全 文 ：广 西 植 物 Guihaia 22(4)：352— 356 2002 年 7月
大鹅观草与阿拉善鹅观草杂种的
形态学和细胞学研究
张海琴 ，周永红，郑有 良，杨瑞武，丁春邦
(四川I农业大学小麦研究所，四川都江堰 6l1830)
摘 要 ：为探讨阿拉善鹅观草(R．口z口 口(Keng)S．I ．Chen)的染色体组组成 ．进行 了大鹅观草( ．grandis
Keng(2n=24，StStYY))和阿拉善鹅观草种间杂交 ．对这两个种及其杂种 F 的形态学 、繁育学和减数分裂染
色体配对行为进行 了研究 。结果表 明：杂种 F 的形态特征介于父母本之间，其花粉母细胞减数分裂染色体平
均构型为：2O．40 I+3．69Ⅱ+O．O9Ⅲ+O．04Ⅳ。表明阿拉善鹅观草含有一个修饰的 St基因组 ，即St‘。
关键词：大鹅观草；阿拉善鹅观草；种间杂交 ；减数分裂 ；基因组；染色体配对
中图分类号 ：Q942 文献标识码：A 文章编号 ：1000—3142(2002)04—0352—05
M orphol ogical and cytol ogical studies on hybrid
。 alashanica
(Poaceae：Triticeae)
ZHANG Hai—qin，ZHOU Yong—hong，ZHENG You—liang，
YANG Rui—WU，DING Chun—bang
(Triticeae Research Institute，Sichuan Agricultural Universit3·，Dujiangyan 611830，China)
Abstract：Interspecific hybridization was carried OUt between Roegneria alashanica (Keng)S．I ．Chen and R．
grandis Keng (2n一 24．StStYY)．The morphology，fertility and chromosome pairing behavior in meiosis of the
parents and their hybrid F1 were studied tO determine the genomic constitution of R．alashanica．The Fl hybrid
appeared intermadiate between the two parents．The meiotic configuration of the hybrid was 20．40 I+ 3．69Ⅱ
+ 0．09Ⅲ+ 0．04 lV．The results indicated that R．alashanica has one slightly modified St genome，be designated
as st‘．
Key words：Roegneria grandis；R．alashanica；interspecific hybrid；meiosis；genome}chromosome pairing
Roegneria C． Koch of the tribe Triticeae large number of species，subspecies，and the morpho一
(Gramineae)is a very large and widely distributed
genus which includes about 130 species in the world
and nearly 7O in China ”．Since this genus contains a
logical limits among some of them are not SO clear，SO
taxonomy seems very difficult within the genus．
Roegneria grandis Keng and R． alashanica
收稿 日期 ：2001—1O一26
作者简介 ：张海琴(1977一)，女 ，浙江金华人 ，硕士，主要从事小麦族生物系统学研究 。
基金项 目：教育部骨干教师资助项 目教技 司[2000-163号
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4期 张海琴等：大鹅观草与阿拉善鹅观草杂种的形态学和细胞遗传学研究 353
(Keng)S．I ．Chen are tetraploid (2n= 4x 28)
Derennial species，awnless on the lemma and distribut—
ed in northwest China．R．grandis has two genomes
StStYY ，while the genomic constitution of R．
口Z盘 ^口 is stil unknown。 ． Morphological1y，R．
口Z口 ，2ica has a solitary spikelet at each rachis node
and haye the disarticulation of the rachila above the
glumes and beneath the florets．All of these charac—
ters were considered to be similar to R．elyt, igioides
Yen et J．I ．Yang．Considering the morphological
similaritv between R．alashan／ca and R．elytrigioides，
Cai‘ treated it as R． alqshanica var． elyt,‘igioides
(Yen Pt J⋯I Yang)I ．B．Cai．I u reported that
R． elytrigioides contained two slightly modified St
genomes—St。St。，and transferred it to Psedoroegn—
eria． In order to investigate the genomic constitution
of R．alashanica (Keng)S． I ． Chen，intergeneric
hybridization was carried out between R．grandis and
R．aeashnn cn．Morphological comparison and chro—
mosome pairing behavior at metaphase I of polen
mother cells (PMCs)were observed in the parental
species and their Fr hybrid．
Materials and M ethods
Roegneria grandis Keng was collected in Shanxi，
China and R．alashanica (Keng)S．I ．Chen was
collected from Ningxia，China．Both of them were
grown in the field at the Triticeae Research Institute，
Sichuan Agricultural University．
For hybridization was：the spikes of尺．grandL~
were emasculated and put in a cellulose bag． Two
days after the maternal stigmas were brushed，the
hand—emasculated spikes were polinated with newly
broken anthers of R． alashanica． The hybrid seeds
were germinated on filter paper in petri—dishes and
then the seedling were sowed in pots at two leaf
stage．
Sixteen morphological characters were compared
between the parents and their F1 hybrid．For cytolog—
ical observation， the young spikes were fixed in
