全 文 ：The Flowering Biology of Plum (Prunus
domestica L.) in Aksu Region, Xinjiang
Beibei FENG1, Wenwen LI1, Yajun WEI1, Yeyong XU2, Ming WANG2, Hongli YANG2, Yingying
NIU1, Kang LIAO1*
1. Research Center for Xinjiang Characteristic Fruit Tree, Xinjiang Agricultural University, Urmqi 830052, China;
2. Aksu Jamu Experiment Station, Academy of Forestry Science in Xinjiang, Urumqi 843100, China
Supported by High-quality and Highly-efficient Cultivation Technology Promotion of
Prunus simonii.
*Corresponding author. E-mail: 13899825018@163.com
Received: April 3, 2016 Accepted: May 16, 2016A
Agricultural Science & Technology, 2016, 17(6): 1371-1375
Copyright訫 2016, Information Institute of HAAS. All rights reserved Agronomy and Horticultrue
P runus domestica L. has sever-al aliases names as Damson,Yangli, Prune (USA) and
Suanmei (Xinjiang) that leaves small
tree, which belongs to the genus of
Prunus and family of Rosaceae.
Prunus domestica has more than 2000
cultivars that originating in Yili, Xin-
jiang, West Asia and Europe and other
places [1]. In recent years, Xinjiang has
introduced several excellent plum va-
rieties as Ximei, which has become a
favorite of costumers with their good
qualities as big fruit, bright color, rich
flavor etc. Prunus domestica fruit in
addition to fresh marketing, can be
made into jam, fruit juice, candy etc.
The investigation on flowering biology
of Prunus domestica in Xinjiang arid
area can provide a theoretical basis for
the high quality and yield cultivation.
Zhu et al. [2] found that Prunus domes-
tica has the strong comprehensive re-
sistance, followed by P. cerasifera am-
ong, P. cerasifera, P. ussuriensis, P.
nibs, P. salicina, P. spinosa and Nai
Plum. Sun et al. [3] studied the genetic
relationships in prunus domestica in
xinjiang using ISSR markers, mainly
discusses the pollen morphology and
Abstract [Objective] The aim of this study was to understand the flowering biologi-
cal characteristics of different varieties Prunus domestica L. in Aksu Jiamu Experi-
ment Station. [Method] Three varieties of plum as Prunus domestica L. cv.
‘French’, Prunus domestica L. cv. ‘Victoria’ and Prunus domestica L. cv. ‘Stan-
ley’ were the research subjects. The phenophases of blooming, and the morpholog-
ical and developmental characteristics of floral organs were observed. The amount
and viability of pollen and the stigma receptivity were measured. [Result] Three va-
rieties of plum all blossomed in early April, and the florescence had lasted for 9-10
d. The average number of anthers was 28.48% . The average number of pollen
grains in each flower was 28 700.0. The pollen germination rate was 27.4% .
86.75% of the flowers were medium-styled or long-styled. There was no significant
difference in the petal shape, color, anther number, calyx color, and flower ovary
among the three varieties of plum. The stigma receptivity was strongest on the first
day of flowering, and then dropped gradually. Therefore, the flowers of plum were
manual pollinated on the first day of flowering. The duration time of stigma recep-
tivity of Prunus domestica L. cv. ‘Victoria‘ was longer than the other cultivars, the
duration time of stigma receptivity was about 5 d. [Conclusion] The pollen number
and pollen germination rate of Prunus domestica L. cv. ‘Stanley’ were higher than
the other cultivars, which be considered the most suitable pollinizer for plum. In
contrast, the complete flower rate of Prunus domestica L. cv. ‘French’ and Prunus
domestica L. cv. ‘Victoria’ was relatively high, but the pollen number and pollen
germination rate of Prunus domestica L. cv. ‘French’ and Prunus domestica L. cv.
‘Victoria’ were lower than Prunus domestica L. cv. ‘Stanley’, which are not suit-
able to use as a pollinizer for plum.
