Ancient Chinese Literature Reveals Pathways of Eggplant Domestication
JIN-XIU WANG1,*, TIAN-GANG GAO1and SANDRA KNAPP2
1State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing
100093, China and2Department of Botany, Natural History Museum, Cromwell Road, London SW7 5BD, UK
Received: 13 May 2008 Returned for revision: 9 June 2008Accepted: 7 August 2008 Published electronically: 26 September 2008
†Background and Aims Changes in key traits occurring during the processes of plant domestication have long been
subjects of debate. Only in the case of genetic analysis or with extensive plant remains can specific sets of changes
be documented. Historical details of the plant domestication processes are rare and other evidence of morphological
change can be difficult to obtain, especially for those vegetables that lack a substantial body of archaeological data.
Botanical records chronicled in the ancient literature of established ancient civilizations, such as that of China, are
invaluable resources for the study and understanding of the process of plant domestication. Here, the considerable
body of ancient Chinese literature is used to explore the domestication process that has occurred with the eggplant
(Solanum melongena), an important vegetable in Old World.
†Methods Information about eggplant domestication in the ancient Chinese literature was retrieved using a variety
of methods. The information obtained was then sorted by taxon, examined and taxonomic identifications verified.
†Key Results It was found that the earliest record of the eggplant documented in ancient Chinese literature was in a
work from 59 BC. As far as is known, this is the earliest reliable and accurately dated record of eggplant in cultiva-
tion. The analysis reveals that the process of domestication of the eggplant in China involved three principal aspects
of fruit quality: size, shape and taste. These traits were actively and gradually selected; fruit size changed from small
to large, taste changed from not palatable to what was termed at the time sweetish, and that over time, a wider variety
of fruit shapes was cultivated.
†Conclusions The results indicate that, in addition to data gleaned from archaeology and genetics, evidence as to
changes in key traits occurring during the process of plant domestication and selective forces responsible for these
changes can be traced through the ancient literature in some civilizations.
Key words: Solanum melongena, ancient Chinese literature, domestication process, domestication traits, selective forces.
Changes in key traits occurring during the process of plant
domestication are mostly inferred by reference to wild rela-
tives or to primitive land races, especially for those crops
outside western Eurasia, and therefore are subject to con-
tinuous debate (Ladizinsky, 1998; Diamond, 2002). Only
in the case of genetic analysis (e.g. Doebley et al., 1997)
or with extensive plant remains (e.g. Zohary and Hopf,
2000) can specific sets of changes be documented.
Historical details of the plant domestication processes are
rare and other evidence of morphological change can be
difficult to obtain, especially for those vegetables that
lack a substantial body of archaeological data.
With potato (Solanum tuberosum) and tomato (Solanum
lycopersicum ¼ Lycopersicon esculentum), eggplant or
aubergine (Solanum melongena) is one of the three most
important cultivated crops in the Solanaceae (FAO, 2007).
The origin and domestication of eggplant, however, is
still a challenging question. Hypotheses about the origins
and evolution of eggplants have in the past been based on
inference (Lester and Hasan, 1991; Choudhury, 1995)
owing to the lack of archaeological evidence for origins
and early domestication. Lester and Hasan (1991) suggested
that the eggplant was derived from the subtropical species
S. incanum, native to north Africa and the Middle East.
They suggested that the wild progenitor developed as a
garden weed, and through human selection in south-east
Asia, progressively more advanced cultivars were selected.
They divided S. melongena into a series of morphological
types or gene pools, identified as A (putative wild progeni-
tors) to G (advanced cultivars), and suggested eastwards
movement of cultivated forms, with subsequent movement
westwards complicating patterns of character change
(Lester and Hasan, 1991). AFLP (Mace et al., 1999) and
DNA sequence (T. Weese and L. Bohs, University of
Utah, USA, pers. comm.) data sets support the broad
relationships between S. incanum and its African relatives
and the eggplants, but the data sets used were relatively
small and did not include Chinese samples. None of the
S. incanum s.s. group as currently understood is present in
China or adjacent south-east Asia; Lester and Hasan
(1991) hypothesized that the true wild progenitor of the
eggplant was an undiscovered species in the savanna eco-
systems of the region. No Asian prickly solanums have
been included in any phylogenetic analyses of relationships
in Solanum, making this a priority for understanding not
only evolution in Solanum in general, but the origin of
the eggplant in particular.
