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Dynamic changes in endogenous hormones in Taxus chinensis var.mairei seed during stratification

南方红豆杉种子综合处理过程中内源激素的动态变化



全 文 :广 西 植 物 Guihaia 31(3):370— 376 2011年 5月
DOI:10.3969/).issn.1000—3142.2011.05.017
南方红豆杉种子综合处理过程中
内源激素的动态变化
张艳杰 ,鲁顺保 ,高捍东
(1.江西师范大学 生命科学学院,南 昌 330022;2.南京林业大学 森林资源与环境学院,南京 2l0037)
摘 要:为揭示南方红豆杉种子内源激素与休眠的关系,采用酶联免疫吸附法(EI IsA)测定了经过层积处理
的种皮和胚乳的脱落酸(ABA)、赤霉素(GA3)、吲哚乙酸(IAA)、玉米素核苷(ZR)4种 内源激素含量的变化
情况。结果表明:种子胚乳中内源 ABA的含量随着层积时间的延长而逐渐下降,GA含量增加,IAA和 ZR的
含量先增加后降低,GA/ABA、IAA/ABA和zR/ABA逐渐增大,休眠随之解除。种皮中内源ABA、GA、IAA
和 ZR的含量均随着层积时间的延长而逐渐下降。其中GA/ABA的变化较大,因此,南方红豆杉种子休眠的
限制因子很可能是 ABA和 GA的平衡调控作用。
关键词:南方红豆杉 ;种子;内源激素
CLC Number:Q945 Document Code:A Article ID:1000—3142(20l1)03 0370—07
chinensis Var.mairei seed during stratification
ZHANG Yan-JieI,LU Shun-Bao1,GAO Han-Donge
(1.College of Life Science,Jiangxi Normal University,Nanchang 330022,China;2.Colege of Forest
Resources and Environment,Nanjing Forestry University,Nanjing 210037,China)
Abstract:In order to characterize relationship between endogenous phytohormones and dormancy of Taxus chinensis
var.mairei,changes in contents of four kinds of endogenous phytohormones,i.e.,abscisic acid(ABA),gibberelin
(GA3),indole acetic acid(IAA)and zeatin riboside(ZR)in spermoderm and endosperm of Taxus mairei were analysed
by enzymeqinked immunosorbent assays(ELISA).The results showed that ABA contents were gradualy decreased
in endosperm during stratification,while GA contents were increased and IAA and ZR contents firstly increased and
then decreased.Calculated ratio data of GA/ABA,IAA/ABA and ZR/ABA exhibited gradualy increasing trends
within stratification time and dormancy was relieved accordingly with them.In spermoderm,endogenous ABA,GA,
IAA and ZR contents were gradually decreased through stratification.The change in GA/ABA ratio was greater than
the other two ones.So it was suggested that the tradeoff between ABA and GA contents could control the dormancy
process in chinensis var.mairei seed.
Key words:Taxus cinensis var.mairei;seed;endogenous hormones
Taz“s chinensis var.maire iS one of rare Taxace—
ae Taxus species,widely distributing in China,such as
in Yangtze River basin,Henan Nanling Mountains,and
mountains or valleys in Shanxi,Gansu and Taiwan
Provinces.It has attracted global attention for extrac—
ting significant anticancer activity of taxol from barks。
Received date.2010—08—10 Accepted date:2011 01—1 3
Foundation items:Supported by Youth Foundation of Education Department of Jiangxi Province(GJJ11069)
University(2230)
Biography:ZHANG Yan Jie(1978一),female,Ph.D.,associate professor,Engaged in study on seedling.
