全 文 ：Agronomic Characters of Elymus dahuricus ,
Hordeum brevisubulatum and Their BC1F3
LI Wei-wei1 , LI Zao-zhe1＊ , WANG Lei2
1.College of Ecology and Environmental Science , Inner Mongolia Agricultural University , Hohhot 010019;2.Keshan Branch , Heilongjiang Academy of
Agricultural Sciences, Keshan 161600
Abstract [Objective] This study was to explore the inheritance of BC1F3 to excellent characteristicsof parents(Elymusdahuricus , Hordeum brevisubulatum),
heterosis and fert ility restoration.[ Method] The agronomic characters such as growth period , growth rate , pollen fertility, fecundity , reproducibility and fresh
grass output of Elymusdahuricus , Hordeumbrevisubulatum and BC1F3were comparatively studied.[ Result] The growth dynamic of theBC1F3 linedtended to like
that of its parent Hordeumbrevisubulatum;although growing periodsof various lineswere different , they were close to that of Hordeumbrevisubulatum.There
were differences in pollen fert ility and seed setting , the pollen fertility rate of YF3-93 was higher than recurrent parent Hordeumbrevisubulatum , its seed sett ing
rate in open pollinat ionwasalso higher.Some lines had lowpollenfert ility rate and seed setting rate , such as the pollen fertility rate of PF3-52.Therewere signif-
icant differences in the output among BC1F3 lines, e.g.the heterosis(HP)of YF3-64 , YF3-74 and YF3-83 were 75.53%, 75.12%and 66.16%, respect ively;
however , the yieldsof PF3-52, PF3-15 and PF3-42were lower than their parents.[ Conclusion] Thisstudy provided reference for breeding new varietiesof forage
which is suitable to drought and saline environment.
Keywords Elymus dahuricus;Hordeumbrevisubulatum;Backcross posterity;Agronomic characters
Received:February 11 , 2009 Accepted:April 27 , 2009
Supported by Special Fund fromGrass Industry Research Institute aff ilia-
ted to Department of Science and Technology of Inner Mongolia of China
(20071923);the Startup Foundation of Scient if ic Research for Returned
Scholars of China.
＊Corresponding author.E-mail:zaozheli@hotmail.com
Elymus dahuricus and Hordeum brevisubulatum are perennial
Triticeae grasseswith important economic value and ecological val-
ue.Elymus dahuricus has the properties of high output and strong
drought tolerance;while Hordeumbrevisubulatum has the properties
like strongtillering abil ity , good grass qual ity andtolerance to salt.In
order to combine the properties of these two forages for breedingno-
vel forage variety(novel type)that can adapt to drought and sal ine
environment , WANG Zhao-lan et al carried out the wide hybrid be-
tween Elymus dahuricus and Hordeumbrevisubulatum during 1992-
1993 , and successfully obtained the highly sterile F1 generations of
reciprocal cross between Elymus dahuricus andHordeumbrevisubu-
latum.On the basisof preliminary study onthemorphology andcytol-
ogy of rare and limited obversecross of F1 generation(Elymus dahur-
icus ×Hordeum brevisubulatum)[ 1-2] , we planed to overcome the
sterility of F1 hybrid by back cross.Using the plants propagated by
division , we propagated the F1 hybrids of Elymus dahuricus ×Hor-
deum brevisubulatum and Hordeumbrevisubulatum×Elymus dahuri-
cus to independent l ines , and successfully obtainedthe F1 back cross
generations of(Elymus dahuricus and Hordeum brevisubulatum)×
Hordeum brevisubulatum and (Hordeum brevisubulatum ×Elymus
dahuricus)×Hordeum brevisubulatum by adjacent plantation of F1
generation and parents , and back cross using bagging[ 2] .In order to
reveal the heredity , heterosis and fertility recovery of BC1F3 inherited
from parents , we principally compared the agronomic traits of BC1F3
including growthperiod , growthrate , pollenfertility , seedsetting rate ,
regeneration and freshgrass output per clump , aiming at providing ref-
erence for breeding new varieties[ 3] .
