全 文 ：一种特殊的长寿细胞 : 毛竹茎秆纤维细胞?
甘小洪1 , 丁雨龙2
(1 西华师范大学生命科学学院 , 珍稀动植物研究所 , 四川 南充 637002;
2 南京林业大学竹类研究所 , 江苏 南京 210037 )
摘要 : 利用显微和细胞化学方法 , 对毛竹 ( Phyllostachys edulis) 茎秆纤维次生壁形成过程中超微结构变化
以及 ATP酶、Ca2+ -ATPase和酸性磷酸酶的超微细胞化学定位进行了研究。研究发现 , 次生壁形成早期 ,
细胞核具有双层核膜 , 染色质凝聚 , 可见大量的线粒体、粗面内质网和高尔基体等细胞器存在于纤维细胞
中 ; 随后 , 双层核膜消失 , 细胞器将逐渐解体 , 多泡体开始出现在纤维细胞的细胞质 ; 随着年龄的增加 ,
纤维细胞壁逐渐增厚 , 并出现多层结构现象 , 而运输小泡、细胞膜、胞间连丝和凝聚的染色质将持续存
在。在次生壁形成的整个过程中 , ATP 酶、Ca2+ -ATPase和酸性磷酸酶在运输小泡、细胞膜、质膜内陷、
胞间连丝和凝聚的染色质中将持续存在。结果表明 , 毛竹茎秆纤维细胞是一种不同于木本双子叶植物的长
寿细胞 , 纤维原生质体中 ATP 酶和酸性磷酸酶的持续存在与次生壁的持续增厚密切相关。
关键词 : 纤维 ; 长寿细胞 ; 毛竹 ; 次生壁 ; 超微细胞化学定位 ; 超微结构
中图分类号 : Q 942 文献标识码 : A 文章编号 : 0253 - 2700 (2008) 02 - 151 - 08
A Special Long-Lived Cell: the Culm Fiber Cell
of Phyllostachys edulis (Gramineae) *
GAN Xiao-Hong1 , DING Yu-Long2
(1 Collegeof Life Science, Instituteof Rareand Precious Animals and Plants,
China West Normal University, Nanchong 637002 , China;
2 Bamboo Research Institute, Nanjing Forestry University, Nanjing 210037 , China)
Abstract : With several microscopic andcytochemical methods, the ultrastructuremodification and ultracytochemical local-
ization about some enzymes like H
+
-ATPase, Ca
2 +
-ATPase and acid phosphatase (APase) of fiber in Phyllostachys edulis
culms during secondary wall formation were studied . Early, intact doublekaryotheca remained infiber nucleus, while nu-
clear chromatin agglutinated . A lot of organelles such as mitochondria, rough endoplasmic reticula and Golgi bodies were
observed in fiber cell . Then, double karyotheca disappeared, and the organelles disintegrated . Multivesicular bodies ap-
peared in fiber cytoplasm . With thefurther development of fiber, fiber wall underwent continual thickening withaging, and
polylaminate structure gradually appeared . While the agglutinated nucleus, transfer vesicles, plasma membrane and plas-
modesmata still presented . During the secondary wall formation of fiber, H+ -ATPase, Ca2 + -ATPase and APase also sus-
tainablely remained in plasmamembrane, plasmamembrane invagination, plasmodesmata, transfer vesicles and agglutinat-
ed nucleus . The results demonstrated that (1) the fiber cell of P . edulis culm is a special long-lived cell, which differs
from that of woody dicotyledons, and ( 2) thesustainable persistenceof ATPaseandAPase infiber protoplast is closely cor-
related to thecontinual thickening of secondary wall with ageing .
