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薏苡生物学研究进展(英文)



全 文 :Research Progress of Biology in Coix
Guoqing LIU*,Wei LI,Haiquan LI,Jianan ZHANG,Jinying XIANG,Yucui HAN,Lingling GENG,
Shenglin HOU
Minor Cereal Crops Laboratory of Hebei Province, National Foxtail Millet Improvement Center, Institute of Millet Crops of Hebei
Academy of Agriculture and Forestry Sciences, Shijiazhuang 050035, China
Supported by the Special Financial Program of Hebei Province, the Fund for Agricultural
Science and Technology Innovation Talent Team Construction (2011055001).
*Corresponding author. E-mail: guoqingliu@hotmail.com
Received: September 30, 2015 Accepted: January 28, 2016A
Agricultural Science & Technology, 2016, 18(3): 494-498, 554
Copyright訫 2016, Information Institute of HAAS. All rights reserved Molecular Biology and Tissue Culture
C oix is an ancient food and ec-onomic crop, serving as amedicinal material and a kind
of greenfeed. Coix has a cultivation
history of 2 500 years in China[1], and is
widely distributed in all parts of China,
and China has been the greatest Coix
production country and export country
in the world currently. In recent years,
with continuous exploration of the ef-
fect of Coix in resisting cancer, the ec-
onomic value of Coix has been recog-
nized by people gradually. Coix has a
nearer genetic relationship with Zea. It
has many good agronomic traits, such
as pest resistance, drought tolerance,
lodging tolerance, saline-alkaline tol-
erance and poor soil tolerance, and is
of great significance to the species im-
provement of graminaceous crops
such as maize and sorghum. Howev-
er, there are many problems exiting in
the research and production of Coix,
such as weak fundamental research,
unclear evolutionary process including
origin, species formation and differen-
tiation; not enough attention to collec-
tion, management and utilization of
germplasm resources, and damage
and disappear of partial wild resources
due to factors such as environment
pollution and land development; and
chaos of species, and low yield, easy
lodging, late maturity and poor pest
and disease resistance in exiting
species. This study reviewed the re-
search status of Coix, aiming at provid-
ing information for research and pro-
duction of Coix.
Classification of Coix
Coix belonging to Trib. Maydeae
Dumort. of Panicoideae in Gramineae,
is an annual or perennial C4 herba-
ceous plant. It was named by Carl Lin-
naeus in 1753. The genus includes 9-
Abstract Coix has a long cultivation history as a minor crop in China, but many
problems including unclear origin, chaos of species, loss and inaccurate use of
germplasm resources exist in its research and production due to the lack of sys-
tematic study. In recent years, the importance of Coix has been realized with the
exploration of its effects in treating and preventing cancer. The systematic study of
Coix has met a superb development opportunity with the rapid development of
biotechnology especially the deep-sequencing technique. Therefore, this paper pre-
sented here the current status of origin and evolution, classification, cytology and
molecular biology progress of Coix, analyzed the ploidy of Coix and the genetic re-
lationships with Zea, Sorghum and so on from the point of cytology, and then
based on the molecular biology, analyzed the important significance of DNA molec-
ular marker, genetic mapping and gene library in the research on identification of
genetic relationship between species and genetic breeding, which could provide use-
ful information for further Coix research.