Materials and Methods
Experimental materials
Elymus dahuricus , Hordeumbrevisubulatum , PF3-15 , PF3-52 ,
YF3-23 , YF3-31 , YF3-42 , YF3-43 , YF3-44 , YF3-53 , YF3-64 , YF3-
74 , YF3-75 , YF3-83 , YF3-85 andYF3-93 , ofwhich Prepresents the
F1 back cross generations of(Elymus dahuricus and Hordeum bre-
visubulatum)×Hordeum brevisubulatum and , Y represents the F1
back cross generations of(Hordeumbrevisubulatum×Elymus dahur-
icus)×Hordeumbrevisubulatum , F3 represents F3 generation from
open pollinationof F1 back cross generation.
General situationof experimental plot
The experimental materials were planted in the Forage Experi-
mental Farmof Inner Mongolia Agricultural University at the eastern
outskirt of Hohhot City , where the geographic coordinate is 110°E
and 45°5′N , altitude is 1 063 m , yearly precipitation is about 400
mm , nonfrost period is 145 d.The soil texture is loamydark chestnut
soil with pH value of 7.8-8.2 and medium fertility;the water table
is relatively high.Thus these are conditions for irrigation.
Experimental methods
Observation of growth period , pollen fertility and seed setting
characteristics Growthperiod:using theoverwintered clump prop-
agated by division in the last year as experimental objective , the
growth characteristics from returning green stage in early spring to
withering stage were recorded in detail.Pollen fertility:ten spikes of
each tested materials were randomly sampled at full bloom stage ,
placed into moisturemaintaining boxes and taken back to laboratory for
microscopic examination;anthersweretakenout and placedontoglass
sl ides , followed by dripping a l ittle 1%acetumcarmine;then anther
walls were removed by squeezing using a forceps to make the pollens
adequately stained before covering the microscopic glass;the slides
were examinedunder microscope to record the numbers of fertile pol-
lens and sterile pollens , each slide(material)contained more than 50
visual fields.The criterion to determine the pollen fertil ity was as the
Agricultural Science &Technology , 2009 , 10(2):15-19
Copyright 2009 , Information Inst itute of HAAS.All rights reserved. Agronomy
follows:the pollenswith big size , plump and deeply staining were re-
garded asfertile , that withsmall size , slightly or nostainingwas regar-
ded as sterile.Rate of fertile pollen=(total number of fertile pollens/
total number of pollens observed)×100%[3-6] .
Seed setting rate:thirty spikeswere sampled fromeach tested
material at mature stage to record the seed setting number and flo-
rets.Seed setting rate=(total number of seeds/ total number of flow-
erlets observed)×100%[ 3-6] .
Determinationof growing rate andgrassyield These two indi-
ces were determined by fixed clump observation.Ten well growing
individual clumps from each material after returning green in spring
werenumbered and labeled as experimental objective.
Growing rate:absolute heights of individual clumps tested were
measured every ten days from returning green stage to maturity of
each experimental material.
Grass output:yearly average fresh grass output was shown by
the sumof fresh grass outputs of two cuttings and used to calculate
average heterosis(HP), HP=(F3 -average of parents)/ average of
parents×100%[7-9] .
Results and Analysis
Growing period of parents and the backcrossed hybrid proge-
nies
As shown in Table 1 , parents of Hordeum brevisubulatum re-
turned green 10 days earlier and withered 3 days later than Elymus
dahuricus.The rhythm of Hordeum brevisubulatum grew relatively
quick , and flowered and set seed earlier , its growth period was 13
days longer than Elymus dahuricus.Thereturninggreenstage of each
line of BC1F3 generation was basically identical;the growing period
varied frommaximum214 d to minimum197 d , but tendedto the re-
current parent Hordeum brevisubulatum.The growth and develop-
mental rhythm of BC1F3 generation approximates to Hordeum bre-
visubulatum.These results suggested that the characteristics of quick
growth and developmental rhythm in Hordeum brevisubulatum was
enhanced inBC1F3 generation via back cross.