Key words: Fiber; Long- lived cell; Phyllostachys edulis; Secondary wall ; Ultracytochemical localization; Ultrastructure
云 南 植 物 研 究 2008 , 30 (2) : 151～158
Acta Botanica Yunnanica

? ?Foundation items: TheStartup Fund for Scientific Research of China West Normal University ( No . 05B035) and theNational Natural Science Fund
of China (No . 30271064 )
Received date: 2007 - 07 - 23 , Accepted date: 2007 - 10 - 18
作者简介 : 甘小洪 (1974 - ) 男 , 汉族 , 副教授 , 主要从事发育植物学研究。
Generally, a mature plant fiber related to me-
chanical support was considered as a dead cell . How-
ever, Bailey ( 1953 ) reported that libriform fiber of
trees can remain living protoplast after lignifying and
thickening of secondary wall . Recently, lots of libri-
formfiber or fiber tracheid of many species were ob-
served to have living protoplast, and these fibers were
mainly in the woods of Tamarix, Chenopodiaceae and
other dicotyledon (Fahn, 1982) . A bamboo culmcom-
pletes its growth and reaches the uppermost height in a
few months, and then can keep alive for 16 years
( Zhou, 1998 ) . Different from other woody plants,
bamboo, in its absence of secondary growth, depends
mainly on its primary vascular system during the whole
lifeof a culm . Thereby, the fibers originated fromthe
procambiumcontribute to the mechanical support until
the death of bamboo . Previous observation suggested
that bamboo fibers can keep intercellular linkage
through the pits and plasmodesmata in a mature culm
(Murphy et al. , 1997 ) . Nothing has been reported
about the persistence of the intercellular linkage with
age .
Becauseof the high fiber content of 38% (Gross-
er and Liese, 1974 ) , Phyllostachys edulis is widely
used for making furniture, construction, pulp andother
industries . The morphology, chemical components and
tissue ratio of fibers in P . edulis culmwere studied by
Parameswarn and Liese ( 1976 ) and Xiong et al.
(1980b) . He et al. ( 2000 ) suggested that the suc-
cessive development of bamboo fibers could be divided
into threestages: fiber initials development, the forma-
tion of primarywall (cell elongation) and the formation
of secondary wall ( cell wall thickening and lignifica-
tion) . Xiong et al. (1980a) investigated the early dif-
ferentiation and development of bamboo fibers . There
were some reports about the structural diversification of
bamboo fibers with age ( Liese and Weiner, 1997;
Murphy and Alvin, 1997) , and the continual thicken-
ingof fiber secondary wall with aging was also report-
ed . Mostly recent observation showed that the pro-
grammed cell death (PCD) of bamboo fiber occurred at
the earlier stageof secondarywall formation (He et al. ,
2000) . There were few investigations about the struc-
tural development during the whole formation of fiber
secondary wall .
ATP was considered as an energycarrier of all liv-
ing system including plants, and involved in energy
transform related to metabolism, transportation and in-
formation transfer of living substance . Energy was re-
leased only after hydrolyzing of ATP, so hydrolyzingof
ATP became abasic function for all living cells (Liu et
al. , 2000) . Therefore, the persistence of H+ -ATPase
in cells can become a criterion to validate the living
character (Wang et al. , 2000) .
APase is a non-specific hydrolytic enzyme essen-
tial to phosphate esters, carbohydrate and phosphate
metabolism, which could be important for phosphorus
scavenging and remobilization inplants (Darleen et al. ,
1989; Cashikar et al. , 1997; Yan et al. , 2001) . Mas-
sive evidences have been accumulated to demonstrate
that APase plays crucial roles in metabolism of cell
wall , intercellular nutrient transportation and phosphate
transfer reaction ( Tian et al. , 1999; Wang et al. ,
1999; Ibrahim et al. , 2002; Ferte et al. , 2003) .
Can bamboo fibers keep living protoplasts after
maturation? Why can bamboo fiber wall continually
thickening after PCD ? To answer these questions, the
ultrastructural modification and ultracytochemical local-
ization of ATPase, APase and Ca
2 +
-ATPase of fiber in
P . edulis culms during secondary wall formation were
studied with several microscopic and cytochemical
methods . The results indicated that the culm fiber of
P . edulis is a special long-lived cell , which differs
from that of woody dicotyledons .
1 Materials and methods
1 .1 Plant Materials
Young shoots and culms of 1 to 8-year-old P . edulis were
harvested in May 2003 at the Bamboo Gardenof Nanjing Forestry
University . Young shootswith the heights of 60 cm, 80 cm, 120
cm and 700 cmwere collected, and all samples with the size of
one-cm
3
from themiddle part of the culmwall were taken (Gan
and Ding, 2005 ; Fig . 1) .
