全 文 ：广 西 植 物 Guihaia 28(3)：373— 378 2008年 5月
Regulatory effects of gibberelic acid(GA3)
(Scutellaria baicalensis)in vitro
ZENG Hong-Yan ， ．Saxena Praveen．K．
(1．Institute of Biotechnology，University of Xiangtan，YOangtan 411105，China 2．Department
of Plant Agriculture，University of Guelph，Guelph，Ontario，Canna N1G 2W1)
Abstract：Influence of exogenously applied gibberellic acid(G )on shoot and rot formation and development in
shoot tip and stem tissue cultures of Huang-qin(Scutellaria baicalensis)，an important medicinal plant，was investiga—
ted．G application at the concentrations of 1-20~mol／L resulted in the significant enhancement of shoot formation
and growth while it reduced rot formation and development．In different transferring tests，the exogenously applied
growth regulators G and IAA resulted in significant chang es in the concentrations of major flavonoids in Huang-qin
tissues．The highest concentrations of baicalin，baicalein and wogonin were 14．90，2．70 and 0．54 g·mg-1(dry
weight)，respectively，when the explants were pre-cultured on 2．5／lmol／L IAA medium for 6 days before transfered
to MS0 medium containing G at 5／lmol／L in continuous darkness．
Key words：adventitious rooting；Huang-qin；Scutellaria baicalensis；gibberelic acid；baiealin；baicalein and wogomn
CLC Number：Q943 Document Code：A Artide ID：1000-3142(2008)03-0373—06
Huang-qin(Scutellaria baicalensis Georgi)is one
of the most important medicinal plants used in tradi—
tional Chinese medicine (TCM )． Its plants contain
high levels of flavonoids，a group of low molecular
weight polyphenolic compounds，which are considered
to be the main medicinally active constituents of
Huang-qim Whole plant extracts and flavonoids of
Huang-qin were reported to possess antiviral，antican—
cer，antibacterial，antimutagenic，antioxidative(Gao et
a1．，1 998)，anti—inflammatory and anxiolytic activities
(Hui eta1．，2002；Zhang eta1．，2003)．Recent findings
indicate that Huang-qin has melatonin and many other
secondary metabolites with potential medicinal proper—
ties(Murch et a1．，1997；Pawlicki& Welander，1992)．
Hence，Huang-qin is an ideal plant for scientific studies
aiming to uncover hovel active medicinal constituents．
Although it has been one of the major ingredients
of TCM for over two thousand years，westem ers’inter—
est in Huang-qin is relatively recent． In East Asia，
Huang-qin is generaly wild harvested from nature．
Wild harvest of medicinal plants presents many risks．
Genetic differences among the plants，differences in the
physiological stages at the time of harvest，contamina—
tion with biotic and abiotic agents，differences in har—
vesting and processing procedures and adulteration
with other plant materials can result in inconsistency in
active constituents，medicinal efficacy and even in seri—
OUS toxicity(Li et a1．，2000)．A recent study(Ye et
a1．，2004) reported significant variation in chemical
composition and biological activities of commercial
Huang-qin extracts． Most of these problems can be
solved by use of genetically uniform plant material in
controlled production systems． In vitro propagation
systems can be used as an alternative to wild harvest of
this plant． A considerable amount of efforts has been
made in developing in vitro propagation systems for
Huang—qin by our group(Li eta1．，2000；Murch eta1．，
1997，2006；Zeng et a1．，2007；Zobayed et a1．，2004)．
However，an efficient，reproducible，high—frequency di—
rect regeneration system has to be established to allow
Received date：2006—12—29 Accepted date：2007—06—07
作者简介 ：曾虹燕(1963一)，女 ，河南洛阳人，博士，教授，主要从事植物资源开发和酶工程 ，(E-mail)hyzeng(~xtu．edu．an。
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374 广 西 植 物 28卷
mainstream production of high quality plant material