Table 1 Growth characteristics of tested plants during tested period
Grasses Returning green Tillering Jointing Booting Earing Flowering Maturing Withering Growing period∥d
Elymusdahuricus 03-30 04-27 06-16 06-22 06-30 07-04 08-01 10-12 196
Hordeumbrevisubulatum 03-20 04-24 04-27 05-06 05-14 05-26 06-24 10-15 209
PF3-15 03-20 04-25 04-29 05-10 05-22 05-30 06-30 10-03 197
PF3-52 03-20 04-25 04-29 05-12 05-24 05-30 07-04 10-06 200
YF3-23 03-20 04-24 04-28 05-10 05-22 05-30 07-02 10-22 216
YF3-31 03-20 04-25 04-28 05-10 05-26 06-02 06-30 10-12 206
YF3-42 03-20 04-23 04-27 05-12 05-24 05-30 07-02 10-12 206
YF3-43 03-22 04-24 04-28 05-10 05-22 05-28 06-30 10-12 204
YF3-44 03-22 04-23 04-26 05-08 05-24 06-02 07-04 10-14 206
YF3-53 03-22 04-25 04-29 05-10 05-24 05-28 07-04 10-18 210
YF3-64 03-22 04-24 04-27 05-08 05-24 05-28 07-02 10-22 214
YF3-74 03-22 04-23 04-27 05-08 05-22 05-28 07-02 10-24 214
YF3-75 03-22 04-23 04-28 05-08 05-22 05-28 07-02 10-24 214
YF3-83 03-22 04-25 04-29 05-08 05-22 05-28 06-30 10-22 210
YF3-85 03-22 04-24 04-28 05-10 05-20 05-28 06-30 10-18 210
YF3-93 03-22 04-25 04-28 05-08 05-18 05-26 06-30 10-22 214
Growth rate of parents and the backcrossed hybrid progenies
Both BC1F3 generation and its parents presenteda growing tend
of slowing before jointing-then quickening-slowing down again after
flowering(Scurve)as indicated in Fig.1.Thegrowthand devel-
opmental rhythm of Hordeum brevisubulatum was relatively earl ier
and growth rate was relatively quicker.Most of the BC1F3 lines
showed an identical rhythmwith Hordeum brevisubulatum , of which
six lines including YF3-64 , YF3-74 ,YF3-75 ,YF3-83 ,YF3-85 and YF3-
93 grew slightly quicker than Hordeumbrevisubulatum after June 3 ,
other lines grew slowly , indicating that the growing rate of BC1F3
generation tends to inherit the characteristic of its concurrent parents
Hordeumbrevisubulatum.
Regenerat ion rate of parents and the Backcrossed hybrid
progenies
It could be concluded from Fig.2 that the regeneration rate of
Elymus dahuricus was higher than that of Hordeum brevisubulatum ,
BC1F3 generation regenerated with a similar rate with parent Hor-
deumbrevisubulatum , of whichYF3-74 andYF3-93 regeneratedquic-
ker than Hordeum brevisubulatum , but most lines didnt.There are
differences in regeneration rate among different lines.
Fig.1 The growth rate of tested plants
Pollen fertility and seedsetting characteristics of parents and
the backcrossed hybrid progenies
As shown in Table 2 , the pollenfertility and seedsetting in both
16 Agricultural Science &Technology Vol.10 , No.2 , 2009
the Elymus dahuricus and Hordeum brevisubulatum were relatively
high(90.45%and 92.96% respectively), their natural seed setting
rates under open pollination respectively reached 63.69% and 52.
42%.The pollen fertility and seed setting rate in BC1F3 generation
recoveredand increased todifferent extent , but therewere significant
differences among different l ines , e.g.rates of fertile pollen in PF3-
52 and YF3-44 were 81.50%and85.61%respectively , thenatural
seed setting rates under openpollinationwere 12.88%and 19.90%
respectively;rates of fertile pollen in YF3-64 and YF3-93 were 92.
