全 文 ：作物学报 ACTA AGRONOMICA SINICA 2008, 34(7): 1206−1212 http://www.chinacrops.org/zwxb/
ISSN 0496-3490; CODEN TSHPA9 E-mail: xbzw@chinajournal.net.cn

Foundation items: Supported by Anhui Provincial Natural Science Foundation (01041102), A key project of scientific and technical attack plan of
Anhui Province (06013104B), A Key Project Grant from Anhui Science & Technology Department (04023069), Outstanding talent subsidy
of 10th five-year plan of higher education of Anhui Province (200579), and Key Subject Foundation of Anhui Science and Technology
University (YZD2004-03).
Biography: ZHAN Qiu-Wen (1963−), male, PhD, professor. Research field: crop genetics and breeding. Tel: 0550-6733117; E-mail: qwzhan@163.com.
Received(收稿日期): 2007-08-17; Accepted(接受日期): 2007-11-15.
DOI: 10.3724/SP.J.1006.2008.01206
Cytogenetical Studies on Sorghum (S. bicolor), Sudangrass (S. sudanense),
and Their Hybrid
ZHAN Qiu-Wen1,2, ZHU Li-Meng1, WU Juan-Juan2, ZHANG Wei1, and ZHANG Tian-Zhen2
(1 College of Plant Science, Anhui Science and Technology University, Fengyang 233100, Anhui; 2 National Key Laboratory of Crop Genetics &
Germplasm Enhancement, Nanjing Agricultural University, Nanjing 210095, Jiangsu, China)
Abstract: Sudangrass (Sorghum sudanense) and sorghum (S. bicolor) are two members of the genus Sorghum (Gramineae).
Their hybrids have exhibited favorable forage yields and overall improved quality and disease resistance. They are widely used in
aquaculture, production of livestock food and environmental protection, but some disagreements exist as to whether they actually
belong to the same species. The Karyotypes of two sudangrass varieties, two sorghum varieties, three sorghum-sudangrass F1 hy-
brids were analyzed by the method of cell wall degradation and hypotonic treatment for chromosome preparations, and the meio-
sis behavior of three F1 hybrids and the chromosome numbers of two F2 hybrids were observed. The result showed that the karyo-
typic types of sorghum and sudangrass were 1A, and the karyotype formula was 2n=18m+2sm (sat) for Sa, 2n=20m for 3042A
and 3042A×Sa F1, and 2n=20m(sat) for the others. The difference of chromosomes among three groups (i.e. sudangrass, sorghum,
and sorghum-sudangrass hybrid F1) were not significant in terms of absolute long arm, absolute short arm, absolute total length,
arm ratio (l/s), relative total length (P>0.05); the change of chromosome length was not obvious in sorghum and sudangrass. The
paired chromosome configuration of sorghum-sudangrass hybrid F1 in pollen mother cells at diakinesis and metaphaseIwas
2n=2x=20(10II) and the chromosome pairing behavior was regular, but the frequencies of rod bivalent of Tx623A×S722 F1,
3042A×Sa F1, and Tx623A×Sa F1 were 4.887, 5.710, and 5.126, respectively; those of ring bivalent were 5.113, 4.290, and
4.874, respectively. At anaphase I, the paired chromosomes of sorghum-sudangrass hybrid F1 could separate from each other. The
chromosome numbers of F2 hybrid were 20 (2n=20). Therefore, the sudangrass/sorghum relationship is sufficiently close.
Keywords: Sudangrass [Sorghum sudanense (Piper) Stapf]; Sorghum [Sorghum bicolor (L.) Moench]; Karyotypes; Chromosome;
Cytogenetics
苏丹草高粱及其杂种的细胞遗传学研究
詹秋文 1, 2 朱立猛 1 吴娟娟 2 张 伟 1 张天真 2
(1安徽科技学院植物科学学院, 安徽凤阳 233100; 2南京农业大学作物遗传与种质创新国家重点实验室, 江苏南京 210095)
摘 要: 苏丹草(S. sudanense)与高粱(S. bicolor)均为禾本科高粱属植物, 两者的杂种优势明显, 杂交种品质好, 抗逆
性强, 在水产、畜禽养殖及资源利用与环境保护上有着广阔的开发利用前景, 但两者是否属于同一个种至今存在争
议。本文采用去壁低渗-火焰干燥法, 分析了 2 份苏丹草、2 份高粱及其 3 个杂种 F1有丝分裂核型, 观察了 3 个杂种
F1减数分裂染色体行为和 2 个杂种 F2体细胞染色体数目。结果表明, 苏丹草、高粱及其杂种 F1均为 1A 核型, 但核
型公式不完全相同 , 苏丹草 Sa 为 2n=18m+2sm(sat), 高粱 3042A 和 3042A×Sa F1 为 2n=20m, 其余材料均为
2n=20m(sat)。苏丹草、高粱及其杂种 F1 3 者在 10 条染色体的绝对长臂、绝对短臂、绝对全长、臂比和相对全长上
差异均不显著(P>0.05), 说明苏丹草与高粱在染色体长度上的变化不明显。杂种 F1花粉母细胞减数分裂, 终变期和中
期 I 染色体核型和数目清晰可见(2n=2x=20), 配对行为规则; 棒状和环状二价体的频率因组合不同而异, Tx623A×
S722 F1、3042A×Sa F1和 Tx623A×Sa F1棒状二价体频率分别为 4.887、5.710和 5.126, 环状二价体频率分别为 5.113、
4.290 和 4.874; 在后期 I, 配对的染色体能够正常分离。杂种 F2体细胞染色体数目为 20(2n=20)。因此, 苏丹草与高
粱的亲缘关系非常近。
第 7期 ZHAN Qiu-Wen et al.: Cytogenetical Studies on S. bicolor, S. sudanense, and Their Hybrid 1207


