全 文 ：斑茅种质资源的表型性状及遗
传多样性
徐超华 1，2，陆鑫 1，2，马丽 1，2，刘新龙 1，2，刘洪博 1，2，
苏火生 1，2，林秀琴 1，2，蔡青 1，2，3* （1.云南省农
业科学院甘蔗研究所，云南开远 661699；2.云
南省甘蔗遗传改良重点实验室 ， 云南开远
661699；3.云南省农业科学院生物技术与种质
资源研究所，云南昆明 650223）
摘 要 [目的 ]有效评价和利用斑茅种质资
源，挖掘其优良性状。 [方法]以 162 份斑茅（云
南 74 份，福建 15 份，贵州 19 份，海南 18 份，
四川 14 份，江西 10 份，广东 4 份，广西 4 份，
浙江 4 份）为研究材料，通过 5 个数量性状和
21 个质量性状对其表型性状及遗传多样性进
行研究。 [结果] 表型性状分析结果表明：①斑
茅种质资源质量性状的 Shannon-Wiener 多样
性指数整体偏低，其中，福建斑茅的多样性指
数 (0.762 4) 最高， 广西斑茅的多样性指数
(0.294 2)最低；②数量性状遗传变异较丰富 ，
其中云南地区的变异系数(32.15%)最大，广西
地区的(14.95%)最小；③海拔高度与锤度呈极
显著负相关，纬度与株高呈极显著负相关。 遗
传分化系数和基因流结果显示，斑茅种质资源
群体的遗传变异主要来自于采集地内部，群体
之间存在较大的基因交流，遗传结构分化不明
显。 UPGMA 聚类分析结果表明，各居群间的
遗传距离与采集地之间有一定的相关性。 [结
论]该研究可为资源采集、杂交利用和优异基
因挖掘提供参考。
关键词 斑茅；表型性状；遗传多样性
基金项目 农业部作物种质资源保护项目
（2014NWB017）；国家甘蔗种质资源平台项目
（2012-044）；国家甘蔗产业技术体系开远综合
试验站（CARS-20-6-13）。
作者简介 徐超华(1986-)，男，湖北洪湖人，硕
士，研究实习员，主要从事甘蔗种质资源利用
研究，E-mail：xuchaohua_0435@sina.com。*通讯
作者，博士，研究员，主要从事甘蔗种质资源与
分子生物学研究，E-mail：caiqingysri@163.com。
收稿日期 2014-11-02
修回日期 2014-12-31
Phenotypic Traits and Genetic Diversity of
Erianthus arundinaceum Germplasm
Chaohua XU1,2, Xin LU1,2, Li MA1,2, Xinlong LIU1,2, Hongbo LIU1,2, Huosheng SU1,2, Xiuqin LIN1,2,
Qing CAI1,2,3*
1. Sugarcane Research Institute, Yunnan Academy of Agricultural Sciences, Kaiyuan 661699, China;
2. Yunnan Key Laboratory of Sugarcane Genetic Improvement, Kaiyuan 661699, China;
3. Biotechnology & Genetic Resources Institute, Yunnan Academy of Agriculture Sciences, Kunming 650223, China
Supported by Crop Germplasm Protection Project of Ministry of Agriculture of the
People’s Republic of China (2014NWB017); Fund for National Infrastructure of
Sugarcane Germplasm Resources (2012-044); Fund for Sugarcane Industry Technical
System Construction of Kaiyuan Field Station(CARS-20-6-13).
*Corresponding author. E-mail: caiqingysri@163.com
Received: November 2, 2014 Accepted: December 31, 2014A
Agricultural Science & Technology, 2015, 16(1): 35-39
Copyright訫 2015, Information Institute of HAAS. All rights reserved Agronomy and Horticultrue
S ugarcane (Saccharum L) asthe major sugar crop con-tributes to 90% of sugar pro-
duction in China[1]. At present, sugar-
cane varieties worldwide are mostly
the hybrids of tropical species Sac-
charum officinarum, S. spontaneum
and Indian species S. barberi, so they
have similar genetic background,
which limits the breeding and promo-
tion of new varieties[2]. Erianthus arun-
dinaceum, a species belonging to an
allied genus of sugarcane, widely dis-
tributed in Yunnan, Sichuan, Guizhou,
Fujian, Hainan and Guangxi and other
provinces of China, has excellent
traits, such as good tillering ability,
adaptability, drought tolerance and
abundant ecological types [3 -4]. In re-
cent years increasing attention has
been paid to the collection of E. arund-
inaceum resources [5-8], and their ge-
netic diversity had been evaluated
through morphological markers[9], cy-
tological markers[10-12], protein markers
[13-14] and mo-lecular markers [15-17], etc.