Key words Coix; Classification; Cytology; Molecular biology; Research progress
薏苡生物学研究进展
刘国庆*,李 伟,李海权,张嘉楠,相金英,韩玉
翠,耿玲玲,侯升林 (河北省农林科学院谷子
研究所,国家谷子改良中心,河北省杂粮研究
实验室,河北石家庄 050035)
摘 要 薏苡在我国一直作为小杂粮种植,尽
管栽培历史悠久,但缺少系统性的研究,存在
起源不清、品种杂乱、种质资源丢失和利用效
率低等问题。 随着薏苡保健功能和癌症防治功
效研究的不断挖掘与深入,以及分子生物学技
术尤其是深度测序技术的普及和成本的下降,
系统性研究薏苡遇到了绝好的发展契机。 对薏
苡的起源与分类、 营养和药用价值进行了概
述,从细胞学方面分析了薏苡倍性以及与玉蜀
黍族玉米属、高粱属等的亲缘关系,从分子生
物学方面分析了 DNA 分子标记、 遗传图谱和
基因文库在鉴别物种亲缘关系以及遗传育种
等研究中的重要意义,以期为薏苡的深入研究
提供理论参考。
关键词 薏苡;分类;细胞学;分子生物学;研
究进展
基金项目 河北省财政专项农业科技创新人
才队伍建设基金(2011055001)。
作者简介 刘国庆(1964-),男,河北沧州人 ,
研究员,博士,主要从事植物分子遗传育种研
究。 E-mail:guoqingliu@hotmail.com。 *通讯作
者。
收稿日期 2015-09-30
修回日期 2016-01-28
DOI:10.16175/j.cnki.1009-4229.2016.03.005
Agricultural Science & Technology2016
11 species of different ploidies (2n=10,
12, 20, 30, 32, 40). There are still a lot
of viewpoints and arguments on the
classification of Coix currently, Qiao et
al.[2] deemed that Coix could be classi-
fied to 1 species and 1 variety in Chi-
na, Zhuang et al. [3] reckoned that Coix
could be classified into 3 species and
4 varieties in China, Lu et al. [4] consid-
ered that there were 4 species and 8
varieties in Guangxi, and Li et al. [5]
classified Coix resources in Guangxi
into 4 species and 9 varieties.
It was recorded in “flora of China”
that there were 5 species and 2 vari-
eties of Coix in China, i.e. Coix aquat-
ica Roxb., Coix puellarum Balansa,
Coix stenocarpa Balansa, Coix lacry-
ma-jobi Linn. and its variety Coix lacry-
ma-jobi var.maxima Makino, and Coix
chinensis Tod. var. chinensis and its
variety Coix chinensis var. formosana
(Ohwi) L. Liu. Fifty two Coix species
and varieties were recorded in the re-
vised “World Checklist of Selected
Plant Families”, in which Coix aquati-
ca Roxb., Coix puellarum Balansa,
Coix stenocarpa Balansa, Coix lacry-
ma-jobi Linn. and its variety Coix lacry-
ma-jobi var.maxima Makino, Coix
lacryma-jobi L. var. lacryma-jobi , Coix
lacryma-jobi var. ma-yuen (Rom.
Caill.) Stapf and Coix gasteenii B. K.
Simon, while two varieties, Coix chi-
nensis var. formosana (Ohwi) L. Liu.
and Coix gigantea Koenig ex Roxb.
have not been recorded officially (http:
//apps.kew.org/wcsp/).
Origination and Distribu-
tion of Coix
Coxi is originated from Asia[7], and
mainly distributed in Myanmar, India
and China. China is one the important
origins of Coix. Coix is planted in vari-
ous provinces (regions) in China ex-
cept Qinghai, Gansu and Ningxia [2],
and the provinces including Guangxi,
Guizhou, Yunnan, Zhejiang and Hebei
show higher yield at present[8-9]. Huang
et al.[10] divided China into 3 Coix diver-
sity centers, i.e. south, the middle and
lower reaches of Changjiang River and
north centers, among them, Guangxi,
Hainan, Guizhou and Yunnan are the
primary center of Coix in China, the
middle and lower reaches of
Changjiang River and various northern
provinces and regions are secondary
centers formed in gradual northward
movement, domestication and selec-
tion of Coix lacryma-jobi and Coix
lacryma-jobi var. ma-yuen (Rom.