for the discovery of novel compounds，testing the me—
dicinal efficacy and commercialization．
Gibberellic acid (GA3)is fl plant hormone of
widespread occurrence in higher plants．It is involved
in the control of number of plant growth and develop—
ment responses．On one hand，gibberellic acid has been
used in the microculture of plants for fl number of pur—
poses，including the stimulation of shoot proliferation in
Malus(George& Sherrington，1984)and Atriplex ca—
Kscens(Wochok& Sluis，1980)．But it was found to
inhibit shoot growt h in Ilex paraguariensis(Sansber
ro et a1．，2001)and Hancornia speciosa(Pereira-Netto
Pt a1．，2003)．On the other hand，there is evidence for
a positive effect of GAs on adventitious rooting in a
number of spedes(Le et a1．，2001)．In some cases its
inhibitory effect has been reported (Pawlicki& W e—
lander，1992)．Direct application of GAs tends to de—
crease rooting，and the application of inhibitors of GA
biosynthesis can enhance rooting (Porlingis et a1．，
1996)．So there are different influences of GAs on
shoot and root form ations of different species． For
these reasons，we investigated the effects of GA3 on
root morphogenesis and shoot form ation of the germ —
plasm line HQ11 from Huang-qin．The role of indol~
3-acetic acid(IAA)。indol~3一butyric acid(IB A)and
thidiazuron(TDZ)during different stages of root and
shoot development has been reported as critical in
Huang-qin(Li et a1．，2000；Zeng et a1．，2007)．How—
ever，we do not have fl complete understanding of
whether other phytohorm ones including GA3 play fl
role during root and shoot induction and／or develop—
ment as wel1．
The present research is aimed at finding fl repro—
ducible method for in vitro multiplication of Huang-qin
through shoot proliferation and rhizogenesis from shoot
tip and stem explants of HQ1l1．The tissue cultures
were also examined for the ability to produce major fla—
vonoids compounds．
1 M aterials and methods
1．1 Plant matedal
Huang-qin line，HQ11，was selected because of its
high regeneration capacity in vitro and wel studied
chemi cal profile (Murch et a1． 2004)． The plants
were maintained asepticaly for about 5 years through
subculturing of shoot explant in MSO containing MS
salts，B5 vitamins，30 g·L- sucrose，and 3 g·L- gel—
lan gum (Gelrite，Schweitzerhall，So uth Plainfield，NJ，
USA)in about every two months．AU cultures were
incubated in fl growt h room wi th fl 16 h photoperiod
under col-white light(30~40 tmol／L·m- ·s- )at
25℃ ．Stem and shoot tip were used as explants．Stem
explants(about 1．5cm)were prepared by cutting fl
node and excising the leaves，and shoot tip explants
(1．5—2 cm)were prepared by cutting fl node and the
tip wi th intact leaves．
1．2 Treatment
A two-step culture for shoot or root form ation
was followed in the experiments．Seven explants were
cultured on fl Petri dish containing 25 mL of MSO me—
dium supplemented with 2．5／~mol／L IAA or 2．5
／~mol／L TDZ for 6 days．They were then transferred
to a plate wi th MSO medium．
GAs(1—20／~mol／L)was added to 2．5~mol／L
IAA or 2．5／~mol／L TDZ or MSO medium and the ex—
plants were cultured on the both media without GA3 as
the controls． After 30 day incubation，the number of
shoots or roots per explant was determined．The ex—
plants were cultured on MSO medium for all 30 days
(hereafter referred to as MSO contro1)and those on
MSO medium with 2．5 vtmol／L TDZ or 2．5~mol／L
IAA (hereafter referred to as TDZ-treated controls or
I treated controls)for all 30 days．
1．3 Fiavonoid analysis
Plantlets developed from both stem and shoot tip
explants were collected after 30 days of culture and
dried at 37 ℃ for 2 days in fl drying chamber，then
ground into fine powder and transferred into amber-
colored 20 mL vials．The weight of the plantlets in the