Besides, extensive studies have
been carried out on their tolerance
to cold [18], drought [19 -21] and disease
[22]. In this study , the genetic diver-
sity of phenotypic traits of 162 ac-
cessions of E. arundinaceum from
Abstract [Objective] This study was conducted to utilize and estimate Erianthus
arundinaceum resources and to develop their elite traits. [Method] Phenotypic traits
and genetic diversity were evaluated based on 5 quantitative traits and 21 qualita-
tive traits of 162 accessions of Erianthus arundinaceum collected from nine
provinces in China (74 in Yunnan, 15 in Fujian, 19 in Guizhou, 18 in Hainan, 14 in
Sichuan, 10 in Jiangxi, 4 in Guangdong, 4 in Guangxi and 4 in Zhejiang). [Result]
The Shannon-Wiener indices of qualitative traits among the E. arundinaceum popu-
lations from different provinces were generally low. The accessions from Fujian ex-
hibited the highest genetic diversity index (0.762 4), while those from Guangxi
showed the lowest one (0.294 2). There were great genetic variances in quantitative
traits, with mean variation coefficients ranging from 14.95% to 32.15%. The acces-
sions from Yunnan showed the highest variation coefficient (32.15%) and those from
Guangxi showed the lowest one (14.95%). Brix exhibited extremely significant nega-
tive correlation with altitude and plant height showed extremely significant negative
correlation with latitude. Coefficients of genetic divergence indicated that a high pro-
portion of total genetic variation was retained within the populations from different
regions, and the high gene flow showed that there were active genetic exchanges
among these populations, suggesting no significant genetic divergence among these
populations. According to genetic distance and UPGMA, there was a certain corre-
lation between genetic distance and different sampling regions of E. arundinaceum.
[Conclusion] The results provide theoretical references for resource collection , het-
erosis application and development of excellent genes of sugarcane germplasm.
Key words Erianthus arundinaceum; Phenotypic traits; Genetic diversity
DOI:10.16175/j.cnki.1009-4229.2015.01.012
Agricultural Science & Technology 2015
Table 1 Coefficients of variance of quantitative traits for Erianthus arundinaceum
populations from different sampling regions %
Region
Coefficient of variation
Stalk length Stalkdiameter Brix
Lamina
length
Lamina
width Mean
Yunnan 24.24 17.00 20.92 61.96 36.65 32.15
Sichuan 17.65 9.79 10.93 14.84 25.66 15.77
Guangdong 26.26 9.42 14.11 25.64 55.75 26.23
Guangxi 11.53 19.59 7.59 6.71 29.33 14.95
Jiangxi 15.76 22.04 8.40 4.54 27.29 15.60
Zhejiang 21.89 8.63 13.08 24.16 10.79 15.71
Hainan 13.63 17.60 10.23 12.60 37.04 18.22
Fujian 30.91 26.68 8.78 16.11 67.99 30.09
Guizhou 21.13 14.15 10.87 10.88 37.87 18.98
Mean 20.33 16.34 11.65 19.71 36.48
nine provinces of China was investi-
gated to explore their genetic back-
ground, with an attempt to provide the-
oretical references for resource collec-
tion , heterosis application and devel-
opment of excellent genes of sugar-
cane germplasm.
Materials and Methods
Materials
The 162 accessions of E. arundi-
naceum were preserved in the Na-
tional Nursery of Sugarcane Germ-
plasm Resources, among which, 74
samples were collected from Yunnan,
15 from Fujian, 19 from Guizhou, 18
from Hainan, 14 from Sichuan, 10 from
Jiangxi, 4 from Guangdong, 4 from
Guangxi and 4 from Zhejiang.
Measurement items and methods
The experiment was conducted
at the National Nursery of Sugar-
cane Germplasm Resources in
November 2008 . All the 162 acces-
sions of E. arundinaceum were sepa-
rately grown in cement frames (0.8 m
in diameter and depth), spacing 1 m to
each other. Six normal plants from
each accession were selected and sur-
veyed for five quantitative traits and 21
qualitative traits. The five quantitative
traits were lamina length, lamina width,
stalk length, stalk diameter and brix,
and the 21 qualitative traits were aerial
root, stalk shape, pipe, pith, internode
shape, internode color unexposed and
exposed, growth ring shape, root pri-
mordial, bud shape, bud placement,
sheath detached from culm, hair
group, shape of inner auricle, shape of
outer auricle, angle of lamina to culm,
lamina colour, wax band, corky patch
and growth crack. From which, ten of
the phenotypic traits with diversity: in-
ternode color unexposed, internode
color exposed, growth ring shape, root
primordial, bud shape, bud placement,
hair group, angle of lamina to culm,
lamina colour, wax band were
screened out and analyzed for their
genetic diversity. All the data were
standardized according to the litera-
tures [23] and [24]. The quantitative traits
were grouped into six levels for cluster
analysis together with the qualitative
traits.