Caill.) Stapf. Coix germplasm re-
sources in China are abundant due to
the differences and changes in geo-
graphical conditions, climates and cul-
tivation conditions of the different di-
versity centers, and 3 ecoregions, i.e.
the late-maturing south ecoregion (in-
cluding Hainan, Guangdong, Guangxi,
Fujian, Taiwan, Yunnan, Hunan, south
of Sichuan and Xizang Autonomous
Region), mid-maturing middle and
lower Changjiang River ecoregion (in-
cluding Jiangsu, Zhejiang, Anhui,
Jiangxi, Sichuan, Hubei, southern
Shaanxi and northern Hunan) and
early-maturing north ecoregion (in-
cluding Beijing, Hebei, Shandong,
Henan, Shanxi, Liaoning, Jilin, Hei-
longjiang, Inner Mongolia and Xin-
jiang) have been formed. Coix collect-
ed from 12 populations in China was
subjected to clustering analysis using
13 biological traits, and the 12 popula-
tions were divided into 4 groups: the
first group including Anguo of Hebei,
Chengde in Hebei, Chengdou in
Sichuan, Taiyuan in Shanxi and
Tongcheng in Anhui; the second group
including Nanjing in Jiangsu and
Pucheng mountains in Fujiang; the
third group including Suining in Jiang-
su, Shenyang in Liaoning and
Shuyang in Jiangsu; and the fourth
group including Pucheng Coix aquati-
ca in Fujiang and Xianju in Zhejiang[9].
Nutritional Components in
Coix
Coix has high and comprehensive
nutritional value, with a good reputa-
tion of “the king of world gramineous
plants”. Li et al. [11] performed quality
analysis and determination to Coix
samples collected from Yi County in
Liaoning, etc., and found that Coix
showed a total fatty acid content of
about 7% and a protein content of
about 14%, and an isoleucine content
of 0.62%, a leucine content of 2.24%,
a lysine content of 0.31% , a pheny-
lalanine content of 0.81%, a threonine
content of 0.42%, a proline content of
1.07% , a valine content of 0.84% , a
glutamic acid content of 3.76% , an
oleic acid content of 51.1%, a linoleic
acid content of 33.6%, and a linolenic
acid content of 0.55% . In addition,
Coix contains energy 1 597 kJ, water
12.2 g, protein 14.4 g, fat 3.8 g, dietary
fiber 1.2 g, carbohydrates 66.6 g, vita-
min B1 0.21 mg, vitamin B2 0.16 mg,
vitamin E 0.3 mg, K 311 mg, Ca 51
mg, Mg 127 mg, Fe 3.9 mg, Mn 1.89
mg, Zn 1.89 mg, Cu 0.07 mg and P
299 mg per 100 g. Compared with rice,
the contents of crude protein and
crude fat in Coix are 1.97 and 5.81
times of those in rice, the amino acids
provided by unit mass is 2.2 times of
that by rice, the content of vitamin B1
is 2.54 times of rice, and the microele-
ments are higher than rice by 25% [12].
The non-kernel parts of Coix contain
16 amino acids, including the 7 amino
acids (lysine, phenylalanine, leucine,
isoleucine, valine, methionine and
threonine) essential for human body.
Among the non-kernel parts, the en-
dopleura contains the highest total
content of amino acids and highest to-
tal content of the 7 essential amino
acids, which are 113.42 -139.81 g/kg
and 35.53 -42.60 g/kg, respectively;
and the root and testa are the second,
the total content of aminoacids and total
content of the 7 essential amino acids in
the rootare42.63-67.53g/kgand13.73-
16.83g/kg, respectively, and those in the
testa are 15.73 -68.51 g/kg and 5.06 -
17.51g/kg, respectively[13].
Medicinal Value of Coix
Coix has been widely used as a
medicinal crop long before in China,
for instance, ripe kernels of Coix is one
of traditional Chinese medicinal mate-
rials. Coix was recorded in the first
section of “Chinese Pharmacopoeia”
in 2000, because it has the effects of
tonifying spleen, eliminating damp-
ness, removing paralysis, curing diar-
rhea, and clearing heat, and is used
for treating edema, difficult urination,
diarrhea caused by insufficiency of the
spleen, pulmonary abscess, intestinal
carbuncle, plane warts, etc. Further-
more, it is found by modern medical re-
search that Coix has pharmacological
activity in the aspects of resisting tu-
mor, adjusting immune, regulating
blood sugar, reducing blood pressure
495
Agricultural Science & Technology 2016
and resisting virus, for instance, Coix
seeds are used for treating chapped
skin, rheumatism and neuralgia, and
serve as anti-inflammatory and an-
thelmintics components [14]. Partial stu-
dies also found thatCoix had the effects
of resisting proliferation of nasopharyn-
geal carcinoma cell [15] and Hep-2 cell,
resisting lung cancer and rectal can-
cer[17 -18], inhibiting early-phase rectum
canceration[19] and resisting anaphylax-
is [14], and could promote the health
of angiocarpy and intestinal tract [20].