79%and 93.91% respectively , the natural seed setting rates under
open poll ination were 37.52%and 56.70% respectively;rates of
fertile pollen in other lines reached 86.50%-95.48%, approxima-
ting to that of the parents , the natural seed setting rates under open
pollination were between 24.27% and 37.20%, suggesting that
pollen fertil ity and seed setting rate in BC1F3 generation were both
recovered and enhanced. Fig.2 The regeneration rate of tested plants
Table 2 Pollen fert ility and seed setting character of tested plants(average , Duncan test)
Grasses
Pollen fertility
Fert ility No. Total Fertility rate∥%
Seed setting character
Seed No. Total florets Seed setting rate∥%
Elymusdahuricus 2 699 2 984 90.45 3 148 4 943 63.69
Hordeumbrevisubulatum 2 879 3 097 92.96 2 274 4 338 52.42
PF3-15 3 000 3 219 93.2 1 068 3 907 27.34
PF3-52 2 458 3 016 81.50 445 3 454 12.88
YF3-23 2 797 3 048 91.77 1 263 3 981 31.73
YF3-31 2 434 2 652 91.78 1 035 4 110 25.18
YF3-42 2 842 3 141 90.48 987 3 910 25.24
YF3-43 3 454 3 780 91.38 925 3 812 24.27
YF3-44 2 963 3 461 85.61 699 3 513 19.90
YF3-53 2 063 2 385 86.50 1 129 4 014 28.13
YF3-64 3 345 3 605 92.79 1 806 4 814 37.52
YF3-74 3 324 3 535 94.03 1 226 3 674 33.37
YF3-75 3 804 4 025 94.51 1 241 3 336 37.20
YF3-83 3 486 3 651 95.48 1 247 4 055 30.75
YF3-85 3 085 3 311 93.17 1 594 4 532 35.17
YF3-93 2 774 2 954 93.91 2 158 3 806 56.70
Table 3 The output performance of tested plants(average, Duncan test)
Grasses
First
cutt ing
kg/per clump
Second
cutting
kg/ per clump
Fresh forage
output per clump
kg
Average
heterosis
%
Elymus
dahuricus
336.0 cd 45.4 cd 381.4 cd -
Hordeum
brevisubulatum
354.6 c 47.8 cd 402.4 cd -
PF3-15 308.3 cd 42.9 cd 351.2 cd -
PF3-52 188.0 d 38.0 d 226.0 d -
YF3-23 401.0 bc 64.5 bcd 465.5 bc 18.78
YF3-31 324.6 cd 47.3 cd 371.9 cd -
YF3-42 296.5 cd 45.1 cd 341.6 cd -
YF3-43 333.6 cd 73.6 bc 407.2 cd 3.90
YF3-44 359.6 c 66.1 bcd 425.7 c 8.62
YF3-53 354.5 c 36.8 d 391.3 cd -
YF3-64 597.4 a 90.5 ab 687.9 a 75.53
YF3-74 572.1 a 114.2 a 686.3 a 75.12
YF3-75 533.7 ab 114.0 a 647.7 a 65.27
YF3-83 569.2 a 82.0 b 651.2 a 66.16
YF3-85 387.5 bc 48.8 cd 436.3 c 11.33
YF3-93 535.8 ab 75.2 bc 611.0 ab 55.91
Values in eachcolumn followed by lowercases are significantly different(P<0.
05).
Output character of parentsand the backcrossedhybrid prog-
enies
FromTable 4 we can see that the output of fresh grass of Ely-
musdahuricus and Hordeumbrevisubulatum are equivalent , both are
dominantly contributed by the first cutting and slightly by the second
one.Elymus dahuricus plants are strong and tall , while Hordeum
brevisubulatum has a vigorous tillering ability , so their outputs of
fresh grass are equivalent.
Fourteen lines of BC1F3 generation were significantly different in
grass output , e.g.YF3-64 ,YF3-74 ,YF3-75 , YF3-83 ,YF3-93 showed
higher grass outputs than parents , whose HPs were respectively 75.
53%, 75.12%, 65.27%, 66.16%and 55.91%, showing a re-
markable heterosis;some others l ike PF3-52 , PF3-15 and YF3-42
performed poorer inoutput than parents , their grass outputswere re-
spectively 226.0 , 351.2 and 341.6 g/clump.
Conclusion
(1)All the lines of BC1F3 generation tend to inherit the growth
rhythmof their recurrent parent Hordeumbrevisubulatum , though the
growing periods of various l ines are different.
(2)There are differences in pollen fertility and seed setting
rate among different lines of BC1F3 generation , but the fertil ity of
each l ine increased to some extent after back cross , e.g.the
pollen fertility of YF3-93 exceeded its parent Hordeumbrevisubu-
17LI Wei-wei et al.Agronomic Characters of Elymus dahuricus , Hordeumbrevisubulatum and Their BC1F3
latum , its rate of fertile pollen was 93.91%andnatural seed set-
ting rate under open pollination reached 56.70%;while some
lines did not , e.g.rate of fertile pollenPF3-52 was 81.50%and
natural seed setting rate under open poll ination was just 12.
88%.
