全 文 ：广 西 植 物 Guihaia 25(5)：481— 488 2005年 9月
梅花‘南京红须’、‘南 京红’ 花色的呈现特征
赵昶灵1，2，郭维明3 ，陈俊愉4
(1．南京农业大学生命科学学院，江苏南京 210095；2．云南农业大学农学与生物技术学院。云南昆明
650201；3．南京农业大学园艺学院，江苏南京 210095；4．北京林业大学园林学院，北京 100083)
摘 要 ：梅花 ‘南京红须’、‘南京红 ’的花色主要存在着花发育阶段导致 的时间变化 ，反映其花色受花发育控
制。二者的花色都在蕾期最浓艳 ，在初花期略淡，在盛花期又稍浓，在末花期最淡，尽管花瓣在花开放时便开
始衰老 ；在整个花发育时期 ，同一朵花不同层次花瓣的颜色浓淡均为：外层花瓣>中层花瓣> 内层花瓣 ，即花
瓣在花冠中的具体排列位置决定着该片花瓣的特定颜色深浅；但不同层次花瓣颜色的变化趋势不完全一致。
同时，两个品种外层花瓣的总黄酮含量变化与外层花瓣的色度变化成正相关。而花朵在树冠的着生部位导致
的花色差异极不显著 ，表明‘南京红须 ’、‘南京红 ’的花色的空间变化极微。本 文可为梅花红色花色的机理探
索和花色色素生物合成关键酶基因cDNA克隆中的花朵选择提供参考。
关键词 ：‘南京红须’；‘南京红’；花色；呈现特征
CLC number：Q944．58 Document code：A Article ID：1000—3142(2005)05—048卜O8
一 ‘ ’ ‘ 一
0f ’ flow~ col,15xpresslon CIaracteristics the er ors t
of Prunus mume‘Nanjing Hongxu，and
Prunus mume‘Nanjing Hong，
ZHAO Chang—ling 一，GUO Wei—ming。，CHEN Jun—yu4
(1·College of fJife Sciences，Nanjing Agricultural University，Nanjing 210095，China；2．College of AgricultMral
Sciences and Biotechnology，Yunnan Agricultural University，Kunming 650201，China：3．Colege of
Horticulture，Nanjing Agricultural University，Nanjing 210095，China；4．College of Landscape
Architecture，Beijing Forestry University，Beijing 100083，China)
Abstract：The flower colors of Prunusmume‘Nanjing Hongxu’and P．mulr／e‘Nanjing Hong’exist mainly tem—
poral changes caused by different flower developing stages，reflecting that the flower colors are under the con—
trol of flower development．The flower colors of these two cuhivars are all the strongest in Alabastrum Deri—
od，thin in some sort in Initial flower period，thicken appreciably in Profuse flower period and thin furthest in
Final flower period even if the peta[s senesee as soon as they begin to splay
． At different flower developing
stage，the chroma differences of petal colors of different layers are：outer layer> middle layer> inner layer
，
namely，the concrete collocating site of petal in the corolla decides the specific color chroma of this Deta1
． But
the change trends of the petal colors of different layers are not completely similar． In the mean time，the con—
tent changes of total flavonoids of the outer layers of two cultivars are positively correlated with the Deta1 co1or
changes of outer layers．On the other hand，the flower color difference induced by the inserting site of the
flower in canopy is not remarkable at all，showing that the spatial variations of the flower colors are very Detit
．
Received date：2004—09—1 3 Accepted date：2005—03—2O
Foundation item：Partly supported by t he Research Fund for Young Scholar of Yunnan Agricultural University
Biography：Zhao Chang-ling(1 969一)·Male，Born in Dujiangyan City，Sichuan Province，Doctor of science，Associate professor．
working in Plant physiology，Biochemistry and Molecular biology．
Author Corresponding
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482 广 西 植 物 25卷
This paper could be a reference or a premise for the exploration on the flower co1or mechanism of red Mei
flowers and the flower-selecting in the cDNA cloning of the key enzyme genes determining the biosynthesis of
anthocyanins in Mei flower．
Key w0rds：P．m“me Nanjing Hongxu(Naniing red—bearded)；P．mumeNanjing Hong’(Nanjing red)；flow
er color；expression characteristics
M ei(P，．“n“s mume Sieb．et Zucc．)flower is
beloved deeply by Chinese for thousands of years
because of its beauty，purity and quality of resis—
ting cold．Today，it is one of the candidates of the
national flower of P．R．China． It is well known
that the scientific studies on Mei flower in China
have made splendent achievements and M ei flower
is the first horticuhural plant which is accredited
Chinese scientist，namely Chen Jun—Yu，as Interna—
tional Cuhivar Registration Authority(Chen，2002；
