全 文 ： 2007年 第5卷 第3期
世界竹藤通讯 WORLD BAMBOO AND RATTAN
Study on Flowering Characteristics of Phyllostachys
heteroclada Oliver in China
Wang Xiaohong1; Zhou Zuji2
(Forestry and Horticulture College, Sichuan Agricultural University,
Ya’an, China 625014)
Abstract: The flowering characteristics of Phyllostachy heteroclada Oliv were studied in this
paper. The result showed that the occurrence of bamboo flowering is random in a year, while
the florescence mainly happens in the period of November to next May, which led to the death
of bamboos. Inflorescence is represented as spike, each spathe possesses 5~7 fake pikel ts nd each
spikelet possesses 4~6 flor ts; the flower filament is about 13 mm long with canary anther and
trifid stigma. Anther is relatively rich, but pistil shows hypogenesis, so Phyllostachys heteroclada
Oliver up to now fail to seed. The further research showed that environment factors, such as
time of sun illumination, temperature and humidity of bamboo forest, had the significant impact
on bamboo flowering, and the anatomy of flowering and non-flowering bamboo leaf, stem and
root showed that the cell structure and development status of flowering`bamboo leaf,stem and
root are inferior to those of non-flowering bamboo. In present study, we suggested that the
interior physical change and external environment condition contributed side to the occurrence of
Ph. heteroclada Oliver flowering.
Key Words: Phyllostachys heteroclada; flowering characteristics; Phyllostachys; Ya’an
水竹开花的生物学特性研究
王小红 周祖基
(四川农业大学 雅 安 625014)
摘 要: 水竹，刚竹属散生竹种，为长江流域及其以南最常见的野生竹种之一，也是黄河和
长江流域广为栽培的一个竹种。自2003年以来，中国四川雅安地区的水竹出现开花现象，到目前
已成大范围开花枯死的趋势。对水竹的开花生物学特性进行了研究。结果显示，水竹开花在每个
季节均有发生，但主要集中在12月至次年的5月份，开花以后整个植株枯死。水竹为穗状花序，
每个佛焰苞（花序）有5~7个假小穗，每个假小穗有4~6朵小花；水竹花丝长约1.8cm，花药
黄绿色，柱头三裂。水竹花药非常丰富，但雌蕊发育不良，因此，至今未发现水竹结实。对开花
和未开花竹林环境因子进行调查和分析后发现，竹林内光照时数、气温以及湿度对开花有比较显
著的影响。同时，开花和未开花水竹根、茎、叶的冷冻切片显示，开花水竹根、茎、叶的细胞结
构和发育状况均差于未开花竹。由此看来，内部生理变化和外部环境影响是导致水竹开花的最终
原因。
关键词: 开花特性;水竹;刚竹属;雅安
1.Wang Xiaohong ,a doctor of Forestry and Horticulture College,
Sichuan Agricultural University in China
2.Zhou Zuji, a professor of Forestry and Horticulture College, Sichuan
Agricultural University in China
DOI:10.13640/j.cnki.wbr.2007.03.006
2007 Vol.5 No.3 13
ACADEMIC FIELD 学术园地
Introduction
Bamboo is one of the economically important
plants in south and South-East Asia. It has been de-
veloped and utilized as the food and manufacturing,
so being called as “poor man’s timber”. In China,
bamboo is more important for its use not only as
mentioned above but also the food of Giant Panda.
Bamboo flowering, which used to result in its death,
happens only once during the lifetime and most are
characterized by simultaneous flowering. It is nor-
mal for annual plant to die after flowering, however
both the flowering and death of bamboo are unusual
with the reason that most bamboo plant are long-lived
ones, being one of natures “century plants” (Numata,
1970). Although many bamboo species flower at long
intervals of up to 120 years, few studies have re-
vealed the flowering characteristics by observing.
Phyllostachys heteroclada Oliver, Gramineae,
Bambusoidease Nees, Phyllostachys Sieb. Et Zucc,
growing across the Yellow River valley, is the one of
dominantly agrestal bamboo in the Changjiang River
valley( Institutum of Kunmingense Academiae
Sinicae, 2003). The bamboo has high adaptability,
could grow in both the fertility and lean environment.
could breed up to highest value of dia. 5 cm and height
9 m by breeding artificially, and at the mean time
with many benefit characteristics, such as the resis-
tance for insect or disease, delicious taste of bamboo
shoots, higher production and broad useness(Jin Aiwu,
1999; Ou Jiande,2002). So Phyllos tachys
heteroclada Oliver are regarded as the important
germ plasm of bamboo locally.
