全 文 ：热带作物学报 2011， 32（9）： 1786-1792
Chinese Journal of Tropical Crops
收稿日期： 2011-06-23 修回日期： 2011-08-03
基金项目： 公益性行业（农业）科研专项经费（No. 200903024）； 南亚热作专项（No. 09RZZY-06）； 海南省自然科学基金（No. 310071）。
作者简介： 范 睿 （1983年—）， 女 ， 研究实习员 ， 硕士 。 研究方向 ： 热带作物种质资源与遗传育种 。 *通讯作者 ： 邬华松 ， E-mail:
13807622912@163.com。
Research Progress on Molecular Phylogeny and
Genetic Diversity of Denus Piper L.
Fan Rui1,3,4, Huang Lifang1,3,4, Hao Chaoyun1,3,4, Yan Lin1,3,4, Yang Jianfeng1,3,4, Wu Huasong1,2,3,4*
1 Spice and Beverage Research Institute, Chinese Academy of Tropical Agricultural Sciences, Wanning, Hainan 571533, China
2 National Center of Important Tropical Crops Engineering and Technology Research, Haikou, Hainan 571101, China
3 Hainan Provincial Key Laboratory of Genetic Improvement and Quality Regulation for Tropical Spice and Beverage Crops,
Wanning, Hainan 571533, China
4 Key Laboratory of Genetic Resources Utilization of Spice and Beverage Crops, Ministry of Agriculture,
Wanning, Hainan 571533, China
Abstract Genus Piper, on e member of the Piperaceae, is located mainly in South and Central America and
South Asia. For a large number of species, the genus provides a noteworthy example of an increase in
diversification rate at the basal angiosperms. Meanwhile, the genus also includes the most valuable economically
important spice crops and various species are of ethnobotanical interest. Given the significant scientific and
economic value, many researchers have been dedicated to reveal the intrageneric molecular phylogeny, genetic
relationships and fingerprints of different cultivars, genetic distribution patterns of some endemic species, and so
on. To summary the past molecular researches, this paper systematically reviewed the researches on the molecular
phylogeny and genetic diversity of genus Piper. The results showed that most researches had focused on
intrageneric phylogeny of the genus, and a taxonomic framework had been established. Genetic diversity and
identification of the important cultivated Piper species had also been clarified. However, there are still some
problems remain to be solved: 1) Intrageneric phylogeny needs further refinement. 2) Genetic fingerprint bank
has not been established. 3) Genetic structures of some endemic and rare species have not received sufficient
attention.
Key words Phylogeny status; Intrageneric phylogeny; Genetic distribution pattern; Genetic diversity; Piper
doi 10.3969/j.issn.1000-2561.2011.09.042
胡椒属（Piper L.）分子系统学和
遗传多样性的研究进展
范 睿 1,3,4， 黄丽芳 1,3,4， 郝朝运 1,3,4， 闫 林 1,3,4， 杨建峰 1,3,4， 邬华松 1,2,3,4*
1 中国热带农业科学院香料饮料研究所， 海南万宁 571533
2 国家重要热带作物工程技术研究中心， 海南海口 571737
3 海南省热带香辛饮料作物遗传改良与品质调控重点实验室， 海南万宁 571533
4 农业部香辛饮料作物遗传资源利用重点实验室， 海南万宁 571533
摘 要 胡椒属（Piper L.）作为胡椒科（Piperaceae）中遍布于热带地区的重要作物， 含有近 2 000 个植物种类， 目
前集中分布于南亚、 南美洲和中美洲。 胡椒属内极高的物种多样性在传统的木兰亚纲（Magnoliidae）中显得尤为
独特， 为基部被子植物（Basal angiosperms）分化速率加快的机理研究提供了一个很好的例子； 同时， 属内还含有
众多具有重要经济价值的香料作物和民族植物种类， 如胡椒（Piper nigrum）和蒌叶（P. betle）等。 因其具有重要的
学术研究价值和经济利用价值， 许多研究者一直致力于揭示其属内分子系统关系、 栽培品种的遗传相似性和指
纹图谱、 特有物种的遗传分布特征等。 本文就过去几十年胡椒属的分子系统学和遗传多样性研究进行了系统总
结。 目前胡椒属在胡椒科中的系统地位已经确认； 胡椒属内的分子分类学框架已经基本建立； 一些重要栽培品
种的遗传多样性和鉴定也已明确。 但仍存在一些亟待解决的问题：（1）胡椒属内的分类系统仍显粗糙， 需进一步
细化研究； （2）胡椒种质资源的指纹图谱库尚未建立； （3）一些特有和珍稀种类的种群遗传特征研究未得到足够
重视。
关键词 系统地位； 属内分类系统； 遗传结构； 遗传多样性； 胡椒属
中图分类号 S573.9 文献标识码 A
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Fig.1 Simplified cladogram of perianthless Piperaceae[19].