Zhao et a1．，2003)．
However，the study on the flower color of Mei
has been a blank all the while(Chen，2002；Zhang et
a1．，2003)．
The flower colors of Mei are very various，in—
cluding mauve，pink，pure white，greenish white，
light yellow or double color(Chen，200 1)and the
flower color is the most pivotal characteristics de—
termining the grace beauty and the aesthetic taste
of Mei．P．mume‘Nanjing Hongxu’(Nanjing red—
bearded)and P．muwle‘Nanjing Hong’(Nanjing
red)are the typical representatives of the mauve
and the pink respectively(Chen，1989)． W e have
found that the red flower colors of these two cuhi—
vars result from the existence of the anthocyanins
in their petals(Zhao et a1．，2004)．But nobody has
explored the variations of the flower colors of the
cultivars till today．
This paper deals with the temporal and spatial
variations of the flower colors of P．muDle‘Naniing
Hongxu’and P．；rlume‘Nanjing Hong’at flower de—
veloping stage and with the inserting site of the
flower in canopy． The research results could pro—
vide a reference or a premise for the aesthetic ap—
preciation of the flower color of Mei，the explora—
tion on the flower color mechanism of red M ei
flowers and the flower—selecting in cDNA cloning
of the key enzyme genes determining the biosyn—
thesis of anthocyanins in M ei flower．
1 Materials and methods
1．1 General
The layer classification of the petal was car—
ried out according to the concrete and natural collo—
caring site of the petal in the corolla．The open an—
gle of the petal in the corolla was defined as the de—
parture angle which was formed by the assumptive
connecting beeline of the petal tip and the inserting
point of the petal on receptacle vs the vertical axis
passing through the inserting point．
The relative content of the anthocyanins of
petal was mensurated with spectrophotometry．
Anthocyanins of the petals were extracted with
methanol containing 1 concentrated HC1(v／v)
(M arkham ，1982)． Extracts were diluted properly
with the same acidic methano1． A53o and A657 were
determined at room temperature in a I cm path—
length quartz cell using a Shimadzu UV—Vis spec—
trophotometer． The relative content of anthocya—
nin was calculated by a formula proposed by Rabi—
no and Mancinelli(1986)，namely(A53o一 0．25
A657)／g(FW)．
The content of total flavonoids of petal was
determined according to the method developed by
Zhao t a1．(2004)，and the formula(C一 0．074 9×
A 一 0．002 4)was selected as standard curve e—
quation． The malondialdehyde(MDA)content of
petal was determined with “Thiobarbituric acid
(TBA )method”(Li，2000)． 0．5 g petals were
ground with 5 mL phosphoric acid buffer(pH7．8)
on ice and the TBA reaction was terminated by icy
waterbath for 5 min． The absorbency was deter—
mined using above UV—Vis spectrophotometer．
A1l solvents used were of analytical grade．
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5期 赵昶灵等：梅花‘南京红须’、‘南京红’花色的呈现特征 483
1．2 Plant material
Flowers were obtained at Mei Flower Hill of
Sun Yat—sen Mausoleum Administrative Office of
Nanjing．The distance between plants of the trees
is 3 m X 3 m． The tree with well—balanced vigor
and inerratic canopy was selected as sampling tree．
On Feb 23 of 2004，j ust after the dew evaporated
completely，various flowers were selected and
picked according to different stages of flower de—
veloping and the Profuse flowers were picked ac—
cording to different inserting site of flower in cano—
PY．All flowers were encased i n yellow kraft enve—
lopes and immediately frozen at一2O～ 一22~C until a—
nalysis．
1．3 Temporal variations of the flower color：effects
of flower developing stage on the flower color