Coixenolide has been applied clinically
currently, and could achieve a good
effect on advanced nasopharyngeal
cancer in combination with radiothera-
py[21]. In addition, Coix also has the ef-
fects of reducing blood lipid and resist-
ing oxidation[22-23]. Coix seeds could be
used for treating obesity and serve
as functional food [24], the benzoxazi-
none type ligands contained therein has
an anti-inflammatory effect[25-26], and the
extract of Coix seeds has an antiviral
effect[27].
Cytological Study of Coix
Coix is rich in ploidy, and common
ones are diploid (2n =10), tetraploid
(2n=20), sextuploid (2n=30) and octa-
ploid (2n=40), with a basic number of
chromosomes of 5[28], and the most co-
mmon one is tetraploid, such as Coix
lacryma-jobi L. Except above
genomes, Coix also includes materials
with other ploidies, such as Coix gi-
gantea (2n=12 and 2n=40)[29] and Coix
taxon (2n=32)[30].
The original species of Coix is
Coix aquatic Roxb., which was found
in India firstly, and contains diploid
genome (2n=10)[31]. This species also
exhibits other genomes, including te-
traploid (2n=20) [32] and octaploid (2n=
40)[33]. It has been found in China lately
that there is a kind of perennial Coix
aquatic Roxb. (2n=30), which is highly
male-sterile and relies on vegetative
propagation[4]. 45S and 5S rDNA were
used as probes for hybridization in
situ, and the species was speculated
to be a new kind of sextuploid Coix
aquatic Roxb., whose parents might
be the tetraploid Coix lacryma-jobi L.
and octaploid Coix aquatica (2n=40)[34].
Genomic in-situ hybridization could be
performed with the genomic DNA of
tetraploid Coix lacryma-jobi as a probe
and the metaphase chromosome in
somatic cells of Coix aquatic, and it
was found that the DNA contents of 20
chromosomes were highly homolo-
gous with the genome DNA of Coix
lacryma-jobi[35]. It was found by prelim-
inary genomic sequencing comparison
on Coix lacryma -jobi and Coix aquati-
ca HG (2n=30) that the sextuploid Coix
aquatica was probably a lately-formed
species, and Coix lacryma -jobi suf-
fered diploidization in its evolution pro-
cess [36]. The research on the genetic
relationship between Coix with other
Poaceae species especially maize is
an important direction of Coix re-
search. Coix and Zea both fall into
Maydeae, and among the 7 genera in
the Maydeae, Coxi has the nearest ge-
netic relationship with Zea besides
Tripsacum. Cai et al.[36] performed ran-
dom sequencing on the genomes of 2
Coix species, found by comparison of
Coix with the genomic sequences of
sorghum and maize that the repetitive
sequences of Coix had higher homol-
ogy with those of sorghum, and thus
deemed that Coix had a closer genetic
relationship with sorghum than maize.
It was indicated by genomic in-situ
hybridization with DNAs of wild rela-
tives of maize as probes that Zea had
a closer genetic relationship with Trip-
sacum than that with Coix, while maize
had a closer genetic relationship with
Coix than with sorghum[37]. The cross-
ing barrier between Tripsacum and
maize is embryo sac incompatibility or
hybrid break-down rather than cross
incompatibility; the hybridization of
maize with Coix is stricter in reproduc-
tive isolation than with Tripsacum, and
is very difficult due to embryo sac in-
compatibility and style incompatibility,
and the abnormal rate of pollen tube
observed from the hybridization of
maize and Coix is higher than that ob-
served from the hybridization of maize
and Tripsacum. Therefore, maize has
a closer genetic relationship with Trip-
sacum than with Coix[38].