Author for correspondence
3期 张艳杰等:南方红豆杉种子综合处理过程中内源激素的动态变化 371
twigs,leaves and other organs(Li et a1.,2003;Yuan et
dZ.,2002a,b,c). However,the species have faced
strong pressures of utilization in recent years and in—
deed suffered devastated looting.Thus,biologists were
seeking to increase the chinensis var.mairei popula—
tion within a framework of forest resource management
and genetic conservation. chinensis var.mairei seeds
with morphophysioIogical deep dormancy have under—
developed and dormant embryos,which must grow
continuously to some time before seed dormancy could
be broken.Listed as one of China’s first class key pro—
tected wild plants in 1999(Cao & Chen,1999),T.
chinensis var.mairei has an endangered existing state
because of its low natural reproduction. In recent
years,reports on artificial propagation and cultivation
of丁_chinensis var.mairei research were both rare in
China and other countries,while research on suitable
germination conditions was rudimentary.The develop—
ment and utilization of chinensis var.mairei were se—
riously restricted by its low seed germination rate.So,
it was necessary to study physiological processes of rf.
chinensis var.mairei seed germination.Involved in the
process of plant growth in all physiological regula—
tions,plant hormone was one of the most important
substances in controlling plant growth and develop—
ment(Ge,2004).Previous studies on endogenous hor—
mones were mainly concentrated on asexua1 reproduc—
tion(Han eta1.,1993;Tan eta1.,2008)and physiolog—
ical resistance(Rock et a1.,1999;Shi et a1.,2006;Lan
et a1.,2006;Wang et a1.,2009;Zhang et a1.,2009).
At present,there are many studies on species endoge—
nous hormones,especially on endogenous hormone as a
signal molecule involved in regulation of plant adapta—
tion under stress(Quan eta1.,2003).In order to carry
out in-depth study in this experiment,the paper has
taken a combination of temperature and hormones to
break the seed dormancy of chinensis var.mairei and
discussed changes of several hormones in seed germi—
nation process.These results will be helpful to under—
stand the occurring physiological and biochemical
processes during alternating temperature stratification
and provide reference information of丁.chinensis var.
maire seeds stratification for further study.
1 Materials and methods
1.1 Experimental Materials
丁.chinensis var.mairei fruits consisting of scarlet
or green cuplike arils were collected in Xiushui Coun—
ty。Jiujiang City,Jiangxi Province.With the age of 20
- 40 a,丁.chinensis var.mairei grew in 400— 500 m
sea level of the evergreen and deciduous broad-leaved
mixed forest in the valleys and the slopes.Arils and
empty seeds were floated off after they were colected
and then macerated in water.After naturally dried,the
seeds were sealed into polyethylene bags and stored in
refrigerator.Experimental materials were fresh seeds.
TGW was 65.048 g.
1.2 Experimental design and sampling
Six treatments labeled as A0 to A5 were set in
this experiment.Air-dried seeds A0 were soaked in 25
℃ warm water for 48 h;A1,A2 and A3 were soaked
respectively with 200,500,1 000 mg·L- GA3 for 48
h:A4 were washed by water for 1 week.Followed by
above treatments,all seeds were mixed with moist
sands at the ratio of 1:3 for stratification.The seeds
were firstly placed under a condition of variable tern—
perature of 23℃/10℃(12 h light)warm stratification
for 4 months.and then placed in 5℃ cold stratification
for 4 months. A5 were soaked in 25 。C water for 48
h,and then mixed with moist sand and placed outdoors
for natura1 stratification for 8 months.
All seeds were randomly sampled every two
months during stratification。and placed into一30。C re—
frigerator. In order to determine the hormones con一
tent,seeds were peeled into seed coats and endosperm
(including embryos.Taxus embryo was usually small
and difficult to be separated from endosperm,so the
embryos were not separated to test hormones)before
hormones were tested.(A3 seeds were almost rotten
after stratification,so their hormones were not tested).
1.3 Experimental methods
The seed materials that preserved in refrigerator
were weighed for 0.3 g,and added into 2 mL sample
extraction.These materials were grinded into homoge—
nate in ice bath and transferred to 10 mL tube. The
372 广 西 植 物 3l卷
mortars were washed with 2 mL extraction and trans—
ferred into the tube,then shaken evenly and placed into
4℃ refrigerator to extract for 4 h. Supernatant was
c。11ected after centrifugation of 4 000 rpm for 1 5 min.