(3)There was large variation in grass output among BC1F3
lines e.g.the heterosis of YF3-64 , YF3-74 and YF3-83 were re-
spectively 75.53%, 75.12%and 66.16%, while thegrass out-
puts of PF3-52 , PF3-15 andYF3-42 were inferior to their parents.
References
[ 1] LI ZZ, YUNJF, YUZ, et al.Morphological study onElymus dahuricus ,
Hordeumbrevisubulatum andtheir F1 hybrid and BC1[ J] .Grassland of Chi-
na, 2002, 24(5):24-28.(in Chinese).
[ 2] LI ZZ, YUNJF, YUZ, et al.Studieson cytogenet icsof Elymus dahuricus ,
Hordeumbrevisubulatum and their Hybrid F1 and BC1[ J] .Acta Agrestia
Sinica , 2005, 13(4):269-273.(in Chinese).
[ 3] YUZ, LI ZZ, YUNJF.Studyon the agronomic charactersof several peren-
nial species in Triticeae and their hybrids[ J] .Acta Prataculturae Sinica,
2003, 6(3):83-89.(in Chinese).
[ 4] YUZ, YAMADATOSHIHIKO.Comparat ive researchonmaincharacteristics
of pacetter variet ies[ J] .Chinese Journal of Grassland , 2006, 10(6):1-
6.(in Chinese).
[ 5] LI ZZ, YUZ, MAQZ, et al.Studiesonthe biological and agronomicchar-
actersof Elymus dahuricus , Hordeum brevisubulatum and their hybrid F1
and BC1F1[ J] .Grassland of China , 2004(9):9-14.(in Chinese).
[ 6] LI XL , YUZ, MAYH, et al.Study on the growth, development and cyto-
geneticsetc.of F1 Hybrid derived fromElymus sibiricus andE.purpura-
ristatus[ J] .Journal of Triticeae Crops, 2006, 26(2):37-41.(in Chi-
nese).
[ 7] WANG SY, YUNJF, XUJ, et al.Growth andmorphology characters of
Elymus canadensis , Elymus sibiricus and their hybrid[ J] .Acta Agrestia
Sinica , 2004, 12(4):294-297.(in Chinese).
[ 8] MAYB, RENWB , LI LH.Evaluat iononagronomiccharactersofthreeEly-
mus species[ J] .Animal Husbandry and Feed Science, 2008(3):29-31.
(in Chinese).[ 9] ZHANQW, LINP, LI J, et al.Research and prospect of hybrid between
sorghum(Sorghum bicolor)and smut (Sorghum sudanense)[ J] .Acta
Prataculturae Sinica , 2001 , 10(2):56-61.(in Chinese).
[ 10] WUNE, LI ZZ, ZHAOH, et al.Analysis of peroxidase isozymes and es-
terase isozymesof Elymus dahuricus , Hordeum brevisubulatum and their
hybrid F1 and BC1F1[ J] .Chinese Journal of Grassland , 2007 , 29(3):28-33.(in Chinese).
[ 11] LIUZH, DONGSW , XIONGY , etal.Variat ion of agronomiccharactersof
F2 of predominance cotton combination and its ut ilization evaluation[ J] .
Agricultural Science &Technology , 2007, 8(3-4):38-41.
[ 12] LI JH, YUNJF, YUSY, etal.Studyof esterase isoymesmarker onpurity
identification among Canadian Elymuscandensis, E.sibiricus and their hy-
brid progeny[ J] .Seed , 2007 , 26(11):75-76.(in Chinese).
[ 13] LIXL, YUZ, MAYH, et al.Studieson the tissue cultureand regeneration
systemof hybrid F1 of Elymus canadensis×E.excelsus[ J] .Journal of
Anhui Agricultural Sciences, 2008 , 36(6):2235-2237.(in Chinese).
[ 14] LI Z.Studies on chromosome stability and agronomic traits of the back-
crossdescendants betweenElymusdahuricus andHordeumbrevisubulatum[D] .Hohhot:Inner Mongolia Agricultural University , 2006.(in Chinese).
[ 15] LI SJ.Studies agronomic traits and genet ic diversity on wild germplasm
resourses of Elymus L.[ D] .Xining:Qinghai University, 2007.(in Chi-
nese).