Key words Prunus domestica L.; Phenophase; Floral organs；Pollen viability; Stigma
receptivity
阿克苏地区欧洲李开花生物学
特性研究
冯贝贝 1，李雯雯 1，魏雅君 1，徐业勇 2，王明 2，杨
红丽 2，牛莹莹 1，廖康 1* （1.新疆特色果树研
究中心，新疆农业大学，新疆乌鲁木齐 830052；
2. 新疆林科院佳木试验站 ， 新疆阿克苏
843100）
摘 要 [目的]探讨引进欧洲李品种在阿克苏
的开花生物学特性，为欧洲李的丰产优质栽培
奠定基础。 [方法]调查‘斯坦勒’、‘法兰西’、‘女
神’3 个欧洲李品种的在阿克苏地区开花物候
期、花器官形态特征、花粉量、花粉活力和柱头
可授性。 [结果]3 个欧洲李品种的花期集中在 4
月初，花期持续时间为 9~10 d，单花花药数平
均 28.48 枚/蕾， 单花花粉量平均值是 28 700.0
粒，花粉萌发率平均是 27.4%，中长花柱花平均
比率为 86.75%；3 个品种欧洲李的花瓣形状 、
颜色，花药、花萼颜色子房着生方式和花类型
都基本上一致。 3 个欧洲李品种在开花当天柱
头可授性变强，之后渐渐变弱。 ‘女神’的可授
时期比较长，可持续 5 d。 [结论]‘斯坦勒’的单
花花粉量和花粉萌发率较高，比较适合作为授
粉树。 而‘女神’和‘法兰西’的完全花比例都比
较高，单花花粉量或者花粉萌发率较低 ，不适
合作为授粉树。 3 个品种的花期相近，‘斯坦勒’
可为其他 2 个品种授粉。
关键词 欧洲李；物候期；花器官；花粉；柱头
可授性
基金项目 杏李优质高效栽培技术示范与推
广。
作者简介 冯贝贝 （1990-），女 ，河南驻马店
人，在读硕士研究生，研究方向为果树栽培与
生理 ，E-mail：1527274279@qq.com。 *通讯作
者，廖康（1962-），男，四川梓橦人，教授，博导，
主要从事果树种质资源及栽培生理研究 ，E-
mail：13899825018@163.com。
收稿日期 2016-04-03
修回日期 2016-05-16
DOI:10.16175/j.cnki.1009-4229.2016.06.021
Agricultural Science & Technology 2016
relationship problems. At present,
many domestic and international
scholars carried on a great deal of
empirical research to cultivation man-
agement, pruning and training [4-5], but
less studies on the flowering biology of
Prunus domestica [6-7]. Therefore, this
study observe the flowering phenolo-
gy, pollen number, pollen viability and
stigma receptivity of Prunus domestica
in Akesu area, to provide a theoretical
basis for the selection of optimal polli-
nation period and pollinizer for the
production practice and further re-
search of Prunus domestica.
Material and Methods
The profile of experimental field
The experiment was conducted at
the Jiamu experiment station, affiliated
with Academy of Forestry Science in
Xinjiang, which is located in the Jiamu
Forest Farm in Wensu county, Xin-
jiang Aksu prefecture. The farm (N 41°
19＇08.27＂，E80°31＇00＂) is an altitude of
1103.8 m. Under the control of the typ-
ical continental arid desert climate,
rainfalls are rare, and there is big dif-
ference between day and night. The
seasons are uneven, with a short,
windy and cool spring, and a hot and
dry summer. The average rainfall is
65.4 mm per year with a large inter-
annual variability, while the average
annual evaporation is 1 883.6 mm.
The annual average temperature is
10.1 ℃, with an extreme low tempera-
ture at -27.4 ℃ , and the effective ac-
cumulated temperature is 2 916.8 -3
198.6 ℃.The frost-free period is 195d,
and the annual average sunshine
counts 2 747.7 hours [8]. The three test
varieties are Prunus domestica L. cv.
‘Victoria’ [9], Prunus domestica L. cv.
‘Stanley’ [10], and Prunus domestica L.
cv. ‘French’[11] and Prunus salicina L.
cv. ‘Meilili’ [1] servers as the control.
The grafts are peach rootstocks, which
more than 6 years old with a moderate
growth rate, and are under middle-
level management.