1．3．1 Partition of the flower developing stage
The developing course of M ei flower should be dif—
ferentiated according to 5 concrete indexes inclu—
ding transverse diameter of corolla，flower shape，
petal color，open angle of petal and stamen charac—
teristics．
1．3．2 Determination 0f the variations of flower
Table l Comparison of the visual characteristics
and P．7HuTHe‘Nanjing Hong’at
color From the outer to the inner，the petals of
outer，middle and inner layers of the flowers of va—
rious developing stages were peeled carefully．
Thereinto，the corolla of P．mume ‘Nanj ing
Hongxu’usually consists of 3 layers of petals，there
are 5 petals at outer，middle and inner layer respec—
tively．P．mume‘Nanj ing Hong’often possesses of
4 layers of petals． From outer to inner，the first
and forth layer were respectively regarded as“outer
layer’’and“inner layer”．and every layer composes
of 5 petals．The second and third layers were gen—
erally regarded as“middle layer”composing of 1 0
petals．The flower color variations were quantified
using the dynamic changes of the relative content
of anthocyanins of peta1． Synchronously，the con—
tents of total flavonoids and MDA of petals were
also determined．
1．4 Spacial variations of the flower color：effects of
the inserting site of flower in the canopy on the flow。
er color
The flower COlOrS of the Profuse flowers of
different inserting site of flower in canopy were also
determined by the relative content of anthocyanins．
of the flowers of P．7f／uTf／e‘Nanjing Hongxu’
different developing stages
2 Results and analyses
2．1 VisuaI characteristics of the flowers changed
with flower developing stages
The flower developing of P．mhtme‘Nanjing
Hongxu’and P．mume‘Nanjing Hong’could be di—
vided into 4 stages，namely Stage 工(Alabastrum
period)，Stage Ⅱ(Initial flower period)，Stage m
(Profuse flower period)and Stage IV(Final flower
period)(Table 1)，and the flower shapes are
“bead”，“cup”，“bowl’and“dish”respectively．
Thereinto，the integrated expression of flower shape
and flower color is the culmination at Stage m which
can fully unfurl the aesthetic merit of the cuhivar．
Though the transverse diameters of the corol—
las of P．mume‘Nanjing Hongxu’and P．mulne
‘Nanjing Hong’are almost the same at Stage IV，
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484 广 西 植 物 25卷
the splaying speed of the corolla of P．m TAme‘Nan一
；ing Hongxu’is quicker all the times(Fig．1)，show—
ing that the flower of P．mume‘Nanjing Hongxu’is
bigger and more flamboyant than that of P． “ P
cNanjing Hong’．At Stage IV，many of the petals
of outer layer of P．rluDIe‘Nanjing Hong’are prone
to slouch，droop and abscise．
3．5
3．0
一
g 2．5
2．0
8 1．5
U
1．0
0．5
O
I Ⅱ Ⅲ Ⅳ
D eveloping stage of flower
Fig．1 Changes of the transverse diameters of corollas
of P．，n埘，1e‘Nanjing Hongxu’and P．muT~e‘Nanjing
Hongat the flower developing stage(The transverse
diameter is the average value of 3O flowers)
2．2 Effects of flower developing stage oil the flower
color
2．2．1 Variations of the flower color at flower de—
veloping stages The flower colors of P． “
‘Naniing Hongxu’and P．rgluirle‘Nanjing Hong’all
reach the climax at Stage I and begin to fade
gradually soon after the petals spread despite the
possible second peak at Stage 1I(Figure 2，3 and
4)． It is speculated that the anthocyanins of the
petals start to decompose ceaselessly as soon as the
petals outspread because of light(Sweeny et n ．，
1 98 1：Rabino and M ancinelli，1 986；Beckwith et
nf．，2004)，which results directly in the fading of
t he flower color．
The change trends of the petal colors of differ—
ent layers of P．mume‘Nanjing Hongxu’at flower