Responsible editor:ZHENG Dan-dan Responsible translator:DUAN Yong-bo Responsible proofreader:WU Xiao-yan
披碱草和野大麦及其杂种回交后代的农艺特性研究
李微微1 ,李造哲1＊ ,王 磊2
(1.内蒙古农业大学生态环境学院 ,内蒙古呼和浩特 010019;2.黑龙江省农业科学院克山分院 ,黑龙江齐齐哈尔 161600)
为了使披碱草和野大麦的优良特性综合在一起 ,培育出适
合干旱 、盐渍生境的牧草新品种 ,王照兰等(1992 ～ 1993年)将披
碱草和野大麦组合进行了远缘杂交 ,成功地获得了高度不育的
披碱草和野大麦的正 、反交 F1 代 。在正交 F1 代(披碱草×野大
麦)的植株形态学和细胞学初步研究的基础上 ,笔者拟定了用回
交法克服杂种 F1 代不育性的研究计划 。用分株繁殖方法 ,分别
将披碱草×野大麦和野大麦×披碱草 F1 代扩繁成株系群体 ,并
采取 F1 代与亲本相邻种植 、套袋回交方法 ,成功地获得了(披碱
草×野大麦)×野大麦和(野大麦 ×披碱草)×野大麦的回交一
代。为了探明 BC1F3 代对亲本优良特性的继承情况 、杂种优势
及育性恢复情况 ,笔者重点对其农艺特性 ,包括生育期、生长速
度、花粉育性 、结实性 、再生性和株丛鲜草产量等进行了系统的
比较研究 ,以期为新品种的培育提供依据。
1 材料与方法
1.1 材料 试材包括:披碱草 、野大麦 、PF3-15 、PF3-52 、YF3-
23 、YF3-31 、YF3-42 、YF3-43 、YF3-44 、YF3-53 、YF3-64 、YF3-74 、
YF3-75 、YF3-83 、YF3-85 、YF3-93 , 其中 , Y 表示(野大麦 ×披碱
草)×野大麦的回交 1代;P表示(披碱草×野大麦)×野大麦的回
交1代;F3表示回交 1代的开放授粉第 3代。
1.2 试验地概况 试材种植于内蒙古农业大学牧草试验场内 ,
该试验场位于呼和浩特市东郊 ,地理坐标为 110°E ,45°5′N ,海拔
1 063m ,年降水量 400 mm ,无霜期145 d。土壤为沙壤质暗栗钙
土 ,pH值 7.8 ～ 8.2 ,肥力适中 ,地下水位较高 ,具有灌溉条件。
1.3 方法
1.3.1 生育期 、花粉育性及结实性观察。生育期:以前一年分
株繁殖的越冬株丛为对象 ,从早春返青期至枯黄期进行详细观
察记载 。花粉育性:在开花盛期随机剪取每种植物穗子 10个 ,
放入塑料保湿盒中带回室内镜检 。剥取花药 ,置载玻片上 ,滴加
少许1%醋酸洋红 ,用镊子挤压去掉药壁 ,使花粉充分着色 ,加盖
玻片 ,在100倍显微镜下观察统计可育与不育花粉粒数 ,每种材
料观察 50个视野以上 。确定花粉是否可育的标准是:花粉粒
大 、饱满、着色深者为可育 ,花粉瘦小 、着色浅或不着色者为不
育 。花粉可育率 =(可育花粉粒总数/观察花粉粒总数)×
100%。结实性:在种子成熟期 ,从各供试材料群体中随机取 30
个穗子 ,统计结实数和小花数。结实率=(结实总数/观察小花
总数)×100%。
1.3.2 生长速度 、产草量测定 。采取定株方法 。春季牧草返青
后自每种供试材料群体中选择长势良好的 10个株丛编号挂牌 。
生长速度:从返青期开始 ,每隔 10 d测量 1次株丛绝对高度 ,直
至每种供试材料成熟。产草量:用 2次刈割鲜草重量之和表示
18 Agricultural Science &Technology Vol.10 , No.2 , 2009
一年内每种供试材料的平均鲜草产量 ,并以此计算杂种平均优
势(HP),HP=(F3-双亲平均值)/双亲平均值×100%。
2 结果与分析
2.1 亲本及其杂种回交后代的生育期 亲本野大麦比披碱草
返青早 10 d ,而枯黄晚 3 d。野大麦的生长发育节律较快 ,较早
地开花结实 ,其生长天数比披碱草多 13 d。BC1 F3 代各株系的