With the onset of flowering in 2002,
Phyllostachys heteroclada oliver bamboo has be-
gun to flower in Ya’an district. This place is the home
town of Giant Panda, and the first Giant Panda was
found and named here. In Ya,an Bifengxia Base of
CCRCGP(The China Conservation and Research
Centre for Giant Panda ), there are abundant of
Phyllostachys heteroclada Oliver planted artificially
or wildly for years. They are main food resouce for
Giant Pandas of the base, especially in autumn and
winter. Since 2002, Phyllostachys heteroclada
Oliver has begun to flower individually, and flowered
and died over a large area by now, which led to the
acute reduction of Giant Panda`s food. On the other
hand, the co-existence of flowering and not-flower-
ing Phyllostachys heteroclada Oliver bamboo pro-
vides great test conditions. We observed and recorded
successively the flowering characteristics of
Phyllostachys heteroclada Oliver bamboo in this
paper, in order to discover some premonitory and rule
of this bamboo, which will be useful to more under-
stand the mechanism of bamboo flowering and take
strategy about how to restrain the event.
1 Materials and Methods
1.1 Materials
Ap prox ima tely 1 .0 ha of f lo weri ng
Phyllostachys heteroclada Oliver bamboo forest
near to the Ya,an Bifengxia Base of CCRCGP (The
China Conservation and Research Centre for Giant
Panda )was used for observation for flowering traits.
The bamboo in this site began to flower from Octo-
ber in 2003, and to now, 40% bamboo flowering, 30%
bamboo already died and about 30% bamboo still alive.
Since began flower, these bamboos were never cul-
tivated or managed. At the same time, 15 flowering
and 15 non-flowering Phyllostachys heteroclada
Oliver forest that near to the base were selected for
research of environment factors. The soil type of these
sites is country yellow-brown soil with good fertility.
Annual area rainfall is l 250 mm~l800 mm, the an-
nual average temperature is 14.1℃~18.8℃.
1.2 Methods
1.2.1 The observation for flowering traits The com-
bination of field survey and interview with local resi-
dent were employed during the period of nineteen
months, from November 2004 to June 2006. Four
flowering index, including the flower appearance time,
florescence, development of anther and seeding were
recorded by observing termly the growth of bamboo
in given area.
1.2.2 The research of environment factors in bam-
boo forest Aspect and illumination time: The box and
needle was employed to determine the aspect. The illu-
14 2007年 第5卷 第3期
世界竹藤通讯 WORLD BAMBOO AND RATTAN
mination time per day was recorded on sunshine days
in each sample area for three times successively and
the mean was calculated.
Soil temperature, moisture and pH value: At 9:00
and 14:00, the temperature of ground surface, under-
ground 10 cm, 15 cm and 20 cm were determined by
geothermometer at four random points in each sample
area. At the same time, the soil underground 0 to 20
cm from these points were taken to determine soil
moisture and pH value. It had two months to deter-
mine these index with the interval of fifteen days since
February seventh 2006.
The bamboo forest humidity and temperature:
At 9:00 and 14:00, the forest humidity and tempera-
ture was determined with the psychrometer and ther-
mometer at the interval of fifteen days since Febru-
ary seventh 2006.
1.2.3 The observation for apparatus cells structure
The leaves, young stems and root of bamboos were
collected from flowering and non-flowering bamboo
in September 2006. The specimens were fixed in FAA
(formalin-acetic acid-alcohol) for four weeks, then the
materials were transformed into sections with the thick-
ness of 12~18 μm by freezing microtome( American,
Reichert Histos TAT ) at -20℃, enveloped with anhy-
drous glycerin. The sections were observed with
microscope(Japan, Nikon ECLIPSE 50i) and taken
photo with digital imaging system ACT-2U.
2 Results
2.1 Flowering cycle
It has not been reported about the flowering
cycle of Phyllostachys heteroclada Oliver. The
population lived in Ya’an, ranging from seven to ninety
years old, were interviewed and no flowering events
were seen or heard. Based on the interview, it is sug-
gested that the flowering cycle was over eighty years.
2.2 Flowering time and florescence
The occurrence of bamboo flowering is random
in a year, while the florescence mainly happens in
the period of November to next May. Flower bulb
appears most densely in the period of December to
next March, and usually appears in the top of plant,
then spreads downwards to all body, the bud is ama-
ranth when young, then turn to brown-amethyst when
died (Fig.1 and Fig.2). The development of bulbs was
unconspicuous during the whole winter, but become
bloom after March. The spread of the pollen was
closely dependent upon the season and temperature.
In the other word, whenever the bulbs appearing, the
spread of the pollen mostly occurred in the period of
the last ten-days of March to the middle ten-days of
April. When anthers stick out of the palea, the spike-
let would dry rot soon, followed by the whole plant.
All the flowered plants would end up until the middle
ten-days of May. The whole florescence lasts out
for about six month. The bamboo may flower by ac-
cident in summer and autumn, namely from May to
November, that is only two percent of all the
population, and complete the flowering course at
shorter time about two months with the smaller spike-
let and fewer pollen.
2.3 The figure and structure of flowers
Based on the anthotaxy, that of Phyllostachys
heteroclada Oliver is racemose, three to five spathe
inserting together each main culm and minor branch
(Fig.2), broader and larger below, narrowest above,
without auricle , ligulate obvious; the bottom of spathe
bears a tuft of bracts with dense minutely pubescent
along the edges, each spathe possesses 1~2 fake
spikelets, each spikelet possesses 4~6 florets(Fig.3),
1~2 glumes and the size is not to a certainty, usually
smaller than lemma, menbrano-chartaceous, with long
pubes, apex appears like awn; lemma with long pubes
and many nerres. apex appears like short awn; the
lemma of first floret 12~14 mm long, the palea, sta-
men and pistil is very small because of hypogenesis;
paleas of other florets are shorter than lemmas, with
long pubes, apex bifid. When maturing, paleas and lem-
mas stretch, filament stick out, about 1.5cm long; an-
ther canary, stigma trifid. Anther is relatively rich, but
pistil hypogenesis(Fig.4), so we didn’t found
Phyllostachys heteroclada Oliver seed up to now.
2007 Vol.5 No.3 15
ACADEMIC FIELD 学术园地
Fig.1 Flower bulds
Fig.2 Flower branch, it is already dieda
Fig 3 Spathe and anther
Fig. 4 Floret, showed lemma and anthers
Fig. 5 Flowering branch, showed paleas and lemmas
already stretched, filament sticking out, and leaves are
still green
Fig. 6 Giant Pandas in the Ya,an Bifengxia Base of
CCRCGP is eating Phyllostachys heteroclada Oliver
16 2007年 第5卷 第3期
世界竹藤通讯 WORLD BAMBOO AND RATTAN
Tab.1 The aspect and illumination in flowering and non-flowering bamboo forest
type south slope north slope east and west slope illumination time (average)
(hour/day)
flowering 6 2 2 7.6
non-flowering 3 5 2 5.2
Tab. 2 The relationship between environmental factors and Phyllostachys heteroclada Olive flowering
time Type of Groud Underground Underground Underground Forest Forest Soil Soil
test site surface temp(10 cm) temp(15cm) temp(20 cm) temp humidity pH moisture
temp/℃ /℃ /℃ /℃ /% /℃ /%
9:00 flowering 5.25a 5.70a 5.24 a 6.08 a 10.6b 93.4c 7.15 a 23.16 a
Non- 4.72a 5.56a 5.00a 5.76 a 6.80 b 81.0c 6.47 a 32.98 a
flowering
14:00 flowering 15.72a 13.90a 13.58 a 13.84 a 20.94c 83.4c - -
Non- 14.60a 13.44a 12.88 a 13.10a 19.68c 65.8c - -
flowering
* Remark: a Means not significantly different P > 0.05;b Means mighty significantly different P < 0.01;c Means significantly different P
< 0.05.
2.4 The influence on flowering by environment
factors
It is found that the bamboos growing at the sun-
shiny area are more prone to flower than those grow-
ing at downsun area in the twenty areas researched
(Tab.1). The illumination time received by flowering
bamboo in sunshiny areas is 1.47 times to that in
downsun ones. Moreover, the bamboos grow greatly
and have no signal of flowering at some area where
sunshine seldom arrived. The relevance of flowering
and illumination is consistent with the results reported
about Fargesia robusta Yi and Fargesia denudate
Yi previously(Li Chengbiao,1997; Liao Zhiqin,1990).
Li suggested light was the most effective factors led
to the genesis and flowering of bud. It was found
that Fargesia denudate Yi ,the number of flowering
bamboo plants in some given area increased with
canopy density decreasing, and the flowering rate of
bamboo grown in less density environment was higher
( Liao Zhiqin,1990) . In this study, the fishscale bam-
boos in sunshiny area have superior vitality compared
with those in downsun area based on the number of
plants in given area, height and diameter of culms. It
is indicated that the fishscale bamboo is a species of
light-favored and shade-endurance.