1 . 2 Preparation for transmission electron microscopy
Samples were pre-fixed for 2 h in 0 .025 mol?l phosphate
buffer ( pH 7 .0) solution containing4% ( v?v) glutaraldehyde in
thecaseof young shoots and 1-year-oldculm, or 6 % (v?v) glut-
251 云 南 植 物 研 究 30 卷
araldehyde in the case of 2 to 8-year-old culms . After washing
with the same buffer, the samples were post-fixed in 1% OsO4
(also in thesamebuffer) . Followed by a further washing with the
buffer, the specimensweredehydrated in a graded ethanol series
and embedded in Epon812 resin . After cutting with a diamond
knife on aLKB-V ultramicrotome, ultrathin sectionswere stained
with saturated aqueous uranyl acetate for 5 min, followed by
stainingwith lead citratefor 5 min . Finally all sections were ex-
amined and photographed with an H-600 (Hitachi , Tokyo, Ja-
pan) transmission electronmicroscope (TEM) .
1 . 3 tPreparation for ultracytochemical localization of H+ -
ATPase ( ATPase)
Following pre-fixed in a mixture of 4 % paraformaldehyde
and 2 .5% glutaraldehyde, the samplesweretreated asthemeth-
od by Wang et al. (2000) . Thecontrolled specimenswere dealt
without substrate, or with substrate in thepresence of 10 mmol?L
sodiumfluoride (NaF ) . Made with LKB-V ultramicrotome, ul-
trathin sections of controlledmaterial werestained with uranyl ac-
etate, but normal samples no stained . All sectionswereobserved
and photographed usingan H-600 TEM .
1 . 4 lPreparation for ultracytochemical localization of APase
The samples from young shoots and 1 to 6-year-old culm,
were immediately fixed in amixtureof 4% paraformaldehydeand
2 .5% glutaraldehyde in 0 .05 mmol?L sodium cacodylate buffer
(pH 7 .2 ) for 2 h at 4℃ . After fixation, the samples were
washed in the same buffer at 4℃ for 1 .5 h, and were dissected
with a razor blade into slices of about 0 .5 mminthickness in the
buffer, and then rinsed with 0 .05 mmol?L Tris-maleate buffer
(pH 5.0) for 1 .5 h prior to enzyme reaction .
Theenzyme reaction procedure was performed following the
method by Wang et al. (1999) . The specimens were incubated
in a complete reaction medium (50 mmol?L Tris-maleate buffer,
pH 5 .0 , 3 .6 mmol?L Pb(NO3 )2 and 10 mmol?L sodiumβ-glyc-
erophosphate) at 37℃ for 2 h . The control specimens were re-
spectively incubated in the same medium but without substrate,
or with substrate in the presence of 10 mmol?L sodium fluoride
(NaF) . After incubation, all specimens were rinsed with 0 .05
mmol?L Tris-maleatebuffer ( pH 5 .2) for 1 .5 hat 4℃ , andthen
rinsed with 0 .05 mmol?L sodium cacodylate (pH 7 .2 ) buffer for
1 .5 h at the same temperature . Following this, these samples
werefixed at 4℃ with 1 % osmium tetroxide in the same buffer
overnight . Then all specimens were dehydrated stepwise in an
acetone series at 4℃ from30% to 100% . Specimenswere em-
bedded in Epon 812 . Ultrathin sectionsof control specimenswere
stained with uranyl acetate, while others were not .
1 . 5 ?Preparation for ultracytochemical localization of Ca2+ -
ATPase
The samples were cut and fixed as above mentioned, and
theenzymereaction procedurewas performed following themeth-
od by Jian et al. (2000) .
Fig . 1 Ultrastructural modification of fiber during secondary wall formation
a, the fiber at the early stageof secondary wall formation; b-c, ultrastructureof fiber when PCD occurred; b, showing the agglutinated chromatin, disinte-
grated organelle, and the lomasomebetweenwall and plasmamembrane(arrow) ; c, showing transfer vesicles and disintegration of mitochondrion; d-e, the
fibers in one-year-old culm; d, showing themultivesicular bodies ( arrow) and agglutinated chromatin; e, showing theplasmodesmata ( big arrow) and vesi-
cles(small arrow) ; f , showing the agglutinated chromatin in 6-year-old fibers .
G, Golgi bodies; M , mitochondrion; N, nucleus; R , rough endoplasmic reticulum; SW, secondary wall ; TV , transfer vesicle . Scar bars = 1μm .
3512 期 GAN and DING: A Special Long-Lived Cell : the CulmFiber Cell of Phyllostachys edulis (Gramineae)
2 Results
2 .1 ?Ultrastructural diversification of fiber during
secondary wall formation
As fiber completed elongation, fiber wall began
thickening billowingly indicated the beginningof second-
ary wall formation . At that time, intact double karyotheca
remains in fiber nucleus, wherein the nuclear chromatin
agglutinated revealingtheoccurrenceof PCD . However, a
lot of organelles such as mitochondria, rough endoplasmic
reticula and Golgi bodies remained (Fig . 1: a) .