Then 1 mL extraction.was added into precipitate and
extracted for 1 h at 4 ℃ and centrifuged again. The
supernatants were pooled and its volume was recorded.
The liquid was processed over C-18 solid-phase extrac—
tion column with a process of balance column with 1
mL of 80 methanol loading samples,washing col—
umn with 5 mL of 100 methanol and eluting hor一
-x
C





C



mones with 100 9/6 ethyl ether(5 mL)and lO0 metha—
nol(5 mL). Samples of washe&column were trans—
ferred into 5 mL plastic centrifuge tube.Vacuum was
concentrated to dry with nitrogen to remove methanol
of extraction,and then diluted to constant volume(USU—
a11v about 1.5 mL samples to 1 g fresh sample dilution
constant volume).Endogenous hormones ABA,GA3,
IAA,ZR were measured by enzyme-linked immunosor—
bent assaY(ELISA)(Zhang et a1.,1991;W u et a1.,
1988)Enzyme immunoassay kit was purchased from
China Agricultural University.Every sample was
J J-Ij J山 汕m Jlj jl
_ I1 畸 著4 O U
Fig.1 The changes of hormones content of different treatments in spermoderm
and emhryonic from T.chinensis var.mairei seed
hist0gram of figures from left t0 right is:A0,A1,A2,A4,A5 and the same loll0w
repeated for three times by the Spectra M ax Plus·
2 Results and Analysis
2.1 Change in endogenous hormones content
2.1.1 Change in ABA content ABA could induce
dormancY of developed seeds and inhibit embryo ger—
mination,but how regulation was carried out remained
unknown(Sun et a1.,2004).ABA content of seeds In
treatments rapidly decreased at first during stratifca—
tion and then maintained at a low level(Fig.1:A).The
results showed that ABA content was high in mature
endosperm and this might be one of the main reasons
of seed dormancy. ABA content of endosperm in A0
and A4 treatnlents decreased slowly,while changes in
ABA contents with the other three treatments were
obvious.and the change in A5 treatment was the most
obvious.ABA synthesis was blocked with the embryo
after-ripening,leading to a decrease in ABA content
which benefited to the relieve seeds dormancy.There
was significant difference by ANOVA of treatments,
stratification and their interaction effects. The ABA
content was significantly different by LSD among five
treatn硷nts.Different treatments affected ABA content
of endosperm. The differences of ABA contents be—
tween March and May,April and May,June and July
were not significant by LSD of treatments and stratifi—
cation。while the differences in other stratification were
significant.
The ABA content of seed coats decreased at dif—
3期 张艳杰等:南方红豆杉种子综合处理过程中内源激素的动态变化 373
ferent speeds during stratification in these treatments
(Fig.1:A).The ABA contents of seed coats were sig—
nificantly lower in A0。A1。A2 and A4 than in A5.
These results showed ABA content of seed coats dur=
ing stratification continuously decreased.The decrease
in ABA content was significantly faster in A0,A1,A2
and A4 than in A5,which benefited to seed germina—
tion.According to the decreasing speed of ABA con—
tent,the A0,A1,A2 and A4 treatments could be divid—
ed into two stages:the first 2—3 months were a phase
of sharp decrease and then a stable phase afterwards.
ABA content increased in A4 during stratification of 4
— 5 months. ABA content in A5 rapidly decreased
during 1—2 months。then slowly decreased during 3—
5 months,and then rapidly decreased during 6— 7
months,and finally lasted in stabilized stage. In the
early stratification.the ABA content in seed coat was
significantly higher than that in endosperm. Changes
in ABA content in seed coat and endosperm were few
after stratification for 6 months ANp showed that
there were significant differences in ABA content a—
mong the treatments,stratification time and their inter—
active effects. The LSD result showed there was no
significant difference in ABA content between A0 and
A4,A1 and A4,during 3 to 5,7 to 8 months. The
differences of others’ABA content were significant.