返青期基本一致 ,不同株系的生长期都有所不同 ,最长的为 214
d ,最短的为 197 d ,但都趋于轮回亲本野大麦 。BC1 F3 代的生长
发育节律与野大麦的较接近 ,说明通过回交使野大麦生长发育
节律快的特性在 BC1F3 代中进一步得到了加强(表见第 16页
Table 1:供试植物生育期观测结果)。
2.2 亲本及其杂种回交后代的生长速度 BC1 F3 代和亲本的
生长动态呈“慢、快、慢”的缓“S型”曲线 ,拔节前生长缓慢 ,之后
生长速度加快 ,开花后生长速度变缓(图见第 16页 Fig .1:供试
植物生长速度)。野大麦的生长发育节律较早 ,生长速度较快 。
回交后代大部分株系与野大麦的生长节律一致 ,其中 YF3-64 、
YF3-74 、YF3-75 、YF3-83 、YF3-85 、YF3-93六个株系在 6月 3日
后生长速度稍快于野大麦 ,其他株系的生长速度稍慢 , 表明
BC1 F3代的生长速度呈偏向轮回亲本野大麦遗传的倾向 。
2.3 亲本及其杂种回交后代的再生速度 亲本披碱草的再生
速度比野大麦的速度快 , BC1 F3 代的再生速度与亲本野大麦的
速度相近 ,其中 YF3-74 、YF3-93较野大麦生长快 ,大部分株系的
再生速度较亲本野大麦慢 ,不同株系的再生速度有较大的差异
(图见第 17页 Fig.2:供试植物再生速度)。
2.4 亲本及其杂种回交后代的花粉育性及结实性 亲本披碱
草和野大麦的花粉育性较强 , BC1 F3 代的花粉育性和结实性有
不同程度地恢复和提高 ,但各株系间的花粉育性和结实性仍有
较大差异(表见第 17页 Table 2:供试植物花粉育性和结实性观
测结果)。
2.5 亲本及其杂种回交后代的产量特性 亲本披碱草和野大
麦鲜草产量相当 ,都是以第 1次刈割产草量占绝对优势 ,第 2次
刈割产草量较少 。披碱草植株粗壮、高大 ,而野大麦分蘖能力
强 ,两亲本各具优势 ,故鲜草产量相当。BC1 F3 代 14个株系的
产草量存在着显著差异 , 如 YF3-64 、YF3-74 、YF3-75 、YF3-83 、
YF3-93的产草量较亲本高 ,表现出明显的杂种优势 ,其 HP分
别为 75.53%、75.12%、65.27%、66.16%、55.91%;也有产量较
低的 ,如 PF3-52 、PF3-15 、YF3-42产草量不及亲本 ,其产草量分别
为 226.0 、351.2和 341.6 kg/株丛 。
表3 供试植物产量测定结果(均值 , Duncan test)
材料 1次刈割
kg/株丛
2次刈割
kg/株丛
鲜草量
kg/株丛
平均杂种
优势∥%
披碱草 336.0 cd 45.4 cd 381.4 cd -
野大麦 354.6 c 47.8 cd 402.4 cd -
PF3-15 308.3 cd 42.9 cd 351.2 cd -
PF3-52 188.0 d 38.0 d 226.0 d -
YF3-23 401.0 bc 64.5 bcd 465.5 bc 18.78
YF3-31 324.6 cd 47.3 cd 371.9 cd -
YF3-42 296.5 cd 45.1 cd 341.6 cd -
YF3-43 333.6 cd 73.6 bc 407.2 cd 3.90
YF3-44 359.6 c 66.1 bcd 425.7 c 8.62
YF3-53 354.5 c 36.8 d 391.3 cd -
YF3-64 597.4 a 90.5 ab 687.9 a 75.53
YF3-74 572.1 a 114.2 a 686.3 a 75.12
YF3-75 533.7 ab 114.0 a 647.7 a 65.27
YF3-83 569.2 a 82.0 b 651.2 a 66.16
YF3-85 387.5 bc 48.8 cd 436.3 c 11.33
YF3-93 535.8 ab 75.2 bc 611.0 ab 55.91