Several candidate areas for eggplant domestication have
been proposed: India and south-east China (Doganlar
et al., 2002a), China, India and Thailand (Doganlar et al.,
2002b), Burma to Indo-China (Daunay et al., 2001) and
south-east Asia (Lester and Hasan, 1991). Evidence for
each of these is based on presence of weedy forms (putative
* For correspondence. E-mail Heather@ibcas.ac.cn
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progenitors for many authors) and literature references. The
authors’ field work in recent years has revealed the presence
of wild, weedy forms of eggplants in southern China, sup-
porting a south-east Asian origin, but the possibility of mul-
tiple domestication events has not yet been investigated.
Evidence for an Indian domestication has been drawn
from examination of the Sanskrit literature. Khan (1979)
cited common names for the eggplant from various
works, with the oldest dated between the 3rd century BC
and the 3rd century AD. His citation of the oldest Sanskrit
work from 300 BC, however, was based on a secondary
source (Monier-Williams, 1899), and the time range he esti-
mated cannot be substantiated, due to the many revisions of
the work in question over the centuries (S. Y. Ye, Peking
University, China, pers. comm.). It is essential that the
primary sources of exact dates be re-examined in order to
explore this further. These Sanskrit names have been
regarded as evidence that the eggplant was first cultivated
in India, although no more detailed and continuous evi-
dence about the domestication process can be gleaned
from the Indian ancient literature. In contrast, in depth
examination of the ancient Chinese literature has revealed
a rich set of references to many cultivated plants (Li,
1969; Keng, 1974; Walters, 1989), including the eggplant.
The Chinese literature is especially characterized by its
antiquity, continuity and coherence (Needham, 1986).
Much written evidence about the eggplant can be found
in this vast reservoir. It is a treasure trove of information
about plant domestication (Brestschneider, 1871; Li,
1969) and has previously usually only been referenced
from secondary sources. The purpose of the present study
is to use primary sources to document as many of these his-
torical data as possible and, by piecing them together, to
draw a relatively authentic picture of the process of the
domestication and evolution of the eggplant in China.
MATERIALS AND METHODS
Details of the ancient Chinese literature
The size and scope of the ancient Chinese literature has been a
major obstacle to using it to study both uses of plants and the
domestication process. This literature is huge and complex
and includes books about botany, agricultural practice,
Chinese herbals, historical documents and relics, local chron-
icles, literary works, dictionaries and Leishu. Leishu are a
class of works combing to some extent the characteristics of
encyclopaedias and concordances, embracing the whole field
of literature, methodically arranged according to subjects,
each heading giving extracts from former works on the
subject in question, e.g. Gujin Tushu Jicheng (Imperial
Encyclopaedia) cited in this paper. The complete extent of
this botanical literature is still unclear even now. In an early
magnum opus on ancient Chinese botanical literature
(Brestschneider, 1882), for example, 1148 books were
included; nevertheless this treatment and description was rela-
tively superficial and contained mistakes. The general con-
dition and complexity of the ancient Chinese botanical
literature were discussed to some extent in another Western
compendium (Needham, 1986), but today, work is being
actively undertaken by Chinese botanists in this field.
Therefore, retrieving all the information in the huge ancient
Chinese literature is a great challenge to its use in research.
In the previous studies on eggplant (or aubergine, Solanum
melongena L.) involved in ancient Chinese literature
(Institute of Vegetables and Flowers of the Chinese Academy
of Agricultural Sciences, 2001; Daunay and Janick, 2007),
very few works were considered. The most often cited works
are Qimin Yaoshu and Nanfang Caomu Zhuang. Some
papers (e.g. Daunay and Janick, 2007) cite the Qimin Yaoshu
as two works using alternative spellings; this illustrates the
importance of consultation of the primary Chinese literature.
Information retrieval methods
Here, we attempted to retrieve the information about egg-
plant domestication from the ancient Chinese literature in a
variety of ways, and we are relatively certain that all import-
ant records were retrieved and analysed. Three methods
(1) Retrieval of information from important books on bota-
nical subjects that have been important and significant
in the development of Chinese ancient botany and
therefore partly studied by the scholars. Most of these
have been compiled and reprinted in modern times.
(2) Consultation of a wide variety of Leishu and Congshu
(encyclopaedias and collected books), such as Gujin
Tushu Jicheng (Imperial Encyclopaedia) and Siku
Quanshu (Complete Library of the Four Treasuries),
where many ancient books that have been lost
through historical time have been partly preserved;
these are not easily accessible at present.
(3) Consultation of electronic databases of non-botanical
ancient Chinese literature, for example, the database
for the poetry of the Tang Dynasty (http://chinese.
pku.edu.cn/tangPoem/). These databases are important
for special studies on plant domestication, although
most databases only present partial information and
re-analysis of the primary records is necessary.
Those records that repeated early statements or descrip-
tions and those not closely related to the questions of
changes during domestication that are being addressed are
excluded from the present analysis. In total, 189 records
from 76 books were included in the final analysis.
Information obtained was sorted by taxon, examined and
taxonomic identifications verified. The data presented here
are the most significant to trajectories of character
changes during domestication of the eggplant.
The earliest record and early dispersal in ancient China
The earliest written record of the eggplant documented in
Chinese ancient literature is the statement of Wang Bao in
Wang et al. — Eggplant Domestication892
his work Tong Yue in 59 BC (Wang, 59 BC). He states: ‘In
the second month of the year, the Spring Equinox ... sepa-
rate and transplant seedlings of eggplant and scallion’.
About the same time, Yang Xiong in his famous A
Rhapsody on Metropolitan of Shu (Yang, 1st century BC
to 1st century AD) mentioned: ‘The eggplant is included
as one of the vegetable crops’. Both of these records
referred to the eggplant that was then planted on the
Chengdu Plain of south-west China (Fig. 1). These
records, although rather brief, revealed that the eggplant
was already being specifically cultivated in gardens as a
crop, not a wild form collected from the surrounding
habitat. They also indicate that the domestication of the
eggplant took place not later than 59 BC in China. As far
as is known, this is the earliest record of eggplant, the
time of which is precisely determined.
Eggplant cultivation in southern China and northern
Vietnam is documented in Nanfang Caomu Zhuang
(Nan-fang ts’ao-mu chuang) (Ji, 304; Fig. 1). Thereafter,
from the 4th to the 6th century (Jin Dynasty and Northern
and Southern Dynasties), written records of the eggplant
referred mostly to the crop cultivated in the lower
Yangtze River regions (Shen, 5th and 6th centuries; Li,
659; Sima, 1084) (Fig. 1), and by the early 6th century,
eggplant was widely grown in the middle and lower
Yellow River regions (Jia, 6th century; Fig. 1). By the
Song Dynasty in the early 11th century, eggplant cultiva-
tion had spread all over China, and the crop had differen-
tiated into a number of different cultivars (Su, 1069).
Changes in key traits during the process of domestication
The process of domestication of the eggplant in China
involved three principal aspects of fruit quality: size,
shape and taste.
Size. The Qimin Yaoshu of the early 6th century AD (Jia, 6th
century) first described the size of eggplant fruit as ‘as large
as Danwan’. At that time a Danwan was a mud or an iron
pellet used in a slingshot and the term was often used to
turies (South Song Dynasty), the fruit was described as ‘as
large as Zhan’ (Wu, 12th and 13th centuries), a teacup of
the time of about 10 cm diameter. In AD 1069 (North Song
Dynasty) a drawing (Fig. 2A) of the eggplant appeared in
Tu Jing Bencao (Su, 1069); this is, as far as is known the
first drawing of the eggplant; the plant has no prickles on
the stems or leaves, and the fruits are round and apparently
much larger than a pellet. No more than 200 hundred years
later, a coloured drawing of the eggplant appeared in
6 century AD
4-6 century AD
4 century AD
80E85E 90E95E 100E105E110E 115E120E 125E130E 135E
75E80E85E 90E 95E100E 105E110E 115E120E 125E130E135E
FIG. 1. Distribution map of eggplant references in the early Chinese literature, before the crop’s spread across the entire region by the 6th century. Black
symbols indicate documented sites recorded in ancient Chinese literature.
Wang et al. — Eggplant Domestication 893
Lu ¨chanyan Bencao (Fig. 3) (Wang, 1220), a book produced
Province, eastern China. This painting shows that the fruit
were purple in colour and the ratio of fruit to the flowers
and leaves indicates that the size of the fruit of the eggplant
as ‘as round as Chinese trichosanthes’ (Trichosanthes kirilo-
wii Maxim., Cucurbitaceae, with fruits approx. 7–10 cm
in diameter) in the Bencao Gangmu. During the Qing
Dynasty (17th to 19th centuries) numerous cultivars were
recorded in local chronicles. In the Chronicles of Jiangxian
(Shanxi Province) (Chen, 1726), for example, a kind of egg-
plant is described with a fruit ‘as large as Yu, and the weight
kind of wide-mouthed receptacle for holding tea or water,
and according to the weight system used during the Qing
Dynasty, two jin and ten liang equals 1549 grams. This is a
very large eggplant fruit even by modern standards. All of
these reports indicate that ancient Chinese farmers were
continuously selecting eggplants with larger fruits.
Shape. Eggplants recorded in early Chinese literature were
mostly round-shaped (Jia, 6th century; Su, 1069; Wu,
12th and 13th centuries; Wang, 1220). The first recorded
long fruit of the eggplant appeared in the 14th century
(Yuan Dynasty) work entitled Wang Zhen Nongshu: “a
cultivated ‘Shui Qie (
)’ with long fruit and a slightly
sweet taste, could quench one’s thirst.” (Wang, 14th
century). The Bencao Gangmu (Li, 1590) of the Ming
Dynasty recorded eggplants with fruits ‘as long as four to
five cun (
; approx. 12.8–14 cm in the length system of
the Ming Dynasty)’ (Fig. 2B). In the 17th century (Qing
Dynasty) a number of cultivars with long rod-shaped fruit
named ‘Chansi Qie (
1726). The name refers to the length of the fruit; ‘Qie
)’ is the part of the name indicating an eggplant
variety, while ‘Chansi (
fruit. Today there are still long fruit eggplant cultivars
like ‘Chansi Qie’ grown in East China; ‘Ningbo Teng
Qie’, for example, has a length of 30–40 cm with a dia-
meter of only 1.7–2.0 cm (Institute of Vegetables and
Flowers of theChinese
Sciences, 2001). At about the same time, an egg-shaped
eggplant was drawn in Sancai Tu Hui (Wang and Wang,
1609; Fig. 2C). By the late Qing Dynasty numerous culti-
vars with different shapes were recorded in the many
local chronicles published at that time (Chen, 1726).
These varieties were in large part similar to those grown
today by Chinese farmers, and comprised round types, egg-
shaped types and long, thin types. The most common
)’ were recorded (Chen,
)’ described the long, thin
FIG. 2. Drawings of eggplant from selected works of the ancient Chinese literature: (A) Tu Jing Bencao in AD 1069 (North Song Dynasty); (B) Bencao
Gangmu in AD 1590 (Ming Dynasty); (C) Sancai Tu Hui in AD 1609 (Ming Dynasty); (D) Zhiwu Ming Shi Tu Kao in AD 1848 (Qing Dynasty). Reproduced
by permission of the Chinese Academy of Sciences.
FIG. 3. Coloured drawing of eggplant from Lu ¨chanyan Bencao in AD 1220
(South Song Dynasty). Reproduced by permission of the Chinese Academy
Wang et al. — Eggplant Domestication894
cultivars were pictured in the famous botanical magnum
opus Zhiwu Ming Shi Tu Kao (Wu, 1848; Fig. 2D). It
seems that more different shapes of eggplant fruit were
selected and preserved as the domestication process
Taste. In the early 6th century AD the taste of eggplant fruit
was described as ‘like the flavour of small beans’ in the
Qimin Yaoshu (Jia, 6th century), and according to other
records of that time (Li, 659), the eggplant was not con-
sidered a very palatable vegetable. Later, however, in
Youyang Zazu (Duan, 9th century), a book written in the
Tang Dynasty, it was chronicled that ‘Buddhist monks
roasted and ate eggplants, the taste was very delicious’. In
the 12th and 13th centuries (South Song Dynasty), some
books on agriculture (e.g. Wu, 12th and 13th centuries)
recorded the method of using sulfur to make the fruit
flesh sweeter (less bitter). Huang Ting-Jian, a poet of the
South Song Dynasty even wrote several poems in praise
of silver-coloured (white) eggplant cultivars with a deli-
cious taste that could be eaten raw (Huang, 12th and 13th
centuries). All of these references show that the Chinese
farmers selected for improved taste in a variety of ways,
but particularly for the elimination of bitter flavour in the
Some selective forces during the domestication process
As eggplant cultivation spread across China by the 6th
century AD, many advanced techniques for eggplant were
developed by Chinese farmers. Three main kinds of cultiva-
tion methods were adopted in the 6th century (Jia, 6th
century), including ‘planting in beds (rectangular pieces
of land in a field, separated by ridges) in the second
month’, ‘planting in the 10th month’ and ‘planting in
spring but not in beds’. Water selection of good seeds for
reproduction based on the difference in density between
plump and shrivelled seeds was practiced (Jia, 6th
century). A complex fertilization method involving the
use of ashes from old leaves as mulch called ‘Jia Qiezi
)’ was also developed in the 9th century
(Tang Dynasty) to improve fruit yield (Duan, 9th
century). In the 14th century (Yuan Dynasty), a technique
for growing seedlings in a temperature-controlled room in
winter for early harvest was developed (Lu, 1330).
Diversity of the eggplant in China
Both the intensive and exuberant activities of ancient
Chinese farmers in the domestication of the eggplant, and
the vast territory of China are clearly factors resulting in
the large number of modern eggplant cultivars in China.
Today, more than 200 native eggplant cultivars of fine
quality are formally registered (Institute of Vegetables
and Flowers of the Chinese Academy of Agricultural
Sciences, 2001). The vast expanse of China has also led
to many vernacular names for the eggplant and its varieties.
The following names can be found in ancient Chinese lit-
)’, ‘Zigua ()’, ‘Aigua (
)’, ‘Kunwei( )’,
)’ and ‘Xiaogu (
recorded in the 7th century, is still used in vernacular
dialect of the Shanghai and Zhejiang areas for eggplants
with long, thin fruits.
)’. ‘Luosu (
From the historical evidence we have collected from the
ancient Chinese literature, a vivid picture of the process
of the domestication and evolution of the eggplant in the
region has emerged. These records reveal that the cultiva-
tion of the eggplant took place in China not later than the
1st century BC; that the fruit size of the eggplant was
actively and gradually selected and changed from small to
large; that the taste changed from not palatable to what
was termed at the time sweetish and that, over time, more
kinds of shape of the eggplant were selected and preserved
Some of these domestication traits (fruit size and shape)
have been shown to be controlled by a very limited number
of target loci. Using QTL analyses, Doglanar et al. (2002b)
showed that fruit size of the eggplant was controlled pri-
marily by only two loci (fw2.1 and fw9.1) on linkage
groups 2 and 9, and diversification in fruit shape primarily
controlled by two loci (fs2.1 and fs7.1) on linkage groups 2
and 7. The two other domestication traits investigated –
prickliness and fruit þ plant colour – were shown to be
determined primarily by a single locus on linkage group 6
(lp6.1), and a major locus on linkage group 10 (fap10.1,
Documentation of these latter two domestication traits in
the ancient Chinese literature, however, is too incomplete
to infer any evolutionary trends. This may reflect to some
extent the lower importance given to prickliness and
colour (of both fruit and plant) than to fruit size and
shape during selection of cultivars in ancient China.
Further research is needed to clarify this. Taste of the
eggplant, another important domestication trait, however,
was not examined in Doganlar et al.’s QTL analyses
TABLE 1. Changes during the domestication of the eggplant recorded in ancient Chinese literature
–1 to 15–68–912–1314–1618–19
As large as Danwan
As large as Zhan
Sweetish and delicious
Round, oval, long
As large as Yu, weight up to 1549 g
Round, ovoid, long, and other shapes
Sweeter and more delicious
Wang et al. — Eggplant Domestication895
(Doganlar, 2002b). Our investigations in southern China
confirmed that fruit taste of weedy forms and the primitive
cultivars of eggplant are usually unpalatable and more bitter
than that of more advanced cultivars with larger fruit. Fruit
bitterness in eggplant is caused by two kinds of steroidic
saponosides (Aubert et al., 1989). It will be interesting
and challenging to explore the genetics of taste modifi-
cation from bitter to less bitter (sweet) taste that have
occurred during the domestication of eggplant.
The processes of artificial selection on vegetables are
usually slow and gradual (Ladizinsky, 1998). The specific
stages of eggplant domestication described in the ancient
Chinese literature are not unambiguously delineated. But,
as the Chinese history and civilization is relatively continu-
ous and independent, the stages described may have been
an extension of early domestication in the region. It is
amazing that many details of our findings based on histori-
cal records support the suppositions about the domesti-
cation process inferred by other authors (Choudhury,
1995; Lester and Hasan, 1991; Daunay et al., 2001).
Furthermore, from this detailed ancient literature, we now
have insights into some selective forces by which Chinese
eggplant farmers brought about these changes through agri-
cultural practice. The present data provide the earliest direct
evidence for the domestication process of the eggplant
occurring in Asia, and represent a novel approach to under-
standing the process of plant domestication. These indicate
that, in addition to data gleaned from archaeology and gene-
tics (Zeder et al., 2006), evidence as to changes in key traits
occurring during the process of plant domestication and
selective forces responsible for these changes can be
traced through the ancient literature in some civilizations.
We thank Zhengyi Wu, Xingguo Han, Yancheng Tang,
David Boufford, Zuoming Yang, Qingbing Cheng, Dewei
Zhu, Jin Pan and Song Ge for critical discussions and com-
ments on an early draft, and Yan Tang and Shaoyong Ye for
data collection. This study was funded by the United States
National Science Foundation’s Planetary Biodiversity
Inventory program ‘PBI Solanum: a world treatment’
(DEB-0316614 to S.K.), National Science Foundation of
China (30770159 to J.-X.W) and the National Basic
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