developing stage are almost the same：The relative
contents of the anthocyanins of 3 layers are all the
strongest at Stage I，thin in some sort at Stage
1I．thicken appreciably at Stage Ⅲ and thin fur—
thest at Stage IV(Fig．2)．At Stage 1I andⅣ ，the
co1ors of middle and inner layers are feckly uni—
form． However，the petal color of outer layer is
much stronger than those of middle and inner lay
ers at different flower developing stages(Fig．2)．
I 1I Ⅲ IV
Developing stage of flow er
Fig．2 Variations of the petal color of the different
layers of P．DlUDle‘Nanjing Hongxu’
at flower developing stage
The color of the whole corolla of P．／T／urr／e
‘Nanjing Hongxu’is the strongest at Stage I，and
the color at Stage 1I is the next．The relative con—
tents of anthocyanins at Stage Ⅲ ，1I and Ⅳ are
86 、78％ and 63 0A of that at Stage I respectively
(Fig．3)．
Ⅱ Ⅲ Ⅳ
D eveloping stage of flower
Fig．3 Variations of the flower colors of P．171ulgle
‘Nanjing Hongxu’and P．171ulgle‘Nanjing Hong’at
flower developing stage(The、ralue of relative
content of the anthocyanins is the average
of those of three layers of petals)
The relative contents of the anthocyanins of 3
layers of P．ml／．me‘Nanjing Hong’are also the
strongest at Stage I．The change trends of the
petal colors of middle and inner layers at flower de。
veloping stages are almost the same：The relative
9 8 7 6 5 4 3 2 1 0
一每邑∞／l 《 0占竹 《一
曲a —a 西 uo 矗
utI o au aou u =
8 uu1 ．【 u∞．1u ∞C ．I|．
8 7 6 5 4 3 2 1 O
一事 一 ＼一卜 《 0．o竹 《一
曲a —a 西 uo 矗
u正 o au a0u u 0
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5期 赵昶灵等 ：梅花‘南京红须 ’、‘南京红 ’花色的呈现特征 485
contents thin at Stage II，thicken appreciably at
Stage 1I and thin furthest at Stage lV．But the rel—
ative contents of outer 1ayer decrease continuously
with the flower developing stages (Fig．4)． The
relative contents of anthocyanins at Stage 11，I
and IV are 75 、70 and 6l of that at Stage工．
As to the whole corolla，the chroma difference of
the flower color at 4 stages is not remarkable al—
though the color is also the strongest at Stage 工，
and the color at StageⅢ is the next(Fig．3)．
∞
C
C
叮
U
0
a：a
C
叮
3．O
2．5
2．O
1．5
1．O
O．5
O
I Ⅱ Ⅲ Ⅳ
Developing stage of flower
Fig．4 Variations of the petal color of the different
layers of P．7nulne‘Nanjing Hong’
at flower developing stage
At different flower developing stages， the
chroma differences of petal colors of different lay—
ers of P． “ ‘Nanjing Hong’and P．mume‘Nan—
jing Hong’are：outer layer> middle layer> inner
layer(Fig．2，4)．It has been found to be the most
important phenomena that the petals of the differ—
ent layers of one corolla of P．mufle‘Nanjing
Hongxu’or P． “ ‘Nanj ing Hong’are provided
with obvious chroma difference． However，the in—
serting sites of all petals of one corolla on the re—
ceptacle are almost located in the same plane．The
imbricate collocation of the petal in the corola re—
suits in the specific site where the peta1 is situated
in different layer and expresses the chroma differ—
ence．As a result，the concrete collocating site of
petal in the corolla is t he special characteristic of
petal developing，deciding how the petal expresses
specific chroma under t he control of internal and
external factors．
2．2．2 Variations of the content of tota1 flavonoids
of the petals at flower developing stage The con—
tents of tota1 flavonoids of 3 layers of petals of P．
mume‘Nanjing Hong’and P．mulTce‘Nanjing
Hong’all decrease with flower development，and
the decreases of the contents of tota1 flavonoids of
outer layers exist the“tardiness point”at Stage Il
(Fig．5)，which is consistent with the second peak
of the petal color of outer layers．It can be thought
that the changes of the contents of total flavonoids