Genus Piper L. belongs to Piperaceae, a family
of pantropical distribution concentrated mainly in
South and Central America and South Asia. The
genus includes nearly 2 000 species making it one
of the largest genera of the basal angiosperms [1-3].
The angiosperms are the largest, morphologically
and ecologically most diverse group of land plants,
and still an ‘abominable mystery’ with respect to
the circumstances of their origin and their rapid
diversification in the Early Cretaceous. In addition
to their relevance in applied fields of botany, basal
angiosperms are crucial for understanding the
history of flowering plants and for analyzing the
crucial factors in plant evolution. Piper is one of the
10 most speciose genera of basal angiosperms [4].
Because of a large number of species, the ‘giant’
genus is unique among the traditional Magnoliidae
and presents a great challenge, but also provides a
noteworthy example of an increase in diversification
rate at the basal angiosperms[5].
The genus includes the most valuable
economically important spice crops in which various
species are of ethnobotanical interest[6]. Geographically,
the Western-Ghats of South Indian peninsula is the
primary centre of origin of the King of spices, the
source of medicinally and commercially important
black pepper[7-8]. P. nigrum has been found in vast
altitudinal regions and shows great adaptability to a
wide range of environmental conditions[9]. The humid
climatic conditions and the daily consumption in
the diet make this spice crop cultivated widely in
the Asian continent. Betel vine (P. betle L.) is also
an important, traditional and ancient crop of India.
Leaves of betel vine have been used with
condiments such as arecanut, kattha, cloves,
cardamom, fennel and candied rose for chewing
purposes. The leaves have also been used in Indian
system of medicine and health[10-11]. In addition, P.
auritum, P. cubeba, P. longum, P. methysticum also
have some economic value.
Given the significant scientific and economic
value of genus Piper, many researchers have paid a
good amount of attention to this genus in the past
decades. Many of those researches are dedicated to
reveal the intrageneric molecular phylogeny of Piper,
genetic relationships and fingerprints of different
cultivars, genetic distribution patterns of some
endemic species, and so on. This paper
systematically reviewed the molecular phylogeny
and genetic diversity researches of the domestic
and foreign scholars, which would provide a
reference for related researches.
1 Phylogeny status of the genus Piper
The earliest classifications of Piperaceae
recognized between 7 and 15 genera[12-13], within the
current circumscription of the genus. De Candolle
continued the study of Piperaceae and lumped many
of the early genera into the large genus Piper, but
he preserved some of Miquels groupings at the
level of section or subgenus [14 -15] . De Candolles
classification was followed in the Asian floristic
treatments of the early 1900s[16]. The next monographer
of Piperaceae, William Trelease, abandoned the
infrageneric classification of de Candolle. Trelease
recognized a large and unstructured Piper along
with several small segregates in which some were
described by Milquel and others by himself.