The genetic diversity of the popu-
lations was measured by Shannon -
Wiener index which was calculated
with the formula as follows:
H’= -ΣPiLnPi
Wherein, H’ reflects the genetic
diversity of the populations; Pi is the
frequency of the i-th code value of a
quality trait[25]. The total genetic diversi-
ty (Ht), the genetic diversity within pop-
ulations (Hs), genetic divergence coef-
ficient (Gst) and gene flow value (Nm)
were all calculated using the formulas
described in literature[26].
Data analysis
Spss17.0 software was adopted
to analyze the coefficient of variations
(CV) and degree of dispersion of the
five quantitative traits. Excel and
NTSYSpc2.1 software were used to
calculate the phenotypic frequencies
of qualitative traits, and the Nei’s ge-
netic distance between the populations
from different provinces. Finally, a
dendrogram was constructed from
Nei’s genetic distance based on UPG-
MA (weighted pair group method using
arithmetic averages).
Results and Analysis
Coefficient of variation in the quan-
titative traits
As shown in Table 1, among the
five quantitative traits of the 162 E. ar-
undinaceum accessions, the average
coefficient of variation for lamina width
was the largest (36.48%), followed by
that for stalk length (20.33%) and lami-
na length (19.71%), while the coeffi-
cient of variation for brix (11.65%) was
the smallest. Therefore, quantitative
traits should be considered as an im-
portant index for the assessment of
E. arundinaceum resources.
Besides, by comparing the varia-
tion of the five quantitative traits for the
E. arundinaceum samples from differ-
ent regions, it was found that the popu-
lation from Fujian had the largest vari-
ation coefficient of stalk length
(30.91% ), while the population from
Guangxi had the smallest one
(11.53% ); the population from Fujian
also had the largest variation coeffi-
cient of stalk diameter (26.68%), while
the population Zhejiang had the small-
est one (8.63%); the largest variation
coefficient of brix (20.92%) was found
in population from Yunnan, and the
smallest (7.59% ) in population from
Guangxi; the largest coefficient of vari-
ation for stalk length (61.96% ) was
found in population from Yunnan, and
the smallest (4.54% ) in population
from Jiangxi; the largest coefficient of
variation for stalk width (67.99%) was
found in population from Fujian, and
the smallest (10.97% ) in population
from Zhejiang.
The coefficients of variation for
the five quantitative traits of the sam-
ples in each province were averaged,
and the resulting means ranged from
14.95% to 32.14, as shown in Table 1.
Among them, the mean in Yunnan
was the largest (32.15%), followed by
that Fujian (30.09%) and Guangdong
(26.23% ), while that in Guangxi was
the smallest (14.95%).
Genetic diversity in quality traits
As could be seen from Table 2,
the Shannon-Wiener indices for the 10
qualitative traits of the E. arundi-
naceum samples from each province
were averaged, and the resulting
means ranged from 0.294 2 0.762 4,
which were close to that of the de-
caploids of Saccharum spontaneum,
but lower than the Shannon-Wiener
36
Agricultural Science & Technology2015
Table 2 Shannon-Wiener index of qualitative trait for Erianthus arundinaceum populations from different sampling regions
Region
Internode
colour
unexposed
Internode
colour
exposed
Growth
ring shape
Root
primordial
Bud
shape
Bud
placement
Hair
group
Angle of
lamina to
culm
Lamina
colour
Wax
band Mean
Yunnan 0.656 2 0.293 6 0.608 6 0.313 0 0.603 6 1.035 1 0.609 2 0.377 6 0.763 6 0.754 5 0.601 5
Sichuan 0.519 6 0.410 1 0.682 9 0.519 6 0.257 3 0.651 8 0.690 2 0.656 0 0.876 0 0.876 0 0.613 9
Guangdong 0.000 0 1.039 7 0.693 1 0.000 0 0.562 3 0.562 3 0.693 1 0.000 0 0.000 0 0.000 0 0.355 1
Guangxi 0.000 0 0.000 0 0.562 3 0.000 0 0.000 0 0.562 3 0.000 0 0.562 3 0.693 1 0.562 3 0.294 2
Jiangxi 0.673 0 0.000 0 0.673 0 0.610 9 0.325 1 0.950 3 0.000 0 0.500 4 0.673 0 0.801 8 0.520 7
Zhejiang 0.000 0 0.000 0 0.000 0 0.693 1 0.562 3 1.039 7 0.000 0 0.000 0 0.000 0 1.039 7 0.333 5