of outer layers of P． ume‘Nanjing Hong’and P．
Figurine‘Nanjing Hong’are positively correlated with
the petal color changes of outer layers(Fig．2，5)．
This means that the flower colors of P． “7
‘Nanjing Hongxu’and P．murl~e‘Nanjing Hong’
probably result from the interaction of the antho—
cyanins and the coexisting copigments，e．g．flavo—
nols，because the flower colors of these two cuhi—
vars of M ei are mainly displayed by the petals of
outer 1ayers． On the other hand，the contents of
total flavonoids of middle and inner layers of two
cuhivars almost decrease point—blank(Fig．5)．
2．2．3 Variations of the M DA content of the petals
at flower developing stage The MDA contents of
3 layers of petals of P．wlurl~e‘Nanjing Hong’and
P．mMme‘Naniing Hong’all increase with the flow—
er development(Fig．6)，which implies that the pet—
als senesce as soon as they begin to splay although
their senescence is concealed by the flamboyant
red． Furthermore，the changes of M DA contents
also show that the senescence of the petals of outer
layers is much quicker，being evidenced by the rap—
id depigmentation of outer petals after Stage I
(Fig．2，4)． However，as to P．rlume‘Nanjing
Hong’，it is unknown why the senescence of the
petals of inner layer is quicker than that of middle
layer after Stage II(Fig．6，B)．
2．3 Effects of the inserting site of flower in the can-
opy on the flower color
The corolla chromas of the flowers inserting at
different sites of canopy of 尸．mume‘Nanjing
Hongxu’or P．ITgume‘Nanjing Hong’were not ac—
cordant(Table 2)．The higher the inserting site of
一≥邑 一 。《 0-0∞三 一
o c cou 一 对一
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486 广 西 植 物 25卷
the flower was，the thinner the flower color was．
The flower color
thinner than that
of the outer layer of canopy was
of the inner layer． However，va—
I Ⅱ Ⅲ Ⅳ
Developing stage of flower
I Ⅱ Ⅲ Ⅳ
Developing stage of flower
difference caused by the inserting site of flower in
canopy was not remarkable at all，showing that the
spatial variations of the flower colors are very
petit．It has been confirmed that light is the most
primary factor which influences the biosynthesis
and decomposition of anthocyanins(Sweeny eta1．，
1981；Rabino et a1．，1986；Beckwith eta1．，2004)．
7O
6O
5O
4O
3O
2O
1O
O
7O
喜6o
5O
o
吕 4O
呈3o
三2O
I Ⅱ Ⅲ Ⅳ
Developing stage of flow er
That M ei effloresces before leaves bourgeon leads
to the nicer and even conditions of natural ventila—
tion and daylight illumination in different sites of
one canopy，which results in the almost even con—
tents of anthocyanins in flowers owning different
inserting sites in the same canopy．
3 Discussion
The flower colors of P．mume‘Nanjing
一一≥邑∞＼一0目3《0苫 u 00
加 H 8 6 4 2 o
一≥邑∞＼一0目3《0苫Jo u 00
H 8 6 4 2 O
一≥邑 一0目3《0苫J0 u 00
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5期 赵昶灵等 ：梅花‘南京红须 ’、 ‘南京红’花色的呈现特征 487
Hongxu’and P．mume‘Nanj ing Hong’are mainly
under the control of flower developing，which is
accordant with the research result of Martin and
Gerats(1993)．The colors of the petals of different
layers or the whole corolla all change with flower
developing stages． The colors are the strongest in
Alabastrum period。thin in some sort in Initia1
flower period，thicken appreciably in Profuse flow—
er period and thin furthest in Final flower period e—
ven if the petals senesce as soon as they begin to
splay．At different flower developing stages，the
chroma differences of petal colors of different lay—
ers are：outer layer> middle layer> inner layer，
namely，the concrete collocating site of petal in the
corolla decides the specific color chroma of this
peta1．But the change trends of the petal colors of