2.1.2 Changes in GA3 content GA3 was one of the
strong active GAs,and could promote cell elongation
and break GA3 which played an important role in the
promotion of seed development and germination regu—
lation. The GA3 contents of endosperm in different
treatments were shown in Fig.1:B.GA3 content in—
creased slowly in A0,but rapidly in A1 during stratifi—
cation for 2 months,and remained high in A2.Due to
the fact that A1 and A2 seeds were soaked into high
concentration of exogenous,the content of GA3 was
high.GA3 content firstly increased and then decreased
in A4.GA3 content was higher in A5 than others,but
the reason was still unclear. ANOVA showed that
differences were significant among treatments,stratifi—
cation and their interaction. M ultiple comparisons
showed that the differences
significant,indicating that the
among treatments were
effect of exogenous GA3
on endosperm GA3 content in endosperm was obvious.
Multiple comparisons of different stratification times
showed that the difference of GA content was not sig—
nificant during stratification for 4— 7 months,but oth—
ers’differences were significant.
GA3 content in seed coat was low in A0 and A4
(Fig.1:B). GA3 content rapidly decreased during
stratification for 3 months and then remained stable in
A1 and A2.GA3 content of seed coat fastly decreased
for 2 months。and then remained stable in A5. GA3
content was higher in seed coat than in endosperm at
the beginning of stratification,and then gained opposite
result after stratification for a period.A possible rea—
son was that GA3 might transfer from seed coat to en—
dosperm ANO showed that the differences of GA3
contents were significant among treatments,stratifica—
tion and their interaction effects.The multiple compar—
ative showed that the differences were not significant
between A0 and A4,and between stratification for 3
and 5 months,while the others’differences were signif—
icant.
2.1.3 Changes in IAA content The results of IAA
content in endosperm were shown in Fig.1:C. IAA
content of endosperm firstly increased and then slightly
decreased in treatments during stratification. IAA
change was much obvious in A0。from 2.685 ng/g FW
during stratification for 1 month to 7.645 ng/g FW for
5 months.and then to 3.888 ng/g FW for 8 months.
The change extents were small in other treatments,for
example,IAA content remained stable in A5. At the
beginning of stratification,IAA content increased due
to it has transferred from seed coat into endosperm,
which was corresponding to the decreasing of IAA
content in seed coat.Embryo cell division continued to
strengthen,leading to a decrease of IAA content.
ANoVA showed that the diffe-rences of IAA were sig—
nificant between treatments,stratification and their in—
teraction effects. Multiple comparisons in different
treatments showed that there were significant differ—
ences of IAA between A2 and other treatments.IAA
content was high in A2. The difference between AO
and A1 was not significant,but with the other treat—
ments.A5 had the lease IAA content.There were sig—
374 广 西 植 物 31卷
nificant differences of IAA between A5 and the other
treatments.The multiple comparative also showed that
there was no significant difference of IAA content dur—
ing stratification between 1 and 2 months,among 3,5
and 8 months,and among 4,5 and 7 months as wel1.
The results showed that IAA content change in endo—
sperm during stratification was not obvious.
IAA content in seed coat decreased during stratifi—
cation in different treatments(Fig.1:C).At the begin—
ning of stratification in al treatments,IAA content in
seed coat was higher than that in endosperm,and then
IAA content decreased in seed coat and increased in en—
dosperm ANO。 showed that differences of A
content were significant between treatments,stratifica—
tion and their interaction effects.Multiple comparisons
of treatments showed that there were significant differ—
enees between the treatments except between A0 and
A4.There were significant differences among stratifi—
cation periods except between 4 and 6 months.IAA
content in seed coats obviously decreased during strati—
fication.
2.1.4 Changes in ZR content Changes of ZR content
in endosperm from different treatments were shown in
Fig.1:D.ZR content of endosperm firstly increased
and then slightly decreased during stratification in A0,
and was 6.50 ng/g FW in A1.ZR contents firstly in—
creased to a peak during stratification for 2 months,and
then decreased in A2,A4 and A5 ANO showed
that the differences of ZR contents in endosperm were
significant between treatments,stratification and their
interaction effects.Multiple comparisons of treatments
showed that there was no significant difference be—
tween A1 and A4,A2 and A5,but in A0.The differ—
ence of ZR content was significant during stratification
between the first and the other months,but was not
significant during stratification among 2,4 and 6
months,and between 5 and 6 months.
ZR content in seed coats decreased gradualy dur—
ing stratification in A0,A1 and A5(Fig.1:D).Change
of ZR content was much obvious and got a peak of
27.37 during stratification for 8 months in A5. A
possible reason was that ZR in seed coats was influ—
eneed by external environment in nature condition.ZR
content increased at first for 2 months,and then gradu—
ally decreased for 4 months,and began to increase for
the last 2 months.These changes of ZR were as same
as GA3 content of seed coat and endosperm ANO
showed that there were significant differences of ZR
content in seed coat between treatments,stratification
and their interaction effects. Multiple comparisons of
treatments showed that there were significant differ
ences among the treatments except among A0.Al and
A4.The difference of ZR was significant among strati—
fication periods except between 6 and 7 months,7 and
8 months.These results showed that stratification af—
fected on ZR content of seed coat at the beginning of
the fifth month.
2.2 Changes of endogenous hormones content ratio
2.2.1 Changes of GA/ABA ratio GA/ABA ratio of
species endogenous hormones changed with seed dor—
mancy and germination.The ratio of endosperm slowly
increased during stratification (Fig. 2:A ),which
showed that seed dormancy was gradually relieved

The GA/ABA ratio firstly increased,reaching the peak
during stratification for 3 months,and then decreased
for 4 months,at last increased again for 5 months in
A2.The result was consistent with hormonal balance
hypothesis,which indicated that seed dormancv was
controled by the interaction of ABA and GA
. The
GA/ABA ratio in seed coat did not change during
stratification in A0 and A4.The change of the GA/
ABA ratio in seed coat firstly increased and then de—
creased in A1,A2,obviously because seeds were soaked
by 200,500 mg/I of exogenous GA3.The ratio grad
ually increased during stratification in A5

2.2.2 Changes of IAA/ABA ratio IAA /ABA ratio in
endosperm in treatments was shown in Fig.2:B. The
ratio firstly increased and then decreased in A0 and
A2,while gradually increased in A1,A4 and A5
. The
ratio of the seed coat firstly increased and then de—
creased and finally stabilized in A0,A1 and A2,while
increased during stratification in A4. As the lowest
one,the ratio change in A5 was slowly decreased dur—
ing stratification from l to 5 months,and then in—
creased and decreased again.
2.2.3 Changes of ZR/ABA ratio ZR/ABA ratio in
3期 张艳杰等:南方红豆杉种子综合处理过程中内源激素的动态变化
endosperm gradually increased during stratification
(Fig.2:C).Change of the ZR/ABA ratio in A0 was
the most obvious and,reached a peak for 6 months,but
changes of other ratios were little.Greatly varying a—
mong treatments,ZR/ABA ratio in seed coat firstly in—
creased and then decreased during stratification in AO,
A1,A2,A4 and A5.
2.2.4 Change of(GA+IAA)/ABA ratio The(GA+
16
14
《 1 2
罢1 0
8
a 6
4
2
0
4.5
4 0
3 5
《3 0
2.5
2 0
N 1.5
1 0
0.5
O.0
GA/ABA.
2 3 4 5 6 7 8
Time of st rat
j山 k
2 3 4 5 6 7 8
Tlm6 of st ratif






IAA)/ABA change of endosperm was consistent with
the GA/ABA in general(Fig.2:D).The ratio gradual一
1v increased during stratification,but this change was
different.The ratio change was small in A0 and A4
but 1arge in A1 and A5. The ratio firstly increased
during stratification for 3 months,and then decreased,
but slowly increased again in A2.The(GA+IAA)/
ABA ratio change in seed coat was consistent with the
Fig.2 The changes。f horm。nes ratio am。ng different treatments in endosperm and
embryonic from T.chinensis var.mairei seed
3 Discussion
3.1 Changes of seed’s endogenous hormones
The ABA content in seed coat and endosperm
rapidly decc!ased at the beginning of the stratification
and then changed a little bit,which showed that the
ABA content decreased with the embryo after-ripening
at 23。C/10℃(12 h light)stratification and ABA syn—
thesis was blocked.The IAA content in ABA antago—
nism increased in the early stratification period and that
eould Dromote the degradation of ABA and was benefi—
cial to relieve seed dormancy,which was consistent
with predecessors’study(Jacobsen et a1.,1985).
The relation of seed dormancy and germination
with GA content was noticed wildly(Sun et a1.,
2006).In this study,during stratification,the GA con—
tent of endosperm gradualy increased in A0 and A1,
which could activate the enzyme activity and promote
seed germination. At the early stage of stratification,
the GA in seed coat was significantly higher than that
in the endosperm.GA content in seed coat decreased
during stratification,while its content in endosperm
gradually increased.So the GA content in endosperm
was slightly higher than that in seed coat.A possible
reason was that the part of GA had transferred from
seed coat to endosperm during stratification process.
So it was suggested that the GA contents could control
the dormancy and germination process in seed(Lei et
倪£.,2009).
IAA was a hormone that could promote cell divi—
sion and expansion,weaken the role of germination in—
hibiting substances and regulate seed the material of
development and energy metabolism (Sheng et a1.,
2006:Ding et a1.,2007;Sha et a1.,2007).In general,
IAA content in endosperm firstly increased and then
slightly decreased,while its content in seed coat gradu一

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376 广 西 植 物 31卷
ally decreased. The IAA content in seed coat was
higher than that in endosperm at the early stage of
stratification.Then the change of IAA content was 1it—
tle,might because the hormonal part of the seed coat
has transferred to endosperm.
ZR concentrations in the seed were relatively high
and significantly increased during fuitlet growth(Xiao
et a1.,2007).In this study,ZR content of endosperm
gradually increased during stratification in A0,while
the other ZR contents firstly increased and then de—
creased and finally stabilized.ZR content in seed coat
was consistent with endosperm and decreased during
stratification.At the beginning of stratification,the ZR
content in seed coat was higher than that in endo-
sperm,but then appeared as the opposition.
3.2 al锄 ges of seed endogenous hormones ratio
Research by Khan(1975) illustrated that seed
dormancy and germination were related to the absolute
content of endogenous hormone in plant and the ratio
of various horm ones,especially the ratio of horm ones
that could promote or inhibit growth.The GA/ABA
ratio of endosperm increased during stratification,and
the biggest increase was 25.6 times in A1.The chan—
ges of IAA/ABA,ZR/ABA and(GA+ IAA)/ABA
gradually increased during stratification. It could be
considered as dormancy relieve and germination promo—
tion by increasing IAA/ABA and ZR/ABA in the seed
after-ripening process.
The ratio of GA/ABA in seed coat firstly in—
creased and then decreased in A1 and A2 during strati—
fication,while the ratio gradually increased in A5.The
IAA/ABA and(GA+ IAA)/ABA ratio firstly in—
creased and then decreased during stratification. The
ZR/ABA ratio changed during stratification in A0,Al,
A4 and A5,but the ratio in A2 firstly increased,then
decreased,and at last slowly increased again. The
change in GA/ABA ratio was greater than the other
two ones. So it was suggested that the tradeoff be—
tween ABA and GA contents could control the dor—
mancy and germination process in 丁.chinensis var.
mairei seed,which results were agreement with previ—
OUS research on others(Lei et a1.,2009).
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