Molecular Biology Study of
Coix
The molecular biology study of
Coix has just started, the DNA molec-
ular marker systems for its molecular
biology study have been developed
gradually, and these markers could be
widely applied to the studies on genet-
ic breeding, germplasm diversity de-
tection, identification of genetic rela-
tionship between species, species
evolution analysis, genome mapping,
gene localization, genebank establish-
ment and gene clone. The molecular
marker methods used in the research
of Coix include RFLP (restricted frag-
ment length polymorphism), RAPD
(random amplification of polymorphic
DNA), AFLP (amplified fragment
length polymorphism), SRAP (se-
quence-related amplified polymor-
phism), simple sequence repeats
(SSRs) and so on [39-42]. RFLP requires
molecular hybridization, AFLP and
RAPD are dominant marker methods
though they are both performed on
the basis of PCR, and therefore, MA
et al. [43] selected 17 convenient prac-
tical SSR markers from an estab-
lished gene library with abundant mi-
crosatellite sequences by improved
biotin-streptavidin capture. Studies
on genetic diversity, fingerprint, sys-
tem evolution and establishment of
genetic linkage map of Coix
germplasms from Korea, Vietnam
and Yunnan and Guangxi of China
were conducted with these markers,
conventional STS marker, and SSR
markers from maize and rice [40-42, 44-46].
Genetic map is a basic tool for genetic
research, gene localization and sepa-
ration and marker-assisted breeding.
Qin et al.[46] constructed a genetic link-
age map of Coix using an F2 popula-
tion with 131 individuals obtained from
parents Beijing and Wuhan Coix
lachryma-jobi L. (2n =20) with AFLP
and RFLP markers, and formed 10 ge-
netic linkage groups corresponding to
the chromosome number with 80 AFLP
markers and 10 RFLP markers, with a
total length of 1 339.5 cM and an aver-
age genetic distance between markers
of 14.88 cM; and among the linkage
groups, the longest linkage group had
a length of 379.9 cM, containing 24
markers, and the shortest one had a
length of 30.9 cM, containing 4 AFLP
markers. Li [47] performed linkage anal-
ysis on the segregation population ob-
tained from the combination of Beijing
and Wuhan Coix lachryma-jobi L. and
divided all used molecular markers in-
496
Agricultural Science & Technology2016
to 11 genetic linkage groups with a to-
tal length of 1 275.7 cM and an aver-
age genetic distance between markers
of 15.6 cM; and QTL analysis was also
performed on important Coix agronom-
ic traits, 31 QTL loci were localized,
the related traits included stem diame-
ters (8), plant heights (3), tassel
lengths (6), leaf lengths (7) and leaf
width (7), and the phenotypic variance
explained by each QTL ranged from
12.2% to 64.4% . The construction of
genetic maps lays the foundation of
exploration of beneficial genetic loci in
Coix resources.
Meng et al. [48] established the first
bacterial artificial chromosome (BAC)
library using PI 324059 (Coix lacryma-
jobi L.) in 2010, and the BAC library
contains about 230 400 clones with an
average insert size of 113 kb, and pro-
vides 16.3-fold coverage of the
genome, providing a material basis for
future gene isolation and comparative
genome research. The entire se-
quence of the coding region of waxy
gene of Coix lacryma-jobi has been
isolated successfully currently, and it
was found that there was a 275-bp
deletion in the coding region of this
gene specific to waxy cultivars, and
also shown that this deletion was
commonly found in Japanese and
Korean cultivars [49]. Furthermore, Cai
et al.[36] the genome sizes of Coix lacry-
ma-jobi (2n=20)and Coix aquatic HG
(2n =30), which were 1.684 Gb and
2.335 Gb, respectively, and found that
the later one was 1.4 times of the for-
mer one; and it was found by 1.84 ×
and 2.76×sequencing on the 2 materi-
als that 75.54% of the genome of Coix
lacryma-jobi and 72.88% of the
genome of Coix aquatica HG were
repetitive DNA sequences.