Experimental methods
The observation of blooming
phenophase
The determination of plum’s
blooming phenophase was based on
Li’s standards[12]. We monitored the fl-
ower bud expanding period (about
25% of the flower buds of the entire
tree began to expand, and the scales
became staggered), the period of re-
vealed calyces (calyx came out from
the tip of the scale), the period of re-
vealed petals (petal came out from the
calyx), the initial bloom stage (about
5% of the flowers of the entire tree be-
gan to bloom), the full-bloom stage
(about 25% , 50% and 75% of the
flowers of the entire tree bloomed for
the beginning, middle and ending of
the full-bloom stage), the flower drop
period (about 25% of the flowers start-
ed falling for the beginning of the peri-
od, and more than 95% of the flowers
started falling for the end of the flower
drop period).
The morphological characteristics
of the floral organ
One hundred flowers were ran-
domly selected from each sample. Ac-
cording to the morphology or the rela-
tive position of the pistil stigma to the
anther, the floral organs were charac-
terized to four developmental styles:
the long style (♀＞♂: the pistil stigma
was higher than the position of anther),
the middle style (♀＝♂: the pistil stig-
ma was equal to the position of an-
ther), the short style (♀＜♂: the pistil
stigma was shorter than the position of
anther), and the pistil-degenerated
style (♀≈0: the pistil stigma was
shorter than the position of filament or
the pistil was undeveloped).The shape
and color of petals, the color of calyxes
and anthers, as well as the position of
ovaries were observed, and the corolla
diameter, the vertical and horizontal
diameter of the petal and anther, the
length and width of the sepal and ov-
ary, the style length and filament
length ,were also measured from 10
flowers of each variety[10].
The measurement of pollen amount
The amount of pollens was
measured with the method intro-
duced by Sun[14]. Ten uncracked flow-
er buds in the balloon stage were col-
lected from each sample, and the
number of anthers were counted for
each flower bud. Ten anthers of each
bud were collected in a 1.5 ml volume
tube. After adding 1 ml distilled water,
the tube was centrifuged for 3 min at
10 000 r/min then mixed by vortexing.
One μl mixture was pipetted onto a
glass slide, and the number of pollen
was counted under microscope (Motic
BA400-102M).
The pollen amount of single flower
(Grain/flower) = (Total number of
pollen grains in each 1 μl solution
drops×1 000)
The measurement of pollen viability
The pollen viability was measured
according to Zhang’s method [15]. Un-
cracked flower buds in the balloon
stage were collected from each sam-
ple, and the anthers were stripped off
and spreaded in acid paper boxes and
dried for 1-2 d then stored in -20 ℃ .
Naturally powdered pollens were em-
bedded in agar (0.01% boric acid +
10% sucrose + 1% ager ) [16] and incu-
bated at 25 ℃ for 4 h. The numbers of
all pollens and germinated pollens
were counted under microscope (Mot-
ic BA400-102M). The pollen germina-
tion rate of each treatment was calcu-
lated with the following formula:
Pollen germination rate (% ) =
(Numbers of germinated pollen in each
horizon/Total number of pollen in each
horizon)×100
The measurement of stigma recep-
tivity
Flowering branches were collect-
ed from each sample and cultured in
water. Stigmas were taken on the day
before the day of flowering (-1 d), the
day of flowering (0 d), and 1, 2, 3, 4 or
5 d after flowering (1, 2, 3, 4, or 5 d).
The collected stigmas were evenly
placed on a concave slide separately
with the reaction solution of Ben-
zedrine and Peroxide (1% Benzedrine
and 3% Peroxide mixed with distilled
water in 4:11:22 proportion). After 10
min of reaction, the stigmas were ob-
served with a Shunyu EX-204 micro-
scope and the staining was ranked to
one of the 5 levels as weak (-), normal
(+/-), slightly strong (+), strong (++)
and extremely strong (+++) according
to the blue-black part of the stigma and
the number of vacancies on the stig-
ma. The stigma quantity was recorded
at different levels[17].
Data statistics
The data was analyzed with Mi-
crosoft Excel 2007 and SPSS19.0.