Treleases generic concepts were followed by
Yuncker and largely accepted in the remainder of the
1900s. Callejas [17], who conducted the first cladistic
analysis of the genus using morphological characters,
reconsidered the infrageneric groupings within
Piper. Based on the results of Smiths research [18],
the members of Piper were found to be late
evolved among the selected genera. Using 6 000
basepairs of chloroplast DNA, Wanke recently
merged pacific genus Macropiper into Piper and
drew a tentative cladogram[19]. Only recently has it
become clear that Verhuellia is sister to the other four
genera in the family [20]. Based on the researches
mentioned above, the family consists of five genera:
Piper, Peperomia, Zippelia, Manekia, and Verhuellia,
which has been generally accepted (Figure 1) .
范 睿等： 胡椒属（Piper L.）分子系统学和遗传多样性的研究进展 1787- -
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2 In trageneric phylogeny of the
genus Piper
Piper species are distributed pantropically and
take the form of shrubs, herbs, and lianas common
in the understory of tropical forests around the
world. According to Gentry [21] , Piper reached its
highest diversity in the lowlands of the Neotropical
region ( 1 300 spp.). The second greatest diversity
occurred in the Southern Asia (300 spp.), followed
by the South Pacific (100 spp.). Although the Piper
species are easy to recognize by their nodose
shoots and perianthless flowers arranged in
condensed terminal spikes, the apparent uniformity
of their diminutive flowers and the vast number of
species have hindered the development of a stable
infrageneric classification[22].
The advances on the general biology,
revisionary studies and phylogenetic analyses
had paved the way for constructing a natural
classification for Piper [23 -25]. These studies,
based on exemplar sampling, confirmed the
monophyly of some traditionally recognized
infrageneric groups, while suggesting the
polyphyletic nature of other assemblages.
Based on a phylogenetic analysis of
sequences of the internal transcribed spacers
( ITS), Jaramillo et al. [23] recognized that the
genus Piper could potentially form three
monophyletic groups: the Neotropics clade,
Asia clade, and the South Pacific clade, a
hypothesis first proposed by Callejas , but
also somehow implicited in the de
Candolles key of the Piperaceae [15 ] . Within
the Neotropical clade, this research
recognized several clades that coincide with
infrageneric groups of de Candolle. Recently,
the phylogeny of Piper species had been
addressed through studies of floral
morphology in association with DNA
sequence data derived from rbcL, atpB, and
18S rDNA [26]. There were several well -
supported subclades, including Enckea,
Ottonia, Radula, Macrostachys, Pothomorphe,
and Macropiper. These c lades coincided with
segregates that had been recognized since the
earliest classifications of the genus [12 -13], and also
with clades described in earlier papers [23]. In the
latest research, Jaramillo et al. [27]sampled 575
accessions corresponding to 332 species of Piper
for the ITS region and 181 accessions for the
psbJ-petA chloroplast intron to further test previous
hypotheses about the major clades within Piper.
The combined analysis provided support for ten
monophyletic groups and offered the best hypothesis
for relationships among Piper. Piper was divided
into ten major clades for which a morphological
description was provided (Figure 2). However, some
details about this taxonomic system are still
unclear.
Figure 2 Schematic diagram summarizing results of the
ITS large dataset analysis. Numbers above branches are
Bayesian posterior probabilities and maximum parsimony
analyses. Boxes are proportional to sampling for each clade;
numbers within each box are accessions/species[26].
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第 9 期
3 Genetic dist ribution pattern in
different populations of Piper species
Patterns of distribution of Piper species vary
from being locally endemic to widespread. In the
lowlands of the Neotropical region, most species
exhibit restricted distributions and is rather common
to find numerous related endemic taxa occurring in
small size areas. This phenomenon also takes place
in the South and Southeast Asia. As others have
pointed out, the genetic structure of plant
populations reflects the interactions of many
different processes such as the long -term
evolutionary history, mutation, genetic drift, mating
system, gene flow, and selection[28]. For this reason,
an understanding of the extent and distribution of
genetic variation within and among natural
populations of endemic species is essential for
devising sampling strategies that efficiently capture
adequate genetic diversity for ex-situ conservation
purposes. Based on the allozyme analysis, intra -
populational genetic diversities of three Piper
species ( i.e. P. amalago, P. jacquemontianum and
P. pseudo -fuligineum) in Parque Nacional Santa
Rosa of northwestern Costa Rica were low relative
to other plants[29]. Genetic distances between species
were relatively high compared to other pairs of
congenetic plants. These high genetic distances
were consistent with the idea that this large genus
is relatively old. However, based on the result of
Shis research[30], genetic difference within P. nigrum
populations in China was relatively small, which
related that pepper was non -origin of China, few
introduced resources, limited geographical cultivation,
as well as the long -term adoption of cutting
propagation mainly. P. polysyphonum is an endemic
species in Southeast Asia, in the narrow habitat
located in the Chinese provinces of Guizhou and
Yunnan, and the country of Laos. Liao et al. [31]
developed 11 primer sets of polymorphic
microsatellite DNA loci for P. polysyphonum. Allele
numbers ranged from two to ten, with observed
heterozygosities ranging from 0.222 to 0.889.