关键词: 苏丹草; 高粱; 杂交种; 染色体; 细胞遗传学
Sudangrass [Sorghum sudanense (Piper) Stapf]
and sorghum [Sorghum bicolor (L.) Moench] are two
members of the genus Sorghum (Gramineal)[1]. Their
hybrids have favorable forage yields and overall im-
proved quality and disease resistance. They are widely
used in aquaculture, production of livestock food and
environmental protection[2]. However, some disagree-
ments exist as to whether they actually belong to the
same species. Snowden[3], using a worldwide collec-
tion of sorghum accessions, classified plants according
to spikelet, anthotaxy and plant traits, and treated su-
dangrass as S. sudanense, a species different from
sorghum (S. bicolor). Many other researchers have
also considered sudangrass and sorghum as different
species—S. sudanense and S. bicolor [4-9]. De Wet &
Huckaby[10], based on examination of topical and
primitive samples, suggested sudangrass was derived
from hybrids between wild and cultivated sorghum.
Harlan & de Wet[11] offered a simplified classification
of cultivated sorghum, and placed sudangrass in the
subtribe of the sorghum bicolor race according to the
characteristics of grain, glume and spikelet. However,
de Wet[12] placed sudangrass into the subspecies,
drummondii, of S. bicolor. Some reaserchers, concur-
ring with the separate-taxon argument, have suggested
sudangrass and sorghum are the same species[13-17]. Ma
et al.[18] called sudangrass, forage sorghum, a close
relative of S. bicolor.
The previous study results were basically based
on morphological classification, resulting in the dif-
ferent classification of sudangrass and sorghum. To
clarify taxonomic confusion of the two species, the
mitosis karyotype and meiosis behavior of sudangrass,
sorghum and their hybrids were studied based on
quantitative classification and isoenzyme[19-20]. Yu et
al.[21-22] observed the meiosis behavior of F1 and F2
hybrids between male sterile 11A of S. bicolor and
three kinds of S. sudanense, but didn’t study on mitotic
karyotypes and the differences of chromosomes in-
dexes between sorghum and sudangrass. This paper is
to study the cytogenetic differences between sudan-
grass and sorghum in chromosomes characters of two
sudangrass varieties, two sorghum varieties, three F1
hybrids and two F2 hybrids.
1 Materials and Methods
Plants of two sudangrass varieties (Sa, S722), two
sorghum varieties (Tx623A, 3042A), three F1 hybrids
(Tx623A×S722 F1, 3042A×Sa F1, and Tx623A×Sa
F1) and two F2 hybrids (Tx623A× S722 F2 and
Tx623A×Sa F2) were grown in the experimental gar-
den of Anhui Science and Technology University in
2003–2005. To exempt from the operation excising
stamen of female parent, and guarantee that F1 seeds
were real hybrids sorghum sterile lines (Tx623A and
3042A) were selected as female parents of F1 hybrids.
The experiment for somatic cell chromosome
slides was carried out using the method of cell wall
degradation and hypotonic treatment in chromosome
preparation[23]. Seeds were cleaned, then dipped in
water for 30 min before pregermination in incubator at
25℃. Seeds sprouted after about 12 h; 36 h later root
tips grew to 0.5–2.5 cm long which had the highest
cell mitotic index. Major root tips cut about 0.5 cm
were pretreated in 1:1 mixture of 0.2% colchicine and
0.002 mol L−1 8-hydroxy quinoline for 2 h before they
were dipped in 0.075 mol L−1 KCl for pre-hypotonic
treatment for 20 min; then they were fixed in Carnoy’s
solution (absolute ethanol: glacial acetic acid=3:1) at
4℃ for 24 h, and dissociated in 2% pectase-cellulase
mixture at 32℃ for 0.5–1.0 h; after a post-hypotonic
treatment in distilled water for 10 min they were fixed
again in Carnoy’s solution for 10 min, at last stored in
70% alcohol. 80 cells with dispersed chromosomes
were selected in each material to survey various
chromosomes indexes. Karyotypes analysis was ac-
cording to the method described by Li et al.[23] The
chromosomal data were analyzed by SAS 8.0 rystem.
Young spikes in the pollen mother cell experi-
ment were picked at different growth periods in vari-
ous materials. The spikes were fixed in Carnoy’s solu-
tion (3:1) for 24 h; and washed twice by 95% alcohol
for 10 min each time; afterwards, dipped in 80% al-
cohol for 10 min until without acidity, and stored in
70% alcohol[23]. Chromosome pairing behavior was
observed in meiosis of pollen mother cells (PMCs)
using the acetic-carmine smear method [23]. Micro-
photographs were taken from permanent meiotic
preparations by the Olympus Camera rystem. 150 pol-
len mother cells of each F1 hybrid were observed, and
the chromosome number for each cell was counted.
The chromosome pairing behavior and configuration
were investigated at diakinesis and metaphase I.
2 Results
2.1 Karyotype of sudangrass, sorghum and F1
hybrid
Chromosome photographs and karyotypes of two
sudangrass varieties, two sorghum varieties and three
F1 hybrids are shown in Fig. 1, and the absolute long
arm, absolute short arm, absolute total length, arm
ratio, and relative total length were measured (original
data omitted). The karyotypic types of seven materials
were all 1A. The karyotype formula was 2n=18m+2sm
(2sat) for Sa, 2n=20m for 3042A and Tx623A×S722
1208 作 物 学 报 第 34卷