Hainan 0.687 0 0.425 8 0.450 6 0.348 8 0.556 6 0.936 9 0.633 7 0.687 0 0.683 7 0.450 6 0.586 1
Fujian 0.690 9 0.690 9 0.673 0 0.500 4 1.077 6 1.060 6 1.322 9 0.392 7 0.970 1 0.244 9 0.762 4
Guizhou 0.680 6 0.436 2 0.336 5 0.206 2 1.113 8 0.753 4 1.093 8 0.514 7 1.059 2 0.824 7 0.701 9
Mean 0.434 1 0.366 3 0.520 0 0.354 7 0.562 1 0.839 2 0.560 3 0.410 1 0.635 4 0.617 2
Table 3 Correlation coefficients between quantitative traits and latitude or altitude
Geographical
index
Stalk
length
Stalk
diameter Brix
Lamina
length
Lamina
width
Altitude -0.132 3 0.039 1 -0.437** 0.062 -0.128 2
Latitude -0.296** -0.153 0 0.097 -0.102 0.077 0
*, significant correlation;**, extremely significant correlation.
Table 4 Genetic structure of Erianthus arundinaceum populations from different sampling
regions
Traits Genetic diversitywithin populations
Total genetic
diversity
Genetic differentiation
coefficient Gene flow
Quantitative traits 0.629 6 0.725 4 13.200 0 3.287 8
Quality traits 0.177 1 0.232 8 23.950 0 1.587 9
indices of the sugarcane germpla -
sm[27-28]. Among the nine provinces, the
Shannon-Wiener index in Fujian was
the largest (0.762 4), followed by that
in Guizhou (0.701 9), Sichuan (0.613 9)
and Yunnan Province (0.601 5), while
that inGuangxi was the lowest(0.294 2).
In addition, by comparing the Shan-
non-Wiener indices of the 10 qualita-
tive traits of the 162 E. arundinaceum
accessions, it was found that bud
placement had the largest Shannon-
Wiener index (0.839 2), followed by
lamina colour (0.635 4), and root pri-
mordial had the smallest diversity in-
dex (0.354 7).
Correlations of quantitative traits
with latitude and altitude
It could be concluded from Table
3 that altitude had an extremely signif-
icant negative correlation with brix, but
no significant correlations with stalk
length, stalk diameter, lamina length
and lamina width of E. arundinaceum
samples, indicating that brix deceased
with the increase in altitude. Latitude
had an extremely significant negative
correlation with stalk length, but no
significant correlations with stalk diam-
eter, lamina length, lamina width and
brix of E. arundinaceum samples, indi-
cating that their stalk length increased
with the increase of latitude. There-
fore, E. arundinaceum resources with
large plant and high brix could be easi-
ly found in low-latitude and low-altitude
areas, and those with short plant and
low brix in high-latitude and high-alti-
tude areas.
Genetic diversity of E. arundi-
naceum populations from different
provinces
The genetic diversity index for the
quantitative traits within E. arundi-
naceum populations from the nine
provinces was 0.629 6, and the total
genetic diversity index was 0.725 4.
The genetic diversity index for their
quality traits within the populations was
0.177 1, and the total genetic diversity
was 0.232 8. The results suggested
that the quantitative traits of these
E. arundinaceum resources had richer
genetic diversity than their quality
traits. For quality traits, the genetic di-
vergence coefficient was 23.95% and
gene flow was 1.587 9; for quantitative
traits, the genetic divergence coeffi-
cient was 13.20%, and the gene flow
was 3.287 8, suggesting that there
was active genetic exchanges and no
obvious genetic divergence among the
E. arundinaceum populations from dif-
ferent regions.
Cluster analysis of the E. arundi-
naceum populations from different
regions
The Nei’s genetic distances be-
tween the E. arundinaceum popula-
tions from different regions were rela-
tive small, ranging from 0.083 4 to
0.513 7, with an average value of
0.227 2, indicating a low degree of ge-
netic divergence among the popula-
tions. The genetic distance between
the populations from Jiangxi and
Hainan (0.083 4) was the smallest,
and that between the populations
from Sichuan and Jiangxi (0.097 5)
was the second smallest; the genetic
distance between the populations
from Guangxi and Zhejiang was the
largest (0.513 7), followed by that be-
tween the populations from Zhejiang
and Hainan (0.497 8).