identical treatment was recorded before or after drying．
Analytical methods for quantification of baicalein，ba—
icalin and wogonin were made according to previously
described methods by Murch(Murch et a1．，2004)．
1．4 Statistical aIlalysis
In each experiment，the treatments consisted of
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3期 曾虹燕等：赤霉素 GAs调节黄芩组织培养中芽和根的形成 375
three replications，and the experiments were repeated at
least two times．Data were analyzed using the Statisti—
cal Analysis System (SAS Institute Inc．，2004)with
Student-Newrnan-Keuls means separation test(P≤
0．05)．
2 Results and discussion
2．1 Efect of GA3 on shoot induction and diferentiati0n
Shoot induction of HQll was achieved by pre-in—
cuhating explants on 2．5 t~mol／L TDZ medium for 6
days and subsequently culturing them onⅣ【SO medium
for growth and development of shoots for 24 days．
The addition of GAs to both media significantly stimu—
lated shoot formation． GAs at concentration as low as
1 t~mol／L caused about 1．3一and 0．2一fold increases of
the control in shoot number per tip and stem explant，
respectively，when added to 2．5 t~mol／L TDZ medium．
Approximately 3．0一and 5．0-fold increases of the con-
trol were obtained，respectively，when added to the
MSO medium．GAs added to 2．5~mol／L TDZ medi-
um was similarly effective on shoot number per tip ex—
plant over all the concentrations tested，but the number
of shoots per stem explant increased wi th increasing
concentrations of GAs until it reached fl maximum at
5 0~mol／L and then decreased 晒 en GAs added to
Ⅳ【SO medium．the shoot number per explant reached fl
maximurn at 1．0~mol／L and then decreased．In addi—
tion，there was no clear difference between shoot hum—
ber per explant cultured o13．the control，2．5~mol／L
TDZ and MSO control media(Table 1)．
Our group has demonstrated previously (Li et
a1．．2000)that TDZ effectively induced shoot regener—
ation on cultured intact seedlings，etiolated hypocotyl
explants and sterile stem segments of Huang-qin and
adventitious shoots formed through an interm ediate
callus． The present experiment found that fl sim lar
adventitious shoots were induced by TDZ，but TDZ did
not stimulate shoot form ation on cultured sterile tip
and stem segm ents of HQIu(data no shown)．The
maximum shoot induction was observed at 2．5／~mol／L
TDZ concentration for the explants of HQll(data no
shown )．TKs is presum ably due to differences be—
tween the different Huang-qin germ plasm lines exam—
ined．An d exogenesis GAs promoted sho t form ation
and inhibited the callus formation of HQl1．The stim-
ulatory effect of G was more pronounced when used
during the period of MSo medium than during the ini—
tial 6 days of 2．5 ttmol／LTDZmedium( le 1)．In-
hibition of callus formation by GAs has been reported
previously by Rudug(2002)．The observation of inhi—
bition of sho t multiplication and growth from the axil—
lary meristem of the explants during flu initial 6 days
(Table 1)could be due to inhibition of TDZ on the de—
velopment of axillary primordium． After the intial 6
days，sho ts had developed to fl point where TDZ was
unable to inhibit the shoot formation from axillary pri—
mordiun1．
2．2 Efect of G on root induction and diferentiation
The explants were pre-incubated on 2．5 tmol／L
IAA medium for 6 days and subsequently cultured on
Ⅳ【SO medium for 24 days． The addition of GAs to
both media significantly inhibited root formation (Ta—
ble 2)．GAs at the concentration as low as 1 ttmol／L
reduced root number per tip and stem explant to about
6O 9／5 and 53 9／5，respectively，in comparison with the
control when added to 2．5 ttmol／L IAA medium and
about 48 9，6 when added toⅣ【SO medium The number
of roots per explant decreased wi th increasing concen—