Results and Analysis
The blooming phenophases
The results were as shown in
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Agricultural Science & Technology2016
Table 1 Blooming phenophase of experimental materials
Variety Date ofgermination Sepal showing Petal baring Initial blooming Full blooming Blossom falling
Blooming still
time//d
‘Stanley’ 3/30 4/4 4/7 4/8 4/12 4/17 10
‘French’ 3/26 4/3 4/6 4/8 4/10 4/16 9
‘Victoria’ 4/2 4/5 4/9 4/10 4/12 4/19 10
‘Meilili’ 3/4 3/28 3/30 4/2 4/3 1/10 9
Table 1, for the variety of Prunus do-
mestica L. cv. ‘Stanley’, the germi-
nation was on March 30th, the sepal
showing on April 4th, the petal baring
on April 7th, the initial blooming on April
8th, the full blooming on April 12th, the
start of blossom falling on April 17th,
and the blooming lasted for 10 days.
For the variety of Prunus domestica L.
cv. ‘Victoria’, the germination was on
March 30th, the sepal showing on April
5th, the petal baring on April 9th, the ini-
tial blooming on April 10th, the full
blooming on April 12th, the start of blos-
som falling on April 19th, and the
blooming still time was 10 d. For the
variety of Prunus domestica L. cv.
‘French’, the germination was on
March 26th, the sepal showing on April
3th, the petal baring on April 6th, the ini-
tial blooming on April 8th, the full
blooming on April 12th, the start of blos-
som falling on April 17th, and the
blooming still time was 10 d. The
Prunus domestica L. cv. ‘Victoria’
buds sprouted earlier than Prunus do-
mestica L. cv. ‘Stanley’ and Prunus
domestica L. cv. ‘French’ by 4-6 d.
Due to the temperature rise on April
10th, Prunus domestica L. cv.‘French’
buds swelled rapidly, and showed
blooming in the afternoon, and Prunus
domestica L. cv. ‘Stanley’ fully
bloomed on the same day. The germi-
nation of all three varieties was be-
tween March 30th and April 2th, and the
flowering was relatively concentrated
between April 8th and April 19th, while
the flowering duration was 9 -10 d,
thus they can be mutually pollinated.
Prunus salicina L. cv. ‘Meilili’ flowered
earlier than the three varieties by 6-8
days.
The characteristics of the floral or-
gan
Comparison of the developmental
characteristics of the floral organ
The results were as shown in
Table 2, the variety of Prunus domes-
tica L. cv. ‘Stanley’ had the lowest
rate of “perfect flower” with the pistil
stigma was higher than or equal to the
position of anther about 61%, while the
varieties of Prunus domestica L. cv.
‘French’, Prunus domestica L. cv.
‘Victoria’ and Prunus salicina L. cv.
‘Meilili’ all had a proportion more than
90% . For Prunus domestica L. cv.
‘Stanley’, about 37% of flowers had
small stigma (short style). For Prunus
salicina L. cv. ‘Meilili’, the majority
(89% ) of flowers was middle style,
while for all the other three varieties
the long style flowers were much more
than the middle style.
The comparison of the floral organ
The results were as summarized
in Table 3, Prunus domestica L. cv.
‘Stanley’ and Prunus domestica L.
cv. ‘Victoria’ had oval-shaped petals,
while the petal shape of Prunus do-
mestica L. cv. ‘ French’ was ovoid. All
varieties had univalve flowers and
white petals. Prunus domestica L.cv.
‘French’ had light yellow anthers, and
the others had yellow anthers instead.
The calyx color was yellow-green for
Prunus domestica L. cv. ‘French’,
light green for Prunus domestica L. cv.
‘Stanley’, and green for Prunus do-
mestica L. cv. ‘Victoria’ and Prunus
salicina L. cv. ‘Meilili’. The ovary had
an upper position for all varieties. The
number of stamens was between 24
and 32. Prunus domestica L. cv.
‘French’ had the largest corolla diam-
eter of 25.59 mm. The stamen length
and the style length were in the ranges
of 5.5 -7.5 mm and 8 -9.5 mm, re-
spectively. The styles length had little
variance. Prunus domestica L. cv.
‘Stanley’ had the longest and widest
anthers, while Prunus domestica L. cv.
‘French’ had the longest and widest
calyxes. There was little difference in
ovary size, with the length generally
around 1.5 mm, and width between
1.2 and 1.5 mm.