As shown in Fig.1, the E. arundi-
naceum population from Zhejiang was
most distantly related to the popula-
tions from other regions, followed by
that from Guangdong and Guangxi.
The populations from Jiangxi and
Hainan were most closely related and
clustered together, indicating that they
had the most similar genetic back-
ground, and then they clustered with
the populations from Sichuan, Fujian,
Guizhou and Yunnan. All the results
showed that the phenotypes of the
E. arundinaceum samples in this
study had a certain relationship with
their geographical locations.
Conclusion and Discussion
37
Agricultural Science & Technology 2015
Fig.1 UPGMA dendrogram based on Nei’s
genetic distance for Erianthus arundi-
naceum populations from different
sampling regions
Liu et al.[28] studied the genetic di-
versity of 1 160 sugarcane accessions
from 20 regions of 13 countries, and
provided useful references for the se-
lection of parental materials, hybrid
combinations and construction of core
collection for sugarcane breeding. The
study of Xiao et al. [29] which was about
genetic diversity in morphological
traits of Miscanthus floridulus provided
theoretical information for the breed-
ing and genetic improvement of Chi-
nese M. floridulus resources. The
study of Liu et al.[27] discovered the di-
verse Saccharum spontaneum re-
sources in China. Our findings proved
that abundant genetic variation existed
in the morphological traits of E. arund-
inaceum populations from different re-
gions. The variation coefficients of the
five morphological traits (such as stalk
length and stalk diameter) among the
E. arundinaceum populations from the
nine provinces ranged from 1.65% to
36.48% , with an average of 20.90% ,
indicating the great genetic differences
and abundant genetic diversity of
them. Besides, the quantitative traits of
the E. arundinaceum populations
shared significant correlations with lat-
itude and altitude. Among them, brix
had an extremely significant negative
correlation with altitude, while stalk
length had an extremely significant
negative correlation with latitude. UP-
GMA cluster analysis showed that the
morphological traits of the E. arundi-
naceum populations had certain rela-
tionship with their location, indicating
that they were affected by geographi-
cal environments.
The genetic divergence coeffi-
cient for quality traits among the
E . arundinaceum populations from
different regions was 23.95%, indicat-
ing that only 23.95% of the genetic
variation was caused by their geo-
graphical locations, while 76.05% of
the variation was retained within the
populations. The genetic divergence
coefficient for quantitative traits among
the E. arundinaceum populations was
13.20% , indicating that 86.8% of the
variation was retained within popula-
tions. In summary, extensive genetic
exchanges existed in both the qual-
ity and quantitative traits of these
E. arundinaceum populations, while
the gene flow of quantitative traits was
over two folds of qualitative traits, indi-
cating that there should be more em-
phasis on quantitative traits than
quality traits in resource evaluation of
E. arundinaceum. The large gene flow
also indicated a low degree of genetic
divergence , which may be related
to the propagation and selection of
E. arundinaceum under natural envi-
ronmental conditions[30].
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38
Agricultural Science & Technology2015
(Continued from page 24)
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Responsible editor: Qingqing YIN Responsible proofreader: Xiaoyan WU
was higher than that of non-grazing
population throughout the growth peri-
od of A. cristatum, proving that grazing
stress damaged A. cristatum popula-
tion and exacerbated membrane lipid
peroxidation. SOD activity was signifi-
cantly increased from heading to flow-
ering stage, but was rapidly decreased
at seed maturity stage, but the differ-
ence between grazing and non-graz-
ing populations was not significant, in-
dicating that SOD activity changed lit-
tle under grazing stress, and the in-
creased SOD activity could be a result
of drought and other environmental
stresses. POD activity was increased
from heading stage, and it was higher
in grazing population than in non-
grazing population, suggesting that
POD played an important role for
clearing reactive oxygen and main-
taining cell stability of A. cristatum un-
der drought stress.
SOD activity changed significantly
among the different growth stages, but
had no obvious response to grazing
pressure. POD activity in A. cristatum
was increased under grazing stress.
The soluble sugar content of grazing
population was higher than that of non-
grazing population of A. cristatum at
tillering stage, and the difference was
significant (P<0.05). MDA content of
grazing population was higher than
that of non-grazing population during
throughout the whole growth period of
A. cristatum, indicating that grazing
stress caused the accumulation of
free radicals and exacerbated mem-
brane lipid peroxidation of A. crista-
tum. POD played an important role for
relieving the damage caused by graz-
ing stress and clearing the harmful
substances of A. cristatum.
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