F1, and 2n=20m (2sat) for the others. The karyotype
formulas were different, whether within the two su-
dangrass varieties or within the two sorghum varieties
(Table 1).
2.2 Chromosome index differences among
sudangrass, sorghum, and F1 hybrid
Seven materials were divided into three groups:
sorghum, sudangrass, and F1 hybrid (Table 2). Five
indexes of chromosomes (absolute long arm, absolute
short arm, absolute total length, arm ratio, and relative
total length) were tested using Duncan’s SSR test. The
results showed that the differences of all chromosome
indexes were not significant among three groups
(P>0.05).
2.3 Meiosis behavior of pollen mother cells of
F1 hybrid
The chromosomes of 16.1% of observed cells
looked like long thin strand that overlapped or inter-
twined at prophase I in three F1 hybrids (Tx623A×
S722 F1, 3042A×Sa F1, and Tx623A×Sa F1).
The number of chromosomes could be counted
clearly at diakinesis. The paired chromosome configu-
ration in pollen mother cells of sorghum-sudangrass
hybrid F1 was 2n=2x=20(10II). The chromosome pair-
ing behavior was regular, and the frequency of paired
bivalent was 10, but the frequencies of rod bivalent of
Tx623A×S722 F1, 3042A×Sa F1, and Tx623A×Sa
F1 were 4.887, 5.710, and 5.126, respectively; those of
ring bivalent were 5.113, 4.290, and 4.874, respec-
tively. The chromosomes aligned on the equator of the
cell at metaphaseI. At anaphaseI, the paired chromo-
somes of sorghum-sudangrass hybrid F1 could separate
from each other (Fig. 2). The frequencies of rod biva-
lent of F1 hybrids were about 50%, and the synapses of
homologous chromosomes relaxed, but multivalence.