However, researches in this regard are growing but
still insufficient. Compari ng to the ‘ giant’ genus,
the number of species is so small that it is
difficult to find the inherent regularity.
4 Genetic diversity and identification
of cultivated Piper species
4.1 Black pepper
P. nigrum is often named as Black pepper
for the color of its peppercorn, and is considered
as the ‘ king of spices’ due to its trade in the
international market [32 -33]. The fruits of P. nigrum
contain 1.0%~2.5% volatile oil, 5%~9% alkaloids,
of which the major ones are piperine, chavicine,
piperidine , and piperetine , and a resin [ 34 - 35 ] .
Therefore, its fruits have been widely used as an
important spice since time immemorial in household
spices as flavoring agents, and have also been used
in the treatment of cholera and dyspepsia, as well
as a variety of gastric ailments and arthritic
disorders [ 36 ] . Domestication of black pepper is
believed to have taken place many centuries ago
around its native habitat wherefrom its cultivation
spread to other countries of South and Southeast
Asia, and now it is chiefly cultivated in tropical
regions of the world such as India, Malaysia,
Indonesia, China, and Brazil, and on a smaller
scale in Sri Lanka and the West Indies as well[9].
Morphometrical studies done by various
workers have revealed the existence of extensive
phenotypic variability both in wild forms and in the
cultivated varieties[37-38]. Seven chromosome numbers,
2n =36, 48, 52, 60, 65, 78 and 128, have been
recorded in P. nigrum indicating the contribution of
cytological variation to its genetic diversity. The
majority of landraces were distinctly different from
advanced cultivars in the molecular profile
indicating artificial breeding efforts had brought
distinct genetic changes in the species [ 39 ] . Even
cultivars with the same name from different localities
did not always group together. AFLP analysis of 22
cultivars from South India showed that genetic
proximity among different cultivars could be related
to their phenotypic similarities or geographical
distribution. Greater divergence observed among
landraces than among cultivars could be related to
their ph enotypic similarities or geographical
distribution [ 38 ] . Modern pepper cultivars were
范 睿等： 胡椒属（Piper L.）分子系统学和遗传多样性的研究进展 1789- -
第 32 卷热 带 作 物 学 报
characterized by prominently higher yield than the
landraces [40]. Selection for yield contributing traits
might have contributed to the observed greater
genetic similarity among the advanced cultivars.
Mathew et al.[41] estimated 70 distinct cultivars of black
pepper in Kerala, and revealed that genetic resources
of the crop were getting lost for gene erosion.
One of the problems faced by pepper breeders
is the difficulty in identifying true hybrids from the
crossed progenies. Molecular markers are the most
direct answer to the problem. To date, little work
has been published in fingerprinting studies of
black pepper except for Randomly Amplified
Polymorphic DNA (RAPD) markers[42]. George et al.[43]
studied eleven black pepper accessions and their
hybrid populations maintained at the Indian
Institute of Spices Research using RAPD markers,
and the results showed that this technology was
useful in generating at least one band and to
select true hybrids. Sreedevi et al. [44] studied seven
promising cultivars of black pepper using RAPD
markers, and the results showed that random
primers could generate unique bands in 6 cultivars.