trations of GAs but no clear trend was evident when
added to 2．5 ttmol／L IAA medium． GAs added to
Ⅳ【SO medium had fl sim lar influence on root form ation
in all concentrations tested(Table 2)．
There is considerable evidence that IAA with
maximum enhancement was observed at 2．5 ttmol／L
concentration can promote root regeneration of ex—
plants of HQll(Zeng et a1．，2007)．In contrast to its
positive effect on shot formation，exogenous gibberel—
lin A3 significantly inhibited root growt h (Table 2)．
Root growt h was inhibited to fl lesser extend if applied
duringⅣ【SO medium in stead of during 2．5 ttmol／L
IAA phase．GA3 had flu inhibitory effect after 6 days
of the culture，so the ro ts from the explants could be
induced by IAA before 6 days．This may suggest that
the level of endogenous GAs is sufficient for root
growt h．
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376 广 西 植 物 28卷
1,umol／L G
MS0
2．5 J~mol／LTDZ
2．5 Jamol／L TDZa／MSO
2．5 J~mol／L TDZb／1／*mol／L GA3
2．5 Jamol／L TDZb／5／*mol／L G
2．5 Jamol／L TDZb／10／*mol／L
2．5 J2mol／L TDZb120／imol／L GA3
2．5 Jzmol／L TDZ+1／*mol／L GA；／MSO
2．5 Jamol／L TDZ+5／*mol／L G ／MSO
2．5 J2mol／L TDZ+10／*mol／L G ／MSO
2．5 J2mol／L TDZ+20／*mol／L G ／MSO
3．0476 fl
0．8571 b
1．1905 b
0．9048 b
3．6190 fl
3．5238 fl
2．8095 ab
2．238lab
2．0476 ab
2．4286 ab
2．5238 ab
2．5238 ab
2．4762 bcde
1．4286 cdef
1．2857 def
0．7619 ef
4．5714 fl
4．000 ab
2．7619 bcd
3．0952bc
0．9048 ef
2．4286 bcde
1．4286 cdef
0．5238 f
O．187 b
0．198 fl
0．084 c
0．045 g
O．O39
0．043 g
0．028 h
0．088 c
0．068 e
0．052 f
0．074 d
3．34 i
6．01 e
11．8 fl
5．98 e
5．29 g
5．46 f
3．74 h
7．54 c
7．41 d
9．51 b
7．46 d
0．039
0．035
Trace
Trace
Trace
Tlrace
ND
Tlrace
Trace
Trace
Trace
1／~mol／L G ：Explants were cultured on 1／*mol／L G medium for 30 days)2．5／*mol／L TDZ：Explants were cultured on 2．5／~mol／L TDZ me-
dium for 30 days；MSO：Explants were cultured on MSO medium for 30 days．。Explants were pre-cultured on 2
． 5／~mol／L TDZ medium for 6 days be-
fore transferred to MSO medium． Explants were pre-cultured on 2．5／*mol／L TDZ medium for 6 days before transfered to MSO medium containing G
at the concentrations of 1，5，10 and 2O／*mol／L Explants were pre-cultured on 2．5／~mol／L TDZ medium containing G at the concentrations of 1，5，
10 and 2O／*mol／L for 6 days before transferred to MSO medium． Data recorded after 30 days of culture．
Table 2 Influence of GAa applied on root growth of HQm tissue culture in continuous darkness Treatment
Treatment
Number of shoots per explant
Tip Stem
Content of flavonoids
Baicalein Baicalin Wogonin
( g·rag-0，dry wt)( g·rag-0，dry wt)( g·rng0，dry wt)
1／*mol／L G ：Explants were cultured on 1／*mol／L G medium for 30 days)2．5 IAA：Explants were cultured on 2．5／*mol／L IAA medium for
30 days；MSO：Explants were cultured on MSO medium for 30 days．。Explants were pre-cultured on 2．5／*mol／L IAA medium for 6 days before trans—
fered to MSO medium． Explants were pre-cultured on Z．5 flmol／L IAA medium for 6 days before transferred to MS(3 medium containing G at the
concentrations of 1，5，10 and 2O／*mol／L Explants were pre-cultured on Z．5／*mol／L IAA medium containing G at the concentrations of 1，5，10 and
2O／*mol／L for 6 days before transferred to MSO medium． Da ta recorded after 30 days of culture．
2．3 Efect of GA3 oil flavonoid content
Ultimately，Huang-qin tissues are grown for their
capacity to produce medicinal metabolites．Analyses of
the concentration of bioactive compounds in the ex—
plants demonstrated that the addition of GA3 to 2．5
／lmol／L TDZ or 2．5／lmol／L IAA for 6 days or MSO
media for 24 days had different influences on the con-
centration of bioactive compounds(Table 1，2)．In
IAA-treated contro1．IAA significantly promoted their
concentrations in the explants in continuous darkness．
The concentrations of baicalein，baicalin and wogonin