Carnoy’s Ⅲ solution (6 alcohol：3 chloroform ：1
acetic acid)，and stored in a refrigerator． Chromo—
some pairing was observed at the metaphase I(M 1)
of the pollen mother ceils (PM Cs)using the acetic—
carmine squash technique． M icrophotographs were
taken from permanent meiosis preparations． Pollen
grains of parents and F1 hybrid from mature anthers
were stained in an aqueous 12一KI solution for pollen
fertility studies．
Voucher specimens of the parents and．F1 hybrid
are deposited in Triticeae Research Institute，Sichuan
Agricultural University．
Results
1．1nterspecific cross
A total of 45 florets of R．grandis were pollinat—
ed with R． ala hanica and 9 seeds were obtained．
Seed set was 20．00 ．Two of the nine seeds were
gel-minated and developed into adult plants．The hy—
brid plants grew vigorously and set spikes later．
2． M orphol ogical comparison among the parental
species and the F1 hybrid
Sixteen morphological characters of the parents
and the FI hybrid were measured (Table 1)． R．
grandis was generally larger than R． alashanica in
many aspects，e．g．1ength and width of leaf and flag
leaf，length of spike，glume(first and second)，palea
and anther． R． alashanica had more spikelets per
spike and．florets per spikelet than those of R． 口，卜
d ．Both of them had no awn on lemma，while the
F1 hybrid had an awn，around 1．36 cm．The general
appearance of the F1 hybrid was intermediate between
their two parents or near to one of the parental
species(Fig．1)．
3．M eiosis in the F1 hybrid
The chromosome pairings at the M I of PM Cs in
the parental species and the F1 hybrid are listed in
Table 2． The meiotic chromosome pairings of the
parents were normal and no univalents or multiva—
lents were observed (Fig．
28 chromosomes．A large
found in this hybrid with
2．A—B)．The hybrid has
number of univalents were
2O．40 (from 14-26) per
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354 广 西 植 物 22卷
cell on average．The bivalents were 3．69 (from1—7)
per cel1． Trivalents and quadrivalents were infre-
quent，around 0．09 and 0．04 per cell，respectively．
Of 57 F，PM Cs examined，25 (43．76 )showed 3
to 4 bivalents(Fig． 2．C)． The chiasma frequency
was 4．51 with the range of 1-9 and the C—value was
0．1 6．Chromosome bridges and the lagging chromo—
somes at anaphase 1 were found (Fig．2．D—E)，and
micronuclei were also observed in tetrads of F1 hybrid
(Fig．2．F)．
Table 1 Comparison of morphological characters among R．grandis，R．alashanica and their Fl hybrid
2 3 A 2 3 B
Fig．1 Spikes of R．alashanica，R．grand#and the Fl hybrid A．Spikes；B．Spikes，glumes．1emmas and paleas
1．R．alashanica；2．R．grandis× R．alashanica：3．R．gt‘andi5．
Table 2 Chromosome pairing at MI of PMCs in R．grandis，R．alashanica and their Fl hybrid
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4．Fertil ity
T6e pollen fertility and seed set of the parental
species and Fl hybrid were examined (Table 3)·
Most pollen grains of the parental species were vi—
able，while those of Fl hybrid were not．The seed set
of the parental plants were 83．33 and 91·18 ，
respectively．But no seed was formed in Fl hybrid af-
ter being carefully examined a great number of flo—
rets．
Discussion
Interspecific hybridization between R． grandis
and R． alashan／ca in this study was comparatively
easy to perform without the assistance of embryo res-
cue technique， indicating that crossing barriers are
not very strong be tween these two species．However，
polens of Fl hybrid were completely sterile while
those of the parental species were quite norma1．
Based on the morphology and hybridization data，it is
evident that R． grand~ and R． alashanica are two
dependent biological species．
The degree of chromosome pairing in an inter-
specific or intergeneric hybrid can be used as an index
of chromosome homology as well as specific or generic