The seed of Carica papaya is less or more similar
to the seed of P. nigrum and easily adulterated in
some markets. RAPD markers had been employed
for authentication of P. nigrum from its adulterant
C. papaya, and this technology could clearly
distinguished genuine[45].
4.2 Kava
P. methysticum commonly known as Kava or
Kawa, which is wild or planted perennial medicinal
bush plants of Piper in the South Pacific islands.
As a phytomedicine, Kava is receiving considerable
worldwide interest in its use as a treatment for
anxiety, tension, agitation and insomnia[46]. The use
of Kava has a history of 3 000 years, which form a
unique ‘Kava culture’ in the local[47]. In the process
of introduction and cultivation of Kava, it was
difficult to distinguish among the seedlings of Kava
and its wild relatives which were similar in
morphology. In order to reveal the phylogeny status
of P. methysticum, Shi et al. [30] studied the genetic
relationships of 28 germplasms. The results showed
that Kava should belong to the South Pacific clade
which indicated that the level of genetic
relatedness appears to be high between Kava and
Asia clade. This research provided the basis for
selecting rootstocks molecular identification and the
fingerprint construction of P. methysticum.
4.3 Betel vine
Betel vine (P. betle) is an important, traditional
and ancient crop of India. Leaves of betel vine
have been used with condiments such as arecanut,
kattha, cloves, cardamom, fennel and candied rose
for chewing purposes. The leaves have also been
used in Indian system of medicine and health[10-11].
The betel vine growers invariably named their
cultivars with local or vernacular names. A given
landrace may be named differently in different
regions and more than one landrace may have the
same name. The different landraces were
distinguished earlier on the basis of the leaf
essential oils [10]. However, the extent of variation
among and between them was not easily analyzed
due to its vegetative propagation attributes. Ranade
et al. [48] earlier showed a clear distinction between
the ‘Kapoori’ and ‘Bangla’ landraces on the basis
of RAPD profiles. The landraces belonging to
‘Bangla’ group were more closely related to those
of the other two groups, namely, ‘ Sanchi’ and
‘Others’ than to the ‘Kapoori’ group. Verma et al.[49]
carried out RAPD analysis in several landraces and
considered four groups, namely, ‘Kapoori’, ‘Bangla’,
‘ Sanchi’ and ‘ Others’, in which the ‘ Kapoori’
group was the most diverse.
5 Prospects
(1)In the past decades, various kinds of
molecular technologies had been applied trying to
establish a taxonomic framework for the genus
Piper. Those researches, based on exemplar
sampling, confirmed the monophyly of some
traditionally recognized infrageneric groups and
established a taxonomic framework. However, some
aspects about this taxonomic system still need
explaining, and more attention should be paid to
this target.
(2)Although the genus Piper are easy to
1790- -
第 9 期
recognize in the field, the apparent uniformity of
their diminutive flowers and the vast number of
species in the genus have puzzled many local
farmers, even plant taxonomists. At the same time,
one of the problems faced by black pepper
breeders is the difficulty in identifying true hybrids
from the crossed progenies before planting.
Molecular markers are the most direct answer to
those problems. Currently, researches in this regard
are still insufficient, and the genetic fingerprint
bank for the Piper germplasm has not been
established.
(3)Assessment of genetic variation and its
partitioning within and among populations of
endemic species of genus Piper is necessary for
formulating conservation management strategies.
Whilst, only a few species had been studied, and
the efforts on this aspect should be strengthened.
(4)RAPD is commonly used for the genetic
diversity and distribution of Piper species and has
been proved to be a useful tool. However, this
technique is often criticized because of low level of
repeatability and spurious bands. Some new DNA
markers, such as simple sequence length
polymorphism ( SSLP) and sequence -related
amplified polymorphism ( SRAP), have the
advantages of rapidity, straight, and simplicity of
RAPD, and the stability, reliability, and
repeatability of RFLP. These new techniques will
be presented as a better alternative to RAPD for
the molecular researches of Piper in the future.
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