Fig. 1 Mitotic metaphase chromosomes and karyograms of two sudangrass varieties, two sorghum varieties, and three F1 hybrids (bar=5 μm)

Table 1 Karyotype formulas of two sudangrass varieties, two sorghum varieties, and three F1 hybrids
Name Ratios of chromosomes with arm ratio>2
Chromosomes length ratio
(Longest / shortest) Karyotype Karyotype formula
Sa 0 1.680:1 1A 2n=18m+2sm (2sat)
S722 0 1.075:1 1A 2n=20m (2sat)
Tx623A 0 1.237:1 1A 2n=20m (2sat)
3042A 0 1.310:1 1A 2n=20m
Tx623A×S722 F1 0 1.520:1 1A 2n=20m (2sat)
3042A×Sa F1 0 1.955:1 1A 2n=20m
Tx623A×Sa F1 0 1.594:1 1A 2n=20m (2sat)

第 7期 ZHAN Qiu-Wen et al.: Cytogenetical Studies on S. bicolor, S. sudanense, and Their Hybrid 1209


Table 2 The chromosome indexes and F protected multiple comparisons of three groups
No. of chromo-
some Type
Absolute long arm
(μm)
Absolute short arm
(μm)
Absolute total length
(μm)
Arm ratio
(l/s)
Relative total length
(%)
1 F1 hybrid 2.335 a 1.795 a 4.130 a 1.325 a 13.720 a
Sorghum 2.504 a 1.794 a 4.298 a 1.404 a 11.505 a
Sudangrass 1.958 a 1.592 a 3.550 a 1.225 a 11.422 a

2 F1 hybrid 1.834 a 1.604 a 3.438 a 1.150 a 11.514 a
Sorghum 2.492 a 1.753 a 4.245 a 1.432 a 11.373 a
Sudangrass 2.098 a 1.375 a 3.472 a 1.577 a 11.173 a

3 F1 hybrid 1.822 a 1.334 a 3.155 a 1.375 a 10.676 a
Sorghum 2.337 a 1.624 a 3.961 a 1.455 a 10.625 a
Sudangrass 1.919 a 1.465 a 3.384 a 1.309 a 10.836 a

4 F1 hybrid 1.655 a 1.336 a 2.992 a 1.235 a 10.145 a
Sorghum 2.164 a 1.570 a 3.734 a 1.384 a 10.021 a
Sudangrass 1.768 a 1.421 a 3.189 a 1.235 a 10.079 a

5 F1 hybrid 1.619 a 1.266 a 2.885 a 1.282 a 9.735 a
Sorghum 2.031 a 1.583 a 3.615 a 1.281 a 9.691 a
Sudangrass 1.732 a 1.418 a 3.150 a 1.207 a 9.941 a

6 F1 hybrid 1.542 a 1.235 a 2.777 a 1.250 a 9.341 a
Sorghum 1.965 a 1.602 a 3.568 a 1.224 a 9.562 a
Sudangrass 1.745 a 1.378 a 3.123 a 1.263 a 9.837 a