The characteristics of pollens
The amount of pollen
The results were summarized in
Table 4. For Prunus domestica L. cv.
‘Stanley’, Prunus domestica L. cv.
‘French’, and Prunus domestica L.
cv. ‘Victoria’, the number of anthers
per flower was 31.8, 29.9 and 24.9
pieces/bud, respectively, and for
Prunus salicina L. cv. ‘Meilili’ the
number was 27.3 pieces/bud. The av-
erage number of anthers per flower
was 28.48 pieces/bud. Prunus do-
mestica L. cv. ‘Stanley’ had a pollen
amount of 50 044.4 grains/flower,
which was significantly higher than
those of Prunus domestica L. cv.
‘Victoria’, Prunus domestica L. cv.
‘French’ and Prunus salicina L. cv.
‘Meilili’. The average amount of
pollen was 28 700 grains/flower.
The comparison of pollen viability
The results were as shown in
Table 4. there was large variance in
the pollen germination rate. Prunus
Table 2 Comparison of developmental characteristics of floral prganogenesis in plum varieties
Variety
Perfect flower//% Incomplete flower//%
♀>♂ ♀=♂ Total ♀<♂ ♀≈0 Total
‘Stanley’ 43 18 61.00 37 2 39.00
‘French’ 94 0 94.00 2 4 6.00
‘Victoria’ 95 5 100.00 0 0 0.00
‘Meilili’ 3 89 92.00 8 0 8.00
Average 58.75 28.00 86.75 11.75 1.50 13.25
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Agricultural Science & Technology 2016
Table 5 Comparison of stigma receptivity of tested varieties
Variety
Flowering process
-1 d 0 d 1 d 2 d 3 d 4 d 5 d
‘Stanley’ + + ++ + +/- +/- +/-
‘French’ +/- + + +/- +/- - -
‘Victoria’ +/- + + + ++ + +/-
‘Meilili’ + + ++ ++ + + +/-
weak (-), normal (+/-), slightly strong (+), strong (++) and extremely strong (+++)
Table 4 Comparison of anther number, pollen amount and pollen germination rate among tested varieties
Variety The number of single flower anther(Pieces/bud) The pollen amount of single flower(Grain/flower) Pollen germination rate//%
‘Stanley’ 31.8±0.49 aA 50 044.1±5 671.11 aA 42.4±0.24 aA
‘French’ 29.9±0.547 bA 31 220.0±3 295.92 bB 7±0.32 dD
‘Victoria’ 24.9±0.547 dD 14 490.0±2 327.11 cC 27.4±0.51 cC
‘Meilili’ 27.30±0.578 cC 21 180.0±2 309.05 bcBC 32.8±1.62 bB
Average 28.48 28 700 27.4
Table 3 Comparison of characteristics of the flower-organ of plum varieties
Variety Shape of petal Petal color Anther color Calyx color Ovaryposition
Type of
flower
Diameter of
corolla//mm
Petal length
mm
Petal width
mm
‘Stanley’ ovoid in shape white light yellow light green upper univalve 25.59 12.09 9.18
‘French’ ovoil white yellow yellow-green upper univalve 26.1 11.61 7.33
‘Victoria’ ovoil white yellow green upper univalve 18.6 8.39 6.67
‘Meilili’ ovoil white yellow green upper univalve 18.83 8.71 6.55
Variety Stamen lengthmm
Style lengh
mm
Anther
length//mm
Anther width
mm
Calyx length
mm
Calyx width
mm
Ovary length
mm
Ovary width
mm)
Stamen
number
‘Stanley’ 6.03 9.03 1.34 1.03 4.7 3.12 2.37 1.35 30
‘French’ 7.23 8.57 1.35 1.23 3.94 2.31 2.11 1.32 32
‘Victoria’ 5.98 9.06 1.14 0.83 4.5 2.56 1.58 1.27 24
‘Meilili’ 6.11 7.55 0.8 0.71 4.22 2.62 1.39 1.04 27
domestica L. cv. ‘French’ had the
lowest pollen germination rate of 7% ,
while the average rate was 27.4% .
Prunus domestica L. cv. ‘Stanley’
had significantly higher success in ger-
mination (42.4%) than the other three
varieties.