With the thickening of secondary wall , fiber or-
ganelles gradually disintegrated ( Fig . 1 : b, c ) ,
whereas lomasome remained between fiber wall and
plasmamembrane ( Fig . 1 : b) . Afterwards, multive-
sicular bodies appeared in fiber cytoplasm, and double
karyothecaof fiber nucleus disappeared ( Fig . 1: d) .
Nevertheless, fiber nucleus with the agglutinated chro-
matin persisted in 6-year-old culms (Fig . 1 : f) . Along
with the formation of secondary wall , fiber wall under-
went continual thickening with aging: the thickening
level obviously increased in the first 4 years, and then
gradually decreased (Gan and Ding, 2005) . Polylami-
nate structures arranged in regular alteration of broad
and narrow lamellae appeared with thickening, and the
bordered pits were observed between fibers during the
thickening ( Fig . 1 : e) . Plasmodesmata and vesicles
still presented in thepits (Gan and Ding, 2005) .
2 . 2 FLocalization of H + -ATPase (ATPase) during
the formation of fiber secondary wall
At theearly stage, ATPase activity localizedon the
plasma membrane, tonoplast, nucleus, pits and plas-
modesmata andmitochondrion (Fig . 2: a, b) .With the
Fig . 2 Ultracytochemical localization of ATPase in the fibers during secondary wall formation
a-b, the fibers at the early stageof secondary wall formation; a, showing ATPase activity observed on theplasma membrane, tonoplast, theplasmodesmata
and chromatin; b, showingATPase localized inmitochondrion and nucleus; c, showingATPase localized on the annulate lamella (asterisk) and agglutinat-
ed chromatin in the fibers when PCD occurred; d-e, the distribution of ATPase in 1-year-old fibers; d, showing ATPase distributed in plasma membrane,
plasmalemma invagination and multivesicular bodies; e, showing ATPasedistributed in the plasmodesmata; f , showing the distribution of ATPase on the
plasma membrane and plasmalemma invagination in 4-year-old fibers; g-h, thedistribution of ATPase in 6-year-old fibers; g, showingATPasepersisted in
the plasmamembrane and agglutinated chromatin; h, showing the persistenceof ATPase in theplasmodesmata ( arrow) and transfer vesicles in pit channel ;
I , showing little ATPasedetected on the fibers in the controlled specimen . Ch, chromatin; M, mitochondrion; MB, multivesicular bodies; N , nucleus;
Nu, nucleolus; P, plasma membrane; PI , plasma membrane invagination; SW, secondary wall ; V , vacuole . Scar bars= 1μm .
451 云 南 植 物 研 究 30 卷
occurrence of fiber PCD, ATPase was observed on the
annulate lamella and the agglutinated chromatin in nu-
cleus ( Fig . 2 : c) . With the thickening of secondary
wall , the distribution of ATPase in fibers showed spa-
tial-temporal changes with aging . First, the multi-ve-
sicular bodies and transfer vesicles in pit channel and
plasmamembrane invagination with ATPase activity ap-
peared in the fibers of 1-year-old culms ( Fig . 2 : d,
e) . In the fibers of 4-year-old culms, ATPase was ex-
amined on theplasmamembrane and plasmamembrane
invagination ( Fig . 2 : f ) . Then the multi-vesicular
bodies would disappear gradually, while ATPase was
persistent on the plasma membrane, plasma membrane
invagination, and transfer vesicles in pit channels and
plasmodesmata in the fibers of 6-year-old culms, and
ATPase distributed on the agglutinated chromatin in-
creased with aging (Fig . 2 : g, h) .
Little ATPase activity was detected in the fiber of
the controlled specimens ( Fig . 2: i ) , which demon-
strated that the results above were credible .
2 . 3 ?Localization of APase in fiber during the for-
mation of secondary wall
Early, APasewas distributed on the nuclear chro-
matin and tonoplast and the cleavage cytoplasmof fiber
(Fig . 3 : a) , indicating the in situ self-lyses of cyto-
plasm . With the further development of fiber, the dis-
tribution of APaseshowed dynamic changes with aging .