The statistics also showed that bamboo flower-
ing had the significant correlation with the forest tem-
perature and humidity (Tab.2). Both the temperature
and humidity of flowering bamboo forest were higher
than that of non-flowering forest. The average tem-
perature of flowering forest was 1.5 times at 9:00
and 1.1 times at 14:00 relative to that of non-flower-
ing forest respectively. The humidity of flowering
forest increased by 12.4 percent at 9:00 and 17.4
percent at 14:00 compared with that of non-flower-
ing forest respectively. The other factors, like soil
temperature, soil pH and soil moisture, had no signifi-
cant effect to flowering. The soil temperature of flow-
ering forest, including ground surface, underground
10cm, 15cm and 20 cm temp. were higher than that
of non-flowering forest, while the soil moisture of
flowering forest was lower than that of non-flower-
ing forest. The soil character was neutral in flower-
ing forest and acidic in non-flowering forest. The
above statistics indicated that higher forest
temperature, humidity, soil temperature and lower soil
moisture led to bamboo flowering. On the other hand,
we found that the flowering of Phyllostachys
heteroclada Oliver have occurred far and wide over
the west of Sichuan basin. In despite of the influence
to flowering resulted from environment difference,
the environment factors only effect the occurrence
time of event, physiological mature and aging was
crucial factor about flowering.
2007 Vol.5 No.3 17
ACADEMIC FIELD 学术园地
Fig 7 Anatomy of flowering bamboo leaf
Fig 8 Anatomy of non-flowering bamboo leaf
2.5 Anatomical difference between flowering and
non-flowering bamboo
Epidermic cells of flowering bamboo leaves have
many enation, some interior structure cells rupture
and died, leaf histological differentiation is not very
obvious; cuticle of non-flowering bamboo leaves de-
velop better, the form of epidermic cells is more tidy
and compact (Fig7 and 8).
Young stem of non-flowering bamboo have regu-
lar construction, with big, dense inner vascular bundles
and small, sparse exterior ones, well-growth paren-
chyma cells and developed bundle-sheath, large-size
cross section of veins, tight and regular basic
framework, comparing to that of the flowering bam-
boos (Fig 9 and 10)
Radicel from flowering and non-flowering bam-
boo didn’t showed obvious differences between them.
The most distinctive feature in this organ was the
vascular bundles of little roots of flowering bamboos
are scelerenchyma, and more ageing than that of non-
flowering (Fig11and 12)
Fig 9 Anatomy of flowering bamboo young stem
Fig 10 Anatomy of non-flowering bamboo young stem
Fig 11 Anatomy of flowering bamboo radicel
Fig 12 Anatomy of non-flowering bamboo radicel
18 2007年 第5卷 第3期
世界竹藤通讯 WORLD BAMBOO AND RATTAN
gP
3 Discussion
Based on the interview and observation, the
flowering cycle of Phyllostachys heteroclada Oliver
is about eighty years, the occurrence of bamboo flow-
ering is random in a year, while the florescence mainly
happens in the period of November to next May,
which led to the death of bamboos. Inflorescence
determinate, racemose, each spathe possesses 1~2
fake spikelets, each spikelet possesses 4~6 florets,
1~2 glumes; filament about 13 mm long, anther canary,
stigma trifid. Anther is relatively rich, but pistil
hypogenesis, so we didn’t found Phyllostachys
heteroclada Oliver seed up to now.
The further research showed that environment
factors, such as time of sun illumination, temp and
humidity of bamboo forest, had the significant rel-
evance with bamboo flowering, that higher forest
temperature, humidity, soil temperature and lower soil
moisture led to bamboo flowering.
We also found that there is much difference
between flowering and non-flowering bamboo inte-
rior structure, the anatomy of flowering bamboo leaf,
stem and root is all more inferior than non-flowerings.
Our research showed that the interior physical trans-
formation and external environment condition con-
tributed together to the occurrence of bamboo
flowering.
Acknowledgements
We are thankful to Mr. Huang Yan and Wei
Rongpin (Directors, Ya’an Bifengxia Base of
CCRCGP )for encouragement, and Wang Hanyan
(Sichuan Agricultural University, China )for her help
during the study.
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埃塞俄比亚的竹业发展状况
品和家具制作的培训由 ZOA避难所实施，亚的
斯 亚 贝 巴 的 联 邦 中 小 型 企 业 发 展 局 和
Benishangul-Gumuz地区的中小型企业发展局，
培训项目包括竹加工，乡村技术促进中心和
Assosa技术与职业研究所，他们有竹加工机械
可用于培训。另外，CIDA联合创办的食品安全
项目在竹产品开发方面也有很大的潜力。
——国际竹藤组织
国外动态
在埃塞俄比亚的Benishangul-Gumuz地区，
大约有60万人口，分布在3个管辖区，20个行
政区。它是埃最穷且食品最无保障的地区之一，
贫穷的范围是本地的50%。但它的竹资源丰
富，仅滇竹就有44 万 hm2,主要用于房屋，篱
笆、厨房用品、农用和竹笋，中小型企业在开发
竹胶合板方面潜力很大。一些投资与技术能力
已经存在，如土地与海洋发展公司实行租借的
方式已投资1.36亿，用于竹杆的出口。手工艺