The observation of stigma receptiv-
ity
The stigma receptivities were
summarized in Table 5. Stigmas from
all tested plum varieties had shown
some degree of receptivity on the day
before the day of flowering, among
which the receptivity of Prunus do-
mestica L. cv. ‘Stanley’ was stronger
than those of Prunus domestica L. cv.
‘French’ and Prunus domestica L. cv.
‘Victoria’. From 1 d before flowering
to 4 d after flower, these three varieties
did not show quite strong flowering.
Only on the day of blooming, Prunus
domestica L. cv. ‘Stanley’ exhibited
strong receptivity, but on the third day
there was merely partial receptivity.
The receptivity of Prunus domestica L.
cv. ‘Victoria’ lasted for 5 d, which was
the longest among the three tested
varieties; Prunus domestica L. cv.
‘French’ bloomed on the fourth day,
but had no longer receptivity by the
fifth day. The receptivity of Prunus
salicina L. cv. ‘Meilili’ was stronger
than the other three varieties, and last-
ed from the day before flowering until
four days after the flowering.
Discussion
The flowering of Prunus domesti-
ca is significantly later than P. salicina,
P. simonii, P. usssuriensis, P.
cerasifera, P. nibs, P. americana, etc,
which provides an advantage to avoid
the hazard of late frost and spring cold
spell[1]. In spring, it is always windy and
dusty in Aksu region. The florescence
of Prunus domestica in days of strong
wind may result in poor pollination be-
cause of reduced insect activity and
air-dried stigmas [18]. Measures should
be taken after strong winds to prevent
stigma drying or being blocked by
sand, so that the pollens could germi-
nate.
Zhang et al [15]. studied the differ-
ences of pollen content, pollen germi-
nation and growth among eight
species of fruit trees, and the result
shown that for different species or va-
rieties of fruit trees, the amount of pol-
lens per flower is different, and several
significant differences are consistent
with our findings in the three varieties
of Prunus domestica. Chankun [19]
studied using the pollen variables to
predict the yield of crop, which con-
firmed a significant correlation be-
tween the amount of pollen and the
yield of product. One of the main
causes of low fruit set might be poor
pollen germination rate, which de-
pends not only on the genetic char-
acteristics of the variety, but also on
the nutritional status of the tree and
the development of flower buds,
which are affected by climatic condi-
tions, cultivation and management
levels and other relevant factors [20]. In
order to ensure Prunus domestica pol-
lination, that to obtain the expected
yield and quality, the cultivation and
management should be improved,
good varieties should be chosen for
high pollination.
Stigma is an important part of the
1374
Agricultural Science & Technology2016
Responsible editor: Xiaoxue WANG Responsible proofreader: Xiaoyan WU
floral organ for flowering plants. The
development degree of stigma affect-
ed the germination of pollen grains,
which was related to the fertilization
and fruit setting. In addition to inter-
cepting pollens and providing a breed-
ing ground for pollen, it has an affinity
for pollen[21]. Sun[14] et al. found that the
stigma receptivity of apricot varieties
was viable from one day before the
flowering to six day after flowering, in
the first day of flowering the stigma
receptivity was strong[14]. This study sh-
owed that the stigma receptivity of
Prunus domestica was strong at one
day before the flowering and the first
day of flowering, i.e. these days were
most efficient days for pollination.
Thus, to improve fruit set percentage
by ensuring the pollination effect, the
one day before the flowering or the
first day of flowering is better to
choose to cross-pollination on the pro-
duction practice.
Conclusion
The complete flower rate of
Prunus domestica L. cv. ‘French’ and
Prunus domestica L. cv. ‘Victoria’
was relatively high, but the pollen
number and pollen germination rate of
Prunus domestica L. cv. ‘French’ and
Prunus domestica L. cv. ‘Victoria’
were lower than Prunus domestica L.
cv. ‘Stanley’, which are not suitable to
use as a pollinizer for Prunus domesti-
ca. In contrast, the pollen number and
pollen germination rate of Prunus do-
mestica L. cv. ‘Stanley’ were higher
than the other cultivars, which be con-
sidered the most suitable pollinizer for
Prunus domestica.
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