different layers of one flower are not completely
similar．And the flower colors result from the in—
teraction of the anthocyanins and the coexisting
copigments． On the other hand，the flower color
difference caused by the inserting site of flower in
canopy is not remarkable at all，reflecting that the
spatial variations of the flower colors are very pot—
ty j ust because of the unconspicuous difference of
ventilation and illumination in different sites of one
canopy．
It is estimated that，as to M ei efflorescing in
early spring，the red flower color possesses of spe—
cial biological significance except luring pollina—
tors，namely it can strengthen the ability of M ei to
effloresce against cold．For many years，flower col—
or of higher plant is thought to lure pollinators，
such as insects and birds(Harborne，1973；Driveret
ⅡZ．，1998)．But it has been found that the stamens
and gynoecia of Mei flower are usually petalized
and the M ei which flower is appreciated specifically
by people is not generally fructiferous(Chen，1989；
Chen，200 1)． Furthermore，three remarkable phe—
nomena were observed in our study．(1)The flow—
er colors of P．mume‘Nanjing Hongxu’and P．
mume‘Nanj ing Hong’have been reached the climax
in Alabastrum period when the petals do not
spread at all，and begin to fade as soon as the petals
spread．So，the period when the flower color is the
strongest is not the period when the flower is wait—
ing for pollination． (2)After the petals outspread，
the petals of the outer layer often press close to
branch and can not become the first point of the
line of sight of animals．So，it is apparently impos—
sible that the remarkable flamboyant of the petal
color of the outer layer exists specialy because of
luring animals． (3)M ei flower is provided with
much more characteristics of anemophilous flow—
ers，e．g．the stamens protrude far-forth outside the
corolla after the petals spread．On the other hand，
it has been found that，in many plant tissues，the
structure and control genes controlling the biosyn—
thesis of anthocyanins can be induced to express by
low temperature (Christie et a1．，1994；Leyva et
ⅡZ．，1995；Shvarts et al。，1997)．Therefore，from
the evolutionary viewpoint，the production and tic—
cumulation of the anthocyanins in petals of M ei
may be the result of cold induction and selection of
fl very long period of time． As fl result，the red
flower color may play fl role in protecting the flow—
er of P．mume against cold in winter and early
spring(Driveret a1．，1998；Leng et a1．，2003)，
which directly determines the survival possibility
of this plant on earth．
The identIfications of the molecular structures
of the anthocyanins of the flower color pigments of
P．mume‘Nanjing Hongxu’and P．mume‘Nanjing
Hong’reveal that the key enzyme determining the
red flower color of Mei is flavonoid 3-hydroxylase
(F3 H )(Zhao et a1．，2004a，b)． It has been found
that the transcription of F3 H and the activitv of
F3 H in the flower of Petunia hybrida‘Old Glory
Red’(Brugliera et a1．，1999)and Toreni口hyb，．idⅡ
‘Summerwave Blue’(Ueyama et a1
． ， 2002)a11
reach the climax in Alabastrum period
． Therefore，
in cDNA—cloning of the F3 H of Mei with red
flower color，the flowers being in Alabastrum peri—
od should be the first choice
．
Acknowledgements
The authors thank M s． LIU Xue—fan，M s．LI—
ANG Ying—mei，M r．ZHAO Xing—fa and Mr．LIU
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488 广 西 植 物 25卷
Quan—long of the Research Centre of Met flower of
Sun Yat—sen M ausoleum Administrative Office of
Nanjing for the cuhivar—‘verifying and flower—‘col—-
lecting assistance．
References
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