7 F1 hybrid 1.557 a 1.159 a 2.717 a 1.336 a 9.139 a
Sorghum 2.052 a 1.472 a 3.524 a 1.402 a 9.439 a
Sudangrass 1.691 a 1.366 a 3.057 a 1.222 a 9.595 a

8 F1 hybrid 1.525 a 1.159 a 2.684 a 1.339 a 9.025 a
Sorghum 1.971 a 1.537 a 3.508 a 1.279 a 9.395 a
Sudangrass 1.696 a 1.338 a 3.034 a 1.250 a 9.508 a

9 F1 hybrid 1.411 a 1.135 a 2.546 a 1.243 a 8.548 a
Sorghum 1.957 a 1.535 a 3.491 a 1.272 a 9.351 a
Sudangrass 1.614 a 1.291 a 2.905 a 1.235 a 9.077 a

10 F1 hybrid 1.375 a 1.040 a 2.415 a 1.316 a 8.156 a
Sorghum 1.945 a 1.424 a 3.369 a 1.366 a 9.037 a
Sudangrass 1.559 a 1.206 a 2.764 a 1.294 a 8.533 a
Values followed by the same small letter are not significantly different (P>0.05) in the same chromosome.

and univalent were not observed. The pollen fertility
rates (Fertility pollen number/Total pollen number×
100%) of the F1 hybrids above were 92.62%, 95.21%,
and 96.53%, respectively, the seed setting rates (Seed set
number/Total florets×100%) were 69.15%, 70.32%, and
72.28%, respectively. Therefore, there was a higher ho-
mology between sudangrass and sorghum
2.4 Chromosome numbers and karyotype of F2
hybrid
Eighty cells from each F2 hybrid were observed.
The result showed that a pair of satellites was observed
in F2 hybrids, but the karyotype formulas were differ-
ent possibly because of the segregation of F2 hybrid
population. The chromosome number of sorghum-
sudangrass hybrid F2 was 20 (2n=20), indicating that
the chromosomes for sudangrass and sorghum could
not only pair regularly, but transmit stably. This con-
firmed the experimental results of mentioned above;
therefore, sorghum and sudangrass are very closely
related to each other.
1210 作 物 学 报 第 34卷



Fig. 2 Meiotic chromosome behavior of pollen mother cell of F1 hybrid (×1 600)
A, B, C: Tx623A×S722 F1; D, E, F: 3042A×Sa F1; G, H, I: Tx623A×Sa F1.

3 Discussion
3.1 Karyotype formula of sorghum-sudangrass
hybrid F1 and its parents
He C R et al.[24] found that the karyotypes of sor-
ghum cultivars were 1A, 1B, or 2A by an ASG method.
Wang L Y et al.[25] stated the karyotype of sudangrass
was 1A by using three kinds of sudangrass. Zhan Q W
et al.[2] reported that the karyotypes of sudangrass
varieties were 1A while the karyotypes of sorghum
varieties were 1A, 2A, or 2B. The differences between
karyotypes of sorghum and sudangrass as reported by
different researchers were possibly because of the
cross-pollination, evolution, or the adaptation resulting
in the difference of chromosome shape. In this study,
the karyotypes of sorghum and sudangrass were 1A,
maybe relating to selected experimental material.
As shown in experiment results, the karyotype
formulas were different within the two sudangrass va-
rieties and within the two sorghum varieties, indicating
that sudangrass and sorghum contain some variations.
Similar results were observed by He C R et al.[24] in his
karyotype analysis for different sorghum varieties.
Tx623A×S722 F1 and Tx623A×Sa F1 had the same
karyotype formula with their parents, but the satellite
was not found on 3042A×Sa F1, which had different
karyotype with parent Sa. Perhaps the satellite chro-
mosome was lost or mutated during the procedure of
hybridization. Further study needs to be done to con-
firm this.
3.2 Meiosis behavior of pollen mother cells for
hybrid F2
Meiosis behavior of pollen mother cells for F2
hybrid was not investigated in this study because one
of the parents for F2 hybrids was the sorghum sterile
line. The meiosis behavior for the male-sterile plants
was abnormal (some lagging chromosomes could be
observed at anaphase I). The authors selected sorghum
male-sterile lines Tx623A and 3042A as the male par-
ent of sorghum-sudangrass hybrid based on the fol-
lowing two reasons: these two lines were classic elite
materials for the sorghum genetics and breeding; it
would exempt from the operation excising stamen of
female parent, and guarantee that F1 was a real hybrid.
Further research is required to find out the sorghum
parent from conventional cultivars rather than from
male-sterile lines
3.3 Relationship on Sudangrass (S. sudanense)
and Sorghum (S. bicolor)
In classical studies on sorghum taxonomy, the
discussion of various characteristic shapes was based
mostly on personal experience, making it especially
第 7期 ZHAN Qiu-Wen et al.: Cytogenetical Studies on S. bicolor, S. sudanense, and Their Hybrid 1211


difficult to classify complex groups as well as massive
character relationships. Using several disconnected
shapes, it was impossible to provide the precise
analyses of the genetic differences between sudangrass
and sorghum. The reason is that it is very difficult to
fully explain the relationship and to objectively dif-
ferentiate the phenomena of “tend to separate” and
“tend to homology” between sudangrass and sorghum
in the phylogenetic process. Given the lack of solid,
quantitative data, it is not surprising that various
scholars have failed to reach a consensus on the clas-
sification of sorghum and sudangrass[1,3-5,9-18]. Zhan et
al.[19] found distinct genetic differences between sor-
ghum and sudangrass using quantitative classification,
which had been divided into two groups, based on ob-
serving 66 traits of 10 sorghum and sudangrass varie-
ties. Zhan et al.[20] also analyzed the isoenzymes and
found that the zymograms were not the same between
sorghum and sudangrass; the specific esterase bands of
sorghum and sudangrass were observed, but the analy-
sis of peroxidase isozyme could not distinguish sor-
ghum from sudangrass. Phylogenetic relationships of
sorghum and sudangrass were investigated using
RAPD and SSR Markers (be published), indicating
that it was hard to distinguish between sorghum and
sudangrass with increasing amounts of sorghum and
sudangrass varieties because sorghum and sudangrass
were very closely related to each other. It should be
studied by the cytogenetical method whether dividing
them into two species or into two different subspecies
of the same species. This study showed that the change
of chromosome length was not obvious in sorghum
and sudangrass; in F1 hybrid, the chromosomes paired
and divided regularly in meiosis, the anthers could
dehisce and the plants could set seeds normally, and
the pollen fertility rates were above 92%, as well as
the seed setting rates were above 69%. Our results
appear to support the viewpoint of sorghum and su-
dangrass as the same species[13-17]. It’s unsuitable to
divide them into two species. Sudangrass should be
regarded as a subspecies, S. bicolor (L.) Moench subsp.
sudanense, in S. bicolor.
4 Conclusion
Karyotypic types of sorghum and sudangrass
were 1A, but the karyotype formulas of seven varieties
were not the same. The differences of chromosomes
among three groups (i.e. sudangrass, sorghum, and
sorghum-sudangrass hybrid F1) were not significant in
terms of absolute long arm, absolute short arm, abso-
lute total length, arm ratio (l/s), relative total length
(P>0.05); the change of chromosome length was not
obvious in sorghum and sudangrass. The paired chro-
mosome configuration of sorghum-sudangrass hybrid
F1 in pollen mother cells at diakinesis or metaphase I
was 2n=2x=20(10II) and the chromosome pairing be-
havior was regular, but the frequencies of rod bivalent
of Tx623A×S722 F1, 3042A×Sa F1 and Tx623A×Sa
F1 were 4.887, 5.710, and 5.126, respectively. At ana-
phase I, the paired chromosomes of sorghum-sudan-
grass hybrid F1 could be separated from each other.
The chromosome numbers of F2 hybrid were 20
(2n=20). Therefore, the sudangrass/sorghum relation-
ship is sufficiently close.
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