evolutionary relationship ． Homologous chromo-
somes are defined as having the same nucleotide base
sequences，whereas homoeologous chromosomes have
only residual homology from the originally homolo-
gous chromosomes ”． In the meiosis of Fl hybrid
from the cross of R．grandis and R．alashan／ca，the
number of chromosome pairing varied from 1-7，
showed an average of 3．69 bivalents per cel1． This
suggests that R．alashan／ca has one genome which is
homoeologous to one genome of R． grandis． St
genome is the basic genome in Roegneria，Elymus
and Elytrigia， originating from the genus Psedo-
roeg eria，while Y is mostly found in Asian species
of Roegneria，which come from an unknown diploid
progenitor． Zhang observed that the hybrid of R·
口 ， ／c口and R．ciliaris(Trin．)Nevski had a high—
er amount of chromosome pairing，up to 8．1 7 biva—
lents per cel and drew conclusion that R．alashan／ca
shared at least one set of St genome of R． ciliaris．
Therefore，we could conclude that R．alashanica has
one slightly modified St genome， be designated as
St‘．
I，u made genome analysis of R．el3 trigioides，
and concluded that R． elytrigioides contains two
slightly modified St genomes， StlSt1St2St2． Cai
treated morphologicaly R． elytrighddes as a variety
6f R．日 s^口 ica．Therefore，R．alashanica is related
closely to R． elytrigioMes． The genome of R．
口 ha ica is in our further persuit．If R．alashanica
has two St genomes as R．elytrigioides，according to
the point of view of Dewey[8]1．6ve 。 and Baum㈩ ，
it should be transferred to Psedoroegner&．
References
C13 Keng Y I，，Chen S I，．A revision of the genus Roegne—
r C．Koch of China EJI．Nanjing Univ Acad Rep，
1963，1：15— 92．
C23 Zhang X Q，Yang J L，Yen C．The genome constitu
tion of Roegneria grandis(Poaceae，Triticeae)UI．Pl
Syst Evol，1998，209：67— 73·
C33 Zhang X Q，Yen C，Yang Y I ·et a1．Cytological
studies of hybrids of Roegneria ciliaris with R．alashan—
ica and R．magnicaespes(Poaceae：Triticeae)J-J1．Ac—
ta Bot Yunnanica，2000，22(2)：155— 160．
(43 Cai I B．A taxonomical study on the genus Roegneria
C．Koch from China r-j-1．Acta Phytota Sinica，1997，
35(2)：148—177．
C53 I u B R．Meiosis analysis of the intergeneric hybrids
between Psedoroegneria and tetraploid Elymus[J]．
维普资讯 http://www.cqvip.com
356 广 西 植 物 22卷
Cathaya，1994，6：1——14．
(63 Sakamoto S，Muramatsu M．Cytogeneric studies in the
tribe Triticeae．II．Tetraploid and hexaploid hybrids of
Agropyron[J]．Jap J genet，1966，41：155—168．
[7]Kimber G．Genome analysis in the genus Triticum
[C]．In：Sakamoto S (ed)：Proceedings of the 6 In—
ternational W,heat Genetics Symposium ，Kyoto：Kyoto
University Press，1 983．23——28．
[8]Dewey D R．The genome system of classification as a
guide to intergeneric hybridization with the perennial
Triticeae[M]．In：Gustafson J P(ed．)：Gene Manip—
ulation in Plant Improvement．Proc 1 6th Stadler genet
symp．New York：Plenum Publ Corp，1 984． 209—
280．
[93 L6ve A．Generic evaluation of the wheatgrass[-J~．Boil
ZentealbZ，1982，101：199— 212．
[10]L6ve A．Conspectus of the Triticeae[J]．Feddes
Repertotium ，1984，95：425——521．
[11]Baum B R，Yen C，Yang Y I ．Roegneria：its generic
limits and justification for its recognition[J]．Can J
t．】99】，69：282— 294．
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A．期刊 序号 著者．题 目．刊名，出版年份．卷号(期号)：起止页码
[13周世良，潘开业．洪德元．等．杭州石荠荣(唇形科)小坚果的散布生物学研究[J]．广西植物．1999，19(2)．176一l79．
[23 Pasqua G，Monaceli B．Cuteri A．et n ．Cell suspension cultures of Maclura pomifera：Optimization of growth and metabolite production
[J]．．，Plant Physio1．1991．139：249-251．
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[1]吴征镒主编．新华本草纲要 (第一卷)[M]．上海：上海科枝出版社，l998．14l—l53．
[23 Guowen Xie．Protection and management of mountain forests[M]．Beijing：Science Press．1992．256—260．
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