Prospect
The inflorescence of Coix is
raceme formed by separate flower
and female and male spikelets. The
male spikelets are higher than the fe-
male spikelets which are late-matur-
ing, and when the male spikelets be-
gin to spread pollen, the filaments
have already withered, thereby avoid-
ing pollination within the same inflo-
rescence. In the growth process of
Coix, hybridization could be per-
formed easily under the condition of
near flowering phases, resulting in hy-
brid seeds, and such pollination man-
ner greatly increases the genetic di-
versity of Coix resources [9]. Coix was
not a main food and economic crop
before, and its basic research and
production application research has
long been neglected; China is abun-
dant in Coix resources, while there are
only 284 germplams stored and regis-
tered in the National Genebank [8]; and
the native area of Coix is damaged in-
creasingly, for instance, the native
area of wild Coix in Guangxi has been
damaged by 70% since the 1980s [50].
Therefore, the collection and expand-
ing propagation, management and i-
dentification, and protective conserva-
tion of Coix resources have been very
necessary and urgent.
Coix is closer to Sorghum and
Zea, they have many similar traits,
while the resistance and adaptability of
Coix are both better than crops in-
cluding maize and sorghum, and
therefore, Coix is a new genebank for
improving cereal crops including maize
and sorghum. Meanwhile, with the
completion of genomic sequencing of
maize and sorghum having closer ge-
netic relationships with Coix, these re-
search achievements also create op-
portunities for deep research of Coix.
The random sequencing of Coix lacry-
ma-jobi L. and Coix aquatic Roxb. (2n=
30) has been completed already[36], and
analysis of their genomic sequences
could help to find various practical
molecular markers including SSRs,
SNPs (single nucleotide polymor-
phisms) and InDels in Coix and to es-
tablish high-density linkage map for
obtaining important functional genes in
Coix, thereby laying the foundation of
developing molecular biological re-
search and marker-assisted breeding.
Comparative genomics also could be
used for expounding evolutionary rela-
tionships between sorghum and Coix
and between maize and Coix, and illu-
minating the differentiation and evolu-
tion process of Coix species. Good ge-
netic loci including pest resistance, dis-
ease resistance and stress resistance
in Coix could be fully explored, thereby
making a contribution to genetic im-
provement and new species breeding
of Coix and its relatives maize and
sorghum.
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(Continued from page 498)
Responsible editor: Yingzhi GUANG Responsible proofreader: Xiaoyan WU
should be considered as a leading de-
pendent variable. The plant height of
crop could be measured easily with
regional differences, so that the prob-
lem of regional differences in crop co-
efficient is solved and the calculation
results could be closer to the practical
conditions of various regions, provid-
ing a certain basis for the research of
the universal formula for calculating
crop coefficient according to plant
height.
The time interval between the
greatest corrected FAO-56-recom-
mended crop coefficient and the high-
est change rate of plant height was 10
d, the time interval between the
greatest measured crop coefficient
and the highest change rate of plant
height was 7 d, indicating a difference
of 3 d. The FAO-56-recommended
crop coefficient was different from the
measured one though it was correct-
ed. As the key water requirement peri-
ods of potato, the squaring stage and
the tuber expansion stage are very
important, and especially, the squaring
stage is only about 7 d during which
potato plant grew at a highest rate.
Crop coefficient affects crop irrigation
schedule [8-9]. It was found that the key
water requirement period of potato
should be 3 d earlier than the irrigation
time and irrigation schedule deter-
mined according to the crop coefficient
recommended by FAO-56 in Yunnan
currently, so as to ensure normal
growth of potato. This result provides
technical support for accurate irrigation
of potato.
In the future study on simple or
empirical formula calculation of crop
coefficient, plant height should be
considered as a main dependent vari-
able in that the calculation result would
be closer to the measured crop coeffi-
cient and the formula might be simpler.
Currently, the irrigation time of potato
should be 3 d earlier than the irrigation
time determined according to the cor-
rected FAO-56-recommended crop
coefficient and other factors, so as to
ensure normal growth of potato. The
crop coefficient of potato needs?fur-
ther study.
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