In the bamboo fibers of 1-year-old, the plasma mem-
brane and agglutinated chromatin with APase activity
were observed . The enzyme was present in the transfer
vesicles, plasma membrane invagination, and plas-
modesmata between fibers and their adjacent cells
(Fig . 3 : b) . Then APase remained on the plasma
membrane, agglutinated chromatin, andplasmodesmata
between fibers and their adjacent cells up to six years .
The transfer vesicles and plasmamembraneinvagination
with the enzymatic activity would largely persist in the
fibers of former 4 years, and then decreased (Fig . 3 :
c - e) .
Little APase was detected in the fiber of the con-
trolled specimens (Fig . 3 : f) , which indicated that the
results abovewere credible .
2 . 4 MLocalization of Ca2 + -ATPase in fiber during
the formation of secondary wall
Fig . 3 Ultracytochemical localization of APase during fiber secondary wall formation
a, showing the agglutinated chromatin in nucleus and tonoplast and the disintegrated cytoplasm (star) with APase activity; b, the fibers of 1-year-old,
showing the distribution of APaseon the plasmodesmata(arrow) and transfer vesicles, and in thenucleus; c-d, the localization of APase in the 4-year-old
fibers; c, showing theplasmodesmata ( arrow) with APase activity between fiber and parenchyma cell; d, showing theplasmamembrane, plasmalemma in-
vagination and transfer vesiclewith APase activity; e, showing the APaseobserved on the chromatin and plasmamembrane and plasmodesmata (arrow) in
the fiber of 6-year-old; f , showing little APase detected on the fibers in the controlled specimens .
Ch, chromatin; L , cell lumen ( empty space produced during cell death) ; M , mitochondrion; N , nucleus; P, plasma membrane; PC , parenchyma cell ;
PI , plasmalemma invagination; SW, secondary wall ; TV , transfer vesicles; V , vacuole . Scar bars= 1μm .
5512 期 GAN and DING: A Special Long-Lived Cell : the CulmFiber Cell of Phyllostachys edulis (Gramineae)
Fig . 4 Ultracytochemical localization of Ca2 + -ATPase during secondary wall formation of fibers
a-b, the fibers at the early stage of secondary wall ; a, showing Ca2 + -ATPase distributed in nucleus; b, showing the localization of Ca2+ -ATPase in or-
ganelles, cytoplasm and plasma membrane; c-d, the fibers when PCD occurred; c, showing Ca2+ -ATPase examined on thedisintegrated tonoplast and cy-
toplasm (star) ; d, showing thedistribution of Ca2 + -ATPase in plasmamembrane and transfer vesiclesand plasmalemma invagination; e-f , the fibers of 2-
year-old; e, showing Ca2 + -ATPase investigated on the transfer vesicles and plasma membrane; f , showing the plasmodesmata( arrow) with Ca2+ -ATPase
activity; g-h, the fibers of 4-year-old; g, showing Ca2+ -ATPase localized on the plasmodesmata(arrow) ; h, showing the persistence of Ca2 + -ATPase in
cell lumen (star) and agglutinated chromatin; i , showing little Ca2 + -ATPase detected on the fibers in the controlled specimens .
Ch, chromatin; L , lumen; Nu, nucleolus; P , plasma membrane; PI , plasma membrane invagination; SW, secondary wall ; TV , transfer vesicles; V ,
vacuole . Scar bars = 1μm .
Early, Ca2 + -ATPase was present on the plasma
membrane, cytoplasm, nucleusof fiber and someendo-
membrane system like mitochondria and endoplasmic
reticula but theATPase less aggregatedon the tonoplast
(Fig . 4 : a, b) . With the occurrence of PCD, Ca2 + -
ATPasein thecleavage tonoplast and cytoplasmand ag-
glutinated chromatin increased gradually, and the
transfer vesicles with activity of Ca2 + -ATPase appeared
in fibers ( Fig . 4 : c, d) .
In 2-year-old culms, abundant transfer vesicles
with Ca2 + -ATPase activity remained in fibers, and
Ca2 + -ATPase was detected on the agglutinated chroma-
tin and plasmodesmata and disintegrated cytoplasm
( Fig . 4 : e, f) . Whereas Ca2 + -ATPase persisted in
plasma membrane, plasmodesmata, transfer vesicles
and agglutinated chromatin and cleavage protoplast
(Fig . 4 : g, h) .
Little APase was detected in the fiber of the con-
trolled specimens ( Fig . 4 : i ) , which confirmed the re-
ality of the above results .
3 Discussion
In this study, fiber nucleus with the agglutinated
chromatