also reported in the Philippines tobe mildly toxic. S. macrocarpon L. was consumedas a food in India and the Philippines(Appendix 1), and S. torvum Sw. was consumedas food in all three regions. India and China hadcommon uses for S. torvum related to gastroin-testinal wellness (Appendix 1). S. violaceumOrtega was eaten as a food only in India andChina, but interviewed subjects in all threeregions listed several medicinal uses of thisspecies; the attribute of gastrointestinal wellnesswas common for S. violaceum in all regions.

The data contain examples of adoption of non-native species into the same practices implement-ed with indigenous species. S. macrocarpon and S.aethiopicum were both domesticated in Africa buthave spread into Asia. In its native range of sub-Saharan Africa, S. macrocarpon is grown for edibleleaves in addition to fruit. Informants in India

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and the Philippines described it as looking andtasting similar to S. melongena, suggesting thattheir use of the fruit and not the leaves is basedon its similarity to eggplant (Appendix 1). Thefruits of S. aethiopicum are consumed as food andare reported to have a very bitter flavor. People inChina associate the flavor with the local bittermelon (Momordica spp.), and cook it as food anduse it for medicine in similar ways, such as tolower blood sugar (Appendix 1).The most common preparation of Solanum

species, including data from all regions, recordedfor five species, was to use heated and pulverizedor smoked seeds to relieve toothache (attribute“seeds used for toothache”). The next mostfrequently recorded medicinal attributes (4 spe-cies each) were as a tranquilizer/soother, as a skinirritation reliever, and as a component of theIndian Ayurvedic formula “Dasamoola” (seebelow). Induction of vomiting, increasing sexdrive, reducing liver problems, relieving cough,and lowering heat in the body, were eachattributes for three species. The following attri-butes were assigned to only two species: diuretic,laxative, reliever of gastrointestinal discomfort, forlowering blood sugar, and for treating rheuma-tism (Appendix 1).Our data suggest that multiple species in

Solanum can have the same attributes and mayeven have been used as substitutes for oneanother in certain preparations. For instance,Dasamoola is a combination of the roots of tenherbs, including two Solanum species, that is usedas a tonic to normalize the neuroendocrine systemby regulating various hormone functions (Dashand Kashyap 1980). The traditional formulaincludes S. violaceum and S. virginianum L., bothspecies native to India (Dutt and King 1877,1922; Table 3). However, our data revealed thatS. viarum Dunal and S. torvum, both native toSouth America, were being substituted for theoriginal species used in India, possibly since thesame common names are used, because S. viarumand S. torvum are easy-to-find invasive species, orbecause they may share similar secondary metab-olites with S. violaceum and S. virginianum.

MEDICINAL USES OF DOMESTICATED EGGPLANT

IN INDIA, CHINA, AND THE PHILIPPINES

The 77 medicinal attributes identified foreggplant (Appendices 2–3) were classified into15 categories according to which parts or pro-

cesses of the body were affected (Fig. 2). Thecategory with the most attributes was the nervoussystem, followed by the immune and integumen-tary systems (n = 17, 13, and 10 attributes,respectively). Nine attributes were found in thecategories of endocrine system, gastrointestinalsystem, metabolism regulation, and healing. Inaddition to positive or neutral kinds of attributes,negative attributes were identified for all but fourcategories: cancer, healing, metabolism, andcardiovascular system.In each country studied, at least one attribute

was identified for eggplant in each of the 15categories with the exception of the category forcancer, which was not mentioned in thePhilippines (Fig. 3). Among countries, the specif-ic attributes in each category varied extensively.China accounted for the highest number ofattributes in the categories of cancer and thecardiovascular system (Fig. 3). Likewise, thePhilippines accounted for the highest number inhealing, skeletomuscular, and immune systemcategories. India accounted for the highest toxic-ity, metabolism, gastrointestinal, respiratory, andnervous system categories. Relative to the numberof attributes listed for each country, the numberof overlapping attributes was very low: there were16 medicinal attributes common to all threeregions, and these were most numerous in thehomeostasis, immune, and integumentary sys-tems categories (for each category, n = 3attributes; Fig. 3). The only universal attributeswere the use of plant shoots as a skin wash to treatirritations, rashes, and wounds and as an anti-inflammatory. Categories with less than 15% ofuses in common between at least two countrieswere the endocrine system, respiratory system,and cancer. Nearly half (7/15) of the categoriesdid not have a single attribute that was found inall three regions (Fig. 3). Overall the data suggestthat differences in attributes predominate oversimilarities among all three regions.

INTERPRETING COMMON ATTRIBUTES

In spite of the conspicuous differences, pairs ofsites do share attributes, which may be evidenceof influences from one region to another relatedto eggplant cultivation and use. In both India andChina, eggplant was viewed as heavy to digest,able to increase heat in the body, and able toincrease appetite, and was viewed as similar tomeat (i.e., meat replacement). In both countries

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eating eggplant was said to reduce liver problems,remove stagnant food from the colon, preventatherosclerosis, and lower cholesterol levels. Noneof these attributes were unique to eggplant—eachwas mentioned at least once for related Solanumspecies. These commonalities could be based incommon elements of Traditional ChineseMedicine (TCM) and Ayurvedic pharmacopoeias,such as the assignment of hot and cold charac-teristics to food ingredients. For instance, in bothsystems, attributes such as increasing appetite andbeing heavy to digest are linked to foods thatincrease heat (Lad 1984), and cooling foods arerecommended to counterbalance hot tendencies.In both India and China, large eggplants wereassociated with stimulating digestive activity andelevating body heat. These common attributesrelated to heat were absent in data from thePhilippines.

China and the Philippines shared attributesrelated to the integumentary system and healingcategories. In both regions multiple informantsindicated that the fruits are used topically toimprove elasticity of the skin, and as a skin washor compress for various ailments. The sliced fruitscan be applied and left on the skin like a bandage.In contrast, raw or cooked fruits are not used ascompresses in India, although a single informantin India said her family used the fruit peel as acompress for treating burns. Fruit compresses ingeneral were not mentioned for wild species. Thissuggests either the practice of using fruit com-presses developed independently in China and thePhilippines, and because the eggplant wildprogenitor, S. insanum, was not used in com-presses in either region, this parallel use couldhave developed after eggplant was domesticated,suggesting cultural diffusion (Hyp 1).

India and the Philippines shared numerousfeatures of eggplant medicinal use, particularly forvegetative parts. While fruits were by far the mostwidely used plant part for medicine in all threecountries, informants in India and the Philippineslisted many more attributes for vegetative partsthan those in China (Fig. 3). Data from Chinacontained only 5 attributes for shoots and 3 forroots; likewise, data from India contained 19 forshoots and 6 for roots, and the Philippines had 28for shoots and 17 for roots (Fig. 3; Appendix 2).Although only a few attributes for eggplantvegetative parts were recorded for China, wefound evidence that the roots of other Solanum

species were boiled and drunk as a medicinal tea(Appendix 1). Solanum roots were also widelyused in India, as exemplified by their inclusion inthe Dasamoola preparation. However, that thehighest number of root attributes was recordedfor the Philippines is an illustration of Malesia’sunique pharmacopoeia for eggplant and suggeststhat root uses may have ancient origins in theregion. Malesia is the site of domestication ofmany root and tuber crops, and these were themain staples rather than the seed-focused agricul-tural traditions in India and China. Therefore it ispossible that root use knowledge and technologyhad developed much more in Malesia and hadinfluenced other regions (Hyp 2).

DIFFERENCES BETWEEN MEDICINAL ATTRIBUTES

OF WILD AND DOMESTICATED EGGPLANT

In order to assess whether inhabitants of anyregions use eggplant and its wild progenitor, S.insanum, for the same purpose, medicinal attri-butes of the two species were compared. S.insanum fruit is consumed in all three regions,although it is unclear whether it is ever consumedfor nutritional value alone, because the descrip-tions of consumption were always linked withmedicinal attributes (Appendix 1). In India, S.insanum seeds are used to treat toothache, andunspecified plant parts are consumed to relievecough and are topically applied to relieve skinproblems. Attributes of domesticated eggplant inIndia include all those of the wild progenitor andmore. In China, the fruit of both the S. insanumand S. melongena were ingested to relieve liverproblems, and both were associated with increas-ing heat. In the Philippines, gastrointestinalapplications were conserved between the species.In contrast, informants in the Philippines gavewarnings about eating S. insanum. For example, itwas recommended to avoid S. insanum whilepregnant, but domesticated eggplants were asso-ciated with helping the fetus develop, and theroot was used to promote uterus stabilization aftera miscarriage. This difference between wild anddomestic may be related to the chemical changesthat accompanied domestication of eggplant,particularly the lowering of steroidal alkaloids(see examples in Meyer 2012; Wu et al. 2013).

ConclusionsThis study has shown that eggplant has a wide

range of medicinal uses in Asia (Fig. 2) that differ

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substantially across regions (Fig. 3). Seventy-sevenmedicinal attributes were recorded for eggplant,which indicates eggplant is important in localmedicine and has promise as a functional foodand in the natural products industry. Mostsubjects (65%) interviewed mentioned at leastone medicinal preparation for eggplant, and ofthe informants that mentioned medicinal use,89% mentioned more than one. Informants oftendescribed different varieties of domesticated egg-plant as having different medicinal qualities(Appendix 2), implying varietal diversity isimportant for maintaining the full therapeuticcapacity of the species.There was near complete congruence (95%) of

attributes recorded from literature/web sourcesand from interviews (Appendices 2–3). Some ofthese new uses appear to be novel, such as the useof eggplant fruit compresses for healing, and someare based on species substitution, such as theincorporation of non-native species S. torvum andS. viarum in the Dasamoola formula, or the use ofS. macrocarpon fruit because it looks similar toeggplant. These patterns of conserved or noveluses may be indicators of common chemistryamong different species, and may also be a basisfor explorations of therapeutic capacity of theseplants.In addition, many of the attributes mentioned

assigned to the fruit, such as weight loss, may berelated to phytochemical content. For example,eggplants have been shown to be high inchlorogenic acid (mentioned to have anti-obesityeffects in Plazas et al. 2013) and other health-beneficial phenolic compounds (Meyer 2012;Stommel and Whitaker 2003). It would be ofinterest to further explore whether the chemicalmakeup of the fruit, leaves, and roots may helpexplain the basis of the medicinal attributesdescribed in this work.The diverse medicinal attributes of Solanum

species lead to the question of whether culturaldiffusion influenced practices in certain regions.The regional differences in the attributes of wildand domesticated species reinforces the hypothe-sis that selection in these three regions of India,southern China, and Malesia occurred largelypristinely. However, it is possible that the fewuses that are attributed to multiple Solanumspecies including domesticated eggplant, such asuse of seeds to relieve toothache or the use ofplant parts as ingredients for a skin wash tominimize irritation or rash, were transmitted from

one region to another via early demic diffusion,perhaps even before domestication occurred (Hyp3). On the other hand, there is some evidencethat supports the notion that cultural diffusioninfluenced Solanum pharmacological practices.The common utilization of vegetative organs,particularly roots, for medicine in India andMalesia suggests that these regions might haveinfluenced each other in the development ofeggplant uses. Malay medicine is known to havebeen influenced by the practices of various otherAsian medical systems, including those of Indiaand China (Laderman 1993; Winstedt 1944);therefore it is plausible that uses were adoptedfrom India to the Philippines. However, thehigher number of root attributes in thePhilippines demonstrates that uses probablydeveloped in Malesia and not by diffusion basedon patterns from other crops (Table 1), andalthough more work is needed to understand thedevelopment of root use knowledge, diffusion inthe direction from Malesia to India is plausible(Hyp 2). Influence between Malesia and Indiawould also be congruent with what is knownabout early trade, during which India and Malesiahad many interactions but China and Malesia didnot (Appendix 1). Regarding India and China,some shared attributes mentioned reflect com-monalities in traditional medicine such as thosepertaining to heat; there is no apparent trendsupporting a direction of influence (Hyp 1).Expanded archaeolinguistic and archaeobotanicalinvestigations in Tropical Asia that consider cropwild relatives and multiple plant parts mayuncover key evidence to better understand cropevolution as well as human cultural evolution.

AcknowledgmentsThis work was supported by The New York

Botanical Garden Genomics Program, the Chat-ham Fellowship in Medicinal Botany, and TheGarden Club of America. R. Meyer was support-ed by the City University of New York ScienceFellowship, the National Science FoundationGK-12 Program, and the National ScienceFoundation Plant Genome Postdoctoral Fellow-ship during the research or writing of this work.Great thanks are given to the following peoplewho aided with interviews and literature searches:D. Tandang, D. Madulid, A. Tungpalan, D.McClelland, Y.F. Zhao and family, S. Harrell, S.Ahmed, J. Sathe, A. Paul, D. Deb, the faculty ofTamil Nadu Agricultural University, the Kun-

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ming Institute of Botany, the Deccan CollegeSanskrit Dictionary Project, the Asian VegetableResearch and Development Center, and theNational Bureau of Plant Genetic Resources.The authors express their gratitude to thecommunities that have continued to share andcelebrate traditional plant use knowledge.

Literature CitedAgnivesha, Vaidya Yadavji Trikamji, Charaka,

Dridhabala, Chakrapanidutta, ed. 1984 (1st

edn. 1941). The Charakasamhita by Agnivesa.Chaukhamba Sanskrit Sansthan, Varanasi, India.

Anonymous. 2009. Eggplant and food stagna-tion. Yokine, Australia: Ping Ming Health.http://www.pingminghealth.com/article/550/eggplant-vegetable-for-protecting-the-large-in-testine/ (8 June 2012).

———. 2012a. Benefits of brinjal. http://lifestyle.iloveindia.com/lounge/benefits-of-brinjal-2032.html (8 June 2012).

———. 2012b. Eggplant (Vartaka). http://ayurveda.astrologytutorials.com/vegetables-remedies/eggplant-vartaka/ (8 June 2012).

———. 2012c. Philippine herb index. Laguna,Philippines: Mt. Banahaw Health ProductsCorp. http://www.philippineherbs.com/philippineherbindex.htm (8 June 2012).

———. 2012d. Talong herbal medicine. Philip-p i n e s : M e d i c i n a l H e a l t h Gu i d e .www .med i c a l h e a l t h gu i d e . c om/h e r b /talong.htm (8 June 2012).

Ayushveda. 2012. Menstruation—Useful herbsand Ayurvedic treatments. Panchkula, India:A y u s h v e d a I n f o r m a t i c s . h t t p : / /www.ayushveda.com/ayurveda-art icles/menstruation.htm (8 June 2012).

Asouti, E. and D. Q. Fuller. 2008. Trees andwoodlands in South India: An archaeologicalperspective. Left Coast Press, Walnut Creek,California.

Beckwith, C. I. 2009. Empires of the Silk Road:A history of Central Eurasia from the BronzeAge to the present. Princeton University Press,Princeton, New Jersey.

Bellina, B. and I. Glover. 2004. The archaeologyof early contact with India and the Mediter-ranean world, from the fourth century BC tothe fourth century AD. Pages 68–88 in I.Glover and P. Bellwood, eds., Southeast Asia:From prehistory to history. Routledge Curzon,London.

Bellina-Pryce, B. and P. Silapanth. 2008. Weavingcultural identities on trans-Asiatic networks:Upper Thai-Malay Peninsula—An early socio-political landscape. Bulletin de l’Ecole Françaised’Extrême-Orient 93:257–93.

Bellwood, P. 2005. First farmers: The origins ofagricultural societies. Blackwell, Oxford, U.K.

——— 2011. Holocene population history inthe Pacific region as a model for worldwidefood producer dispersals. Current Anthropol-ogy 52:S363–S378.

Bhishagratna, K. 1907. An English translation ofthe Sushruta Samhita based on original San-skrit text. K. Bhishagratna, Calcutta, India.

Blalack, J. and K. Blalack. 2012. Diet—Chinesemedicine. http://www.chinesemedicinedoc.com/boulder-acupuncture/articles-and-handouts/diet-chinese-medicine/ (8 June 2012).

Blust, R. 1990. Summary report: Linguisticchange and reconstruction methodology inthe Austronesian language family. Pages 133–154 in P. Baldi, ed., Trends in linguistics:Studies and monographs 45. Mouton deGruyter, Berlin, Germany.

Boivin, N., D. Q. Fuller, and A. Crowther. 2012.Old World globalization and the Columbianexchange: Comparison and contrast. WorldArchaeology 44:452–469.

Buckler, E. S. and N. M. Stevens. 2005. Maizeorigins, domestication, and selection. Pages67–90 in T. J. Motley, N. Zerega, and H.Cross, eds., Darwin’s harvest. Columbia Uni-versity Press, New York.

Bustorf, D., D. Martinez d’Alis-Moner, D.Nosnitsin, T. Rave, W. Smidt, and E.Sokolinskaia, eds. 2003. EncyclopaediaAethiopica. Harrassowitz Verlag, Wiesbaden,Germany.

Cameron, J. 2010. The archaeological textilesfrom Ban Don Ta Phet in broader perspective.Pages 129–140 in B. Bellina, E. A. Bacus, O.Pryce, and J. Weissman Christie, eds., Fiftyyears of archaeology in Southeast Asia: Essaysin honour of Ian Glover. River Books,Bangkok/London.

——— 2011. Iron and cloth across the Bay ofBengal: New data from Tha Kae, centralThailand. Antiquity 85:559–567.

Castillo, C. 2011. Rice in Thailand: Thearchaeobotanical contribution. Rice 4:114–120.

——— and D. Q. Fuller. 2010. Still toofragmentary and dependent upon chance?

MEYER ET AL.: MEDICINAL USES OF EGGPLANT IN ASIA2014]

Author's personal copy

Advances in the study of early Southeast Asianarchaeobotany. In: Fifty years of archaeologyin Southeast Asia: Essays in honour of IanGlover, eds. B. Bellina, E. A. Bacus, O. Pryce,and J. Weissman Christie, 91–111. RiverBooks, Bangkok/London: River Books.

Cericola, F., E. Portis, L. Toppino, L. Barchi, N.Acciarri, T. Ciriaci, T. Sala, G. L. Rotino, andS. Lanteri. 2013. The population structureand diversity of eggplant from Asia and theMediterranean Basin. PLoS One 8:e73702.

Charaka (dates debated: 200 B.C.E.–100 C.E.).Caraka Samhita.

Co, L. 1989. Common medicinal plants of theCordillera region (northern Luzon, Philip-pines). Chestcore, Baguio City, Philippines.

Dash, B. D. and L. Kashyap. 1980. MateriaMedica of Ayurveda. Conocept PublishingCompany, New Delhi, India.

Daunay, M. C. and J. Janick. 2007. History andiconography o f eggp l ant . Chron icaHorticulturae 47:16–22.

de Candolle, A. 1883. Origine des plantescultivées. Baillière Germer, Paris, France.

——— 1886. Origin of cultivated plants. KeganPaul, Trench and Co., London.

———. 1967 (reprint of 2nd ed., 1886). Originof cultivated plants. New York: Hafner Pub-lishing Company.

Donohoe, M. and T. Denham. 2009. Banana(Musa spp.) domestication in the Asia-Pacificregion: Linguistics and archaeobotanical per-spectives. Ethnobotany Research and Applica-tions 7:293–332.

Duke, J. A. and E. S. Ayensu. 1985. Medicinalplants of China. Reference Publications, Inc.,Algonac, Michigan.

Dutt, U. C. and G. King. 1877. The materialmedica of the Hindus, compiled from Sanskritmedical works. Thacker, Spink, and Co.,Calcutta, India.

———. 1922 (revised ed., originally published in1877). The material medica of the Hindus.Calcutta, India: Adi-Ayurveda Machine Press.h t t p : / / a r c h i v e . o r g / d e t a i l s /mobot31753000838638.

Frachetti, M. D., R. N. Spengler, G. J. Fritz, andA. N. Mar’yashev. 2010. Earliest direct evi-dence for broomcorn millet and wheat in theCentral Eurasia steppe region. Antiquity84:993–1010.

Fuller, D. Q. 2007. Non-human genetics, agri-cultural origins and historical linguistics in

South Asia. Pages 393–443 in M. Petragliaand B. Allchin, eds., The evolution andhistory of human populations in South Asia.Springer, Amsterdam, Netherlands.

———. 2008. The spread of textile productionand textile crops in India beyond theHarappan zone: An aspect of the emergenceof craft specialization and systematic trade. In:Linguistics, archaeology and the human past,Occasional paper 3, eds. T. Osada and A.Uesugi, 1–26. Kyoto, Japan: Research Insti-tute for Humanity and Nature.

———. 2011a. Finding plant domestication inthe Indian subcontinent. Current Anthropol-ogy 52:S347–S362.

———. 2011b. Pathways to Asian civilizations:Tracing the origins and spread of rice and ricecultures. Rice 4:78–92.

——— and M. Madella. 2009. Banana cultiva-tion in South Asia and East Asia: A review ofthe evidence from archaeology and linguistics.Ethnobotany Research and Applications7:333–51.

———, I. Sato, C. Castillo, L. Qin, A. R.Weisskopf, E. J. Kingwell-Banham, J.-X Song,S.-Mo Ahn, and J. van Etten. 2010. Consil-ience of genetics and archaeobotany in theentangled history of rice. Archaeological andAnthropological Sciences 2:115–131.

———, N. Bovin, T. Hoogervorst, and R.Allaby. 2011. Across the Indian Ocean: Theprehistoric movement of plants and animals.Antiquity 85:544–558.

Glover, I. C. and C. F. W. Higham. 1996. Newevidence for early rice cultivation in South,Southeast and East Asia. Pages 413–441 in D.R. Harris, ed., The origins and spread ofagriculture and pastoralism in Eurasia. UCLPress, London.

Harbaugh, D. T. and B. G. Baldwin. 2007.Phylogeny and biogeography of the sandal-woods (Santalum, Santalaceae): Repeated dis-persals throughout the Pacific. AmericanJournal of Botany 94:1028–1040.

Henry, A. 1893. Notes on the economic botanyof China. Presbyterian Mission Press, Shang-hai, China.

———. 1986 (reprint of 1893 issue). Notes oneconomic botany of China. Kilkenny, Ireland:Boethius Press.

Higham, C. F. W. 2003. Languages and farmingdispersals: Austroasiatic languages and ricecultivation. Pages 223–232 in C. Renfrew

ECONOMIC BOTANY [VOL XX

Author's personal copy

and P. Bellwood, eds., Examining the farming/language dispersal hypothesis. McDonald In-stitute for Archaeological Research, Cam-bridge, U.K.

Hung, H.-C., Y. Iizuka, P. Bellwood, K. D.Nguyen, B. Bellina, P. Silapanath, E. Dizon,R. Santiago, I. Datan, and J. H. Manton.2007. Ancient jades map 3,000 years ofprehistoric exchange in Southeast Asia. Pro-ceedings of the National Academy of Sciencesof the U.S.A. 104:19745–19750.

Hurtado, M., S. Vilanova, M. Plazas, P.Gramazio, H. H. Fonseka, R. Fonseka, andJ. Prohens. 2012. Diversity and relationshipsof eggplants from three geographically distantsecondary centers of diversity. PLOS ONE7:e41748.

Hutchinson, J. 1974. Evolutionary studies inworld crops: Diversity and change in theIndian subcontinent. Cambridge UniversityPress, Cambridge, U.K.

Kanginakudru, S., M. Metta, R. D. Jakati, and J.Nagaraju. 2008. Genetic evidence from Indianred jungle fowl corroborates multiple domes-tication of modern day chicken. BMC Evolu-tionary Biology 8:174.

Kashyap, A. 2007. Use-wear and starch-grainanalysis: An integrated approach to under-standing the transition from hunting gatheringto food production at Bagor, Rajasthan, India.Ph.D. thesis, Michigan State University.h t t p : / / g r a dw o r k s . um i . c om / 3 2 / 4 8 /3248560.html (15 January 2011).

——— and S. Weber. 2010. Harappan plant userevealed by starch grains from Farmana, India.Antiquity 84.

Keightley, D. N., ed. 2009. The origins ofChinese civilization. University of CaliforniaPress, Los Angeles.

Kennedy, J. 2009. Bananas: Towards a revisedprehistory. Pages 190–204 in A. Fairbairn andE. Weiss, eds., From foragers to farmers.Papers in honour of Gordon C. Hillman.Oxbow, Oxford, U.K.

Kiritikar, K. R. and B. D. Basu. 1918. Indianmedicinal plants. Sudhindra Nath Basu, MBPanini Office, Bhuwanéswari Asrama,Bahadurganj, Allahabad, India.

Knapp, S., M. Vorontsova, and J. Prohens. 2013.Wild relatives of the eggplant (Solanummelongena L.: Solanaceae): New understandingof species names in a complex group. PLoSOne 8:e57039.

Kumari, R. M. 2004. A taxonomic revision of theSolanaceae. Ph.D. thesis, Bharathiar Universi-ty, India.

Lad, D. 1984. The science of self-healing—Apractical guide. Lotus Press, Twin Lakes,Wisconsin.

Laderman, C. 1993. Taming the wind of desire.Psychology, medicine, and aesthetics in Malayshamanistic performance. University of Cali-fornia Press, Los Angeles.

Lan, M (1397–1476), ed. 1436. Dian-NanBencao (Materia medica of southern Yunana).

———. 1940 (reprint). Diannan bencao. Kun-ming, China: Harrassowitz Verlag.

Lester, R. N. and S. M. Z. Hasan. 1991. Originand domestication of the brinjal eggplant,Solanum melongena, from S. incanum, inAfrica and Asia. Pages 369–388 in J. G.Hawkes, R. N. Lester, M. H. Nee, and N.Estrada, eds., Solanaceae III: Taxonomy,chemistry, evolution. Royal Botanic Gardens,Kew, London, U.K.

Li, Shi-Zhen (1552–1593). (Bencao gangmu)(1596).

———. 1982 (reprint of 1596 original). Bencaogangmu. Beijing, China: .

Lu, H. C. 1986. Chinese system of food cures:Prevention and remedies. Sterling Publishers,New York.

Meyer, R. S., K. G. Karol, D. P. Little, M. H.Nee, and A. Litt. 2012. Phylogeographicrelationships among Asian eggplants and newperspectives on eggplant domestication. Mo-lecular Phylogenetics and Evolution 63:685–701.

Molina, J., M. Sikora, N. Garud, J. M. Flowers,S. Rubinstein, A. Reynolds, P. Huang, S.Jackson, B. A. Schaal, et al. 2011. Molecularevidence for a single evolutionary origin ofdomesticated rice. Proceedings of the NationalAcademy of Sciences of the U.S.A. 108:8351–8356.

Muthaliar, M. 1988. Siddha materia medica(Vegetable section), Vol. 1. Tamilnadu SiddhaMedical Council, Chennai, India.

Mwacharo, J. M., G. Bjørnstad, V. Mobegi, K.Nomura, H. Hanada, T. Amano, H. Jianlin,and O. Hanotte. 2011. Mitochondrial DNAreveals multiple introductions of domesticchicken in East Africa. Molecular Phyloge-netics and Evolution 58:374–382.

Plazas, M., I. Andújar, S. Vilanova, M. Hurtado,P. Gramazio, F. J. Herraiz, and J. Prohens.

MEYER ET AL.: MEDICINAL USES OF EGGPLANT IN ASIA2014]

Author's personal copy

2013. Breeding for chlorogenic acid content ineggplant: Interest and prospects. NotulaeBotanicae Horti Agrobotanici Cluj Napoca41:26–35.

Polignano, G., P. Uggenti, V. Bisignano, and C.Della Gatta. 2010. Genetic divergence analysisin eggplant (Solanum melongena L.) and alliedspecies. Genetic Resources and Crop Evolu-tion 57:171–181.

Quisumbing, E. 1951. Medicinal plants of thePhilippines. Department of Agriculture andNatural Resources, Manila, Philippines.

Raghunatha Ganesa Navahasta (17th cent.).Bhojanakutuhala of Raghunatha.

Raghunatha. 1956 (reprint of 17th Centuryoriginal). Bhojanakutuhala. Suranad KunjanPillai. Page 86. Kerala, India: UniversityManuscripts Library.

Ramalingayya, M. and M. V. Sastry. 1994(compiled by Vaidya Mulugu RamalingayyaGaru, translated into English by M.Vishveshwara Shastri). Vaidya Yoga Ratnavali(Formulary of Ayurvedic medicines). Madras,India: Indian Medical Practitioner’s Co-oper-ative Pharmacy and Stores, Ltd.

Rao, C. K. 2011. Use of brinjal (Solanummelongena L.) in alternative systems of medi-cine in India. FBAE, Bangalore, India.

Sastri, B. N. 1972. The wealth of India. Adictionary of Indian raw materials and indus-trial products. Raw materials. Vol. IX. Councilof Scientific and Industrial Research, NewDelhi, India.

Sękara, A. 2007. Cultivated eggplants—Origin,breeding objectives and genetic resources, areview. In: Folia Horticulturae 19, eds. A.Sękara, S. Cebula, and E. Kunicki, 97–114.

Sharma, R. K. and B. Dash. 1983 (translation).The Caraka Samhita, Vol. 1. Varanasi, India:Chowkhamba Sanskrit Series Office.

Southworth, F. C. 2005. Linguistic archaeologyof South Asia. Routledge Curzon, London.

Stommel, J. R. and B. D. Whitaker. 2003.Phenolic acid content and composition ofeggplant fruit in a germplasm core subset.Journal of the American Society of Horticul-tural Science 128:704–710.

Sushruta (approx. 600 B.C.E.). Susruta Samhita.Swarup, V. 1995. Genetic resources and breeding

of aubergine (Solanum melongena L.). ActaHorticulturae 412:71–79.

Tan, J. Eggplant. Hong Kong, China: JenniferTan Enterprises. http://tcmhealthtalk.com/eggplant (8 June 2012).

Thulaja, N. R. Brinjal. infopedia.nl.sg/articles/SIP_171_2004-12-15.html (8 June 2012).

Umali-Stuart , G. 2012a. Talong-pipit .www.stuartxchange.org/Talong-pipit.html (8June 2012).

——— 2012b.Talong-pipit. www.stuartxchange.org/Talong.html (8 June 2012).

Vardhana, R. 2006. Floristic plants of theworld, Vol. III. Sarup and Sons, NewDelhi, India.

Wang, J. X., T. G. Gao, and S. Knapp. 2008.Ancient Chinese literature reveals pathways ofeggplant domestication. Annals of Botany102:891–897.

Weber, S. A. 1991. Plants and Harappansubsistence: An example of stability andchange from Rojdi. Oxford and IBH, NewDelhi, India.

Weese, T. L. and L. Bohs. 2010. Eggplantorigins: Out of Africa, into the Orient. Taxon59:49–56.

Winstedt, R. O. 1944. Indian influence in theMalay world. Journal of the Royal AsiaticSociety (New Series) 76:186–196.

Wu, S.-B., R. S. Meyer, B. D. Whitaker, A.Litt, and E. J. Kennelly. 2013. A newliquid chromatography-mass spectrometry-based strategy to integrate chemistry, mor-phology, and evolution of eggplant (Sola-num) species. Journal of Chromatography A1314:154–72.

Wu, X.-Y (). 2006. (translated as Edible wildplants of Taiwan, Vol. 1; in Chinese). Taipei,Taiwan: .

Yang, S.-Z. and Y.-J. Gao. 2011. A preliminarystudy on diverse plant uses of Rukai tribe inWutai district of Pingting county, southernTaiwan. Taiwania 56:7–16.

Zhang, Q. 2009. Eggplant: Pretty, purple and perfectfor people. Shanghai, China: Shanghai DailyPublishing House. http://live.shanghaidaily.com/column_detail.asp?type=column&id=73 (8 June2012).

Zhao, X.-M. 1969 (reprint). Bencao gangmushiyi. (Materia Medica supplements). HongKong, China: Shang Wu Yin Shu Guan.

———. () (1719–1805). Written in 1765, firstprinted in 1864. (Bencao gangmu shiyi).

ECONOMIC BOTANY [VOL XX

Author's personal copy