注:纵列小写英文字母不同表示0.05水平存在差异显著性。
3 结论
(1)BC1 F3 代各株系的生长节律趋向于亲本野大麦 ,不同株
系的生育天数有所不同 ,但都接近轮回亲本野大麦。
(2)BC1 F3 代不同株系花粉育性和结实性有较大差异 ,但回
交后各株系的花粉育性都有一定程度的提高 ,如 YF3-93的花粉
育性已经超过亲本野大麦 ,花粉可育率达到 93.91%,自然结实
率也较高 ,达 56.70%。而有的株系花粉育性和开放授粉结实率
较低 ,如PF3-52的花粉育性为 81.50%,自然结实率为 12.88%。
(3)BC1 F3 代各株系产草量有较大变异 ,不同株系间存在明
显的差异 ,如 YF3-64 、YF3-74 和 YF3-83 的杂种优势分别为
75.53%、75.12%和 66.16%,而 PF3-52 、PF3-15和 YF3-42产草
量不及亲本。
基金项目 内蒙古科技厅草业研究院专项(20071923),国家留学回国人
员科研启动基金项目。
作者简介 李微微(1981-),女 ,内蒙古根河人 ,硕士研究生 ,研究方向:
野生植物资源保护与利用。 ＊通讯作者。
收稿日期 2009-02-11 修回日期 2009-04-27
(上接第 8页)
分离采用不连续系统聚丙烯酰胺凝胶垂直平板电泳法 ,浓缩胶
浓度为 5%,分离胶浓度为 7.5%。AMY 同工酶染色参照顾玉
红报道的方法;SOD 同工酶染色参照何忠效的方法 。染色后
在凝胶成像系统(北京六一电泳仪器厂 ,WD4031 型)上观察并
拍照。
2 结果与分析
2.1 香果树种子萌发过程中淀粉酶同工酶的变化 香果树种
子萌发过程中 AMY 同工酶酶谱如图 1 所示(图见第 7 页
Fig .1),共有 3条酶带 ,按分子量由小到大依次为 A 1 、A 2 、A 3 。
香果树种子萌发过程中第 1 ～ 4天有 2条酶带 A 1和 A 3 ,从第4
天开始出现新的酶带 A2 。酶带 A1和 A 3 在各时期颜色均较
深 ,为香果树种子萌发过程中的主酶带 ,表明酶带 A1和 A 3 的
基因表达贯穿于香果树种子萌发全过程 ,是香果树种子萌发过
程中的必需酶带 。
2.2 香果树种子萌发过程中超氧化物歧化酶同工酶的变化 香
果树种子萌发过程中 SOD同工酶酶谱如图 2(图见第 7页 Fig.2),
共有 6条酶带 ,按照分子量由小到大的顺序依次为 S1 、S2 、S3 、S4 、
S5 、S6 。在香果树种子的萌发过程中 ,SOD同工酶酶带在第 1 ～ 2
天颜色最深 ,第 3 ～ 5天逐渐减弱 ,第 6 ～ 9天活性又加深 。酶带
S1 、S4和 S5在香果树种子萌发各个时期均出现 ,酶带颜色较深 ,
为香果树种子萌发过程中的特征酶带 ,表明酶带 S1 、S4 和 S5 的
基因表达贯穿于香果树种子萌发过程的各个时期 ,为香果树种
子萌发过程中的必需酶带 。酶带 S2 和 S3 第 1 ～ 2天颜色较深 ,
从第3天开始颜色变淡 ,酶带开始消失 ,从第 6天开始酶带 S2
又开始出现。酶带 S6 在香果树种子萌发各个时期均出现 ,只是
在萌发中后期活性明显增强 ,说明其控制的代谢在萌发中后期
明显增强 。结果表明 ,在香果树树种子萌发初期 ,有新的 SOD
同工酶合成。进入萌发中期 ,新合成的第 2 、3条酶带消失 ,在萌
发后期第 2 条酶带又重新出现;萌发中后期第 6条酶带活性
增强。
基金项目 三峡大学博士启动基金和国家自然科学基金项目
(30670202)资助。
作者简介 刘国勇(1972-),男 ,湖北汉川人, 博士, 讲师 ,从事动植物
生理生态研究。 ＊通讯作者。
收稿日期 2009-03-11 修回日期 2009-04-21
19LI Wei-wei et al.Agronomic Characters of Elymus dahuricus , Hordeumbrevisubulatum and Their BC1F3