have about 10一，3一and 4-fold increase in compared to
MSO control，respectively． However，GA3 added to
2．5／lmol／L IAA medium had a negative influence on
the concentrations of the three compounds(Table 2)．
The increased concentrations of the three compounds
were seen only at I-5／~mol／L GA3 added to MSO me-
diu札 Maximum enhancement was observed at 5
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3期 曾虹燕等：赤霉素 GA3调节黄芩组织培养中芽和根的形成 377
／~mol／L to about 5．5一，4．2一and 6．2一fold of the con—
trol，respectively，which was higher than those of IAA—
treated contro1． The highest concentrations of baica—
lein，baicalin and wogonin in continuous darkness were
at 5／~mol／L G added to MSO medium with dry
weights of 2．70，14．90 and 0．54 g·mg ，respective—
ly(Table 2)．
In TDZ_treated contro1．TDZ had no effect on the
concentrations of baicalein and wogonin in comparison
th ()-control，but increased the concentration of ha -
icalin to about 2_fold of theⅣIsO contro1．The concen—
trafions of wogonin in all G treatments were trace．In
the control，TDz reduced the concentrations of baicalein
and wogonin but enhanced the concentration of baiealin
to about 3．5一fold of MSO contro1．The addition of G
to 2．5／~mol／L TDZ medium had less significant influ—
ence on the concentrations of the haicalein and wogonin，
but decreased concentration of baicalin in comparison
th the contro1．The addition of G to M SO mediurn
decreased the concentrations of baicalein and baicalin in
comparison with the control(Table 1)．
One of the major limitations to the development of
high-quality plant-based medidnes is the need to adapt
both traditional and high-tech agricultural practices to
unusual species．Yield of medicinal plants needs to be
defined both in biomass，chemi cal composition and the
quality．Our results show that there is a great potential
of using exogenous auxin for the promotion of pharma—
cological active compounds of Huang-qin．GA3 use at
low concentration is expected to be effective，when ex—
plants are cultured on 2．5／~mol／L IAA-treated induc—
tion medium for 6 days and then cultured onⅣ【SO me—
dium supplemented with 1-5／~mol／L GAs for 24 days．
Under the conditions，ma ximum concentrations of ba-
icalein，baicalin and wogonin in the explants of HQ11
were obtained in comparison wi th those reported previ—
ously(Murch et a1．，2004；Zeng et a1．，2007)．There—
fore，interaction between exogenous GA3 and IAA
should be further investigated．
In conclusion，the present investigation implicated
both the exogenous and endogenous G playing a
stimulatory role during shoot development of Huang-
qin germplasm line HQl1．The endogenous content of
GA3 is probably not optima l for shoot formation．
However，the level of endogenous gibberellin for induc—
tion and growth of root presumably is adequate．Tis—
sue culture techniques can play an important role in
clonal propagation of elite genotypes of Huang-qin．
The authors wi sh to express their sincere grati—
tude to Mr．R Nichols，Department of Plant Agricul—
ture，University of Guelph，On tario，Canada for gener—
OUS help in revising ma nuscript．
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378 广 西 植 物 28卷
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赤霉素 GA3调节黄芩组织培养中芽和根的形成
曾虹燕1，2 ，Saxena Praveen．K．2
(1．湘潭大学 生物技术研究所 ，湖南 湘潭 411105；2．Department of Plant
Agriculture，University of G~elph，Guelph，Ontario，加拿大)
摘 要：应用组织培养技术对黄芩进行外源激素调控研究。在培养不同时间进行的不同培养基之间的转移培
养研究表明，组织培养条件下，在培养基中添加赤霉素，可显著刺激黄芩外植体芽的形成，同时抑制根的生长。
在加有GAz的IAA培养基上，GAz显著影响黄芩组织培养物中的黄酮含量。在黑暗条件下，开始在 2．5／lmol／
L IAA培养基中培养 6 d，随后转移到5／lmol／L GAz培养基上培养，黄芩外植体中黄岑苷、黄岑素和汉黄芩苷的
含量最高，分别为 14．9O，2．7O和 0．54 g mg (干重)。
关键词：不定根；黄芩；赤霉素；黄岑苷；黄岑素；汉黄芩苷
(上接第 421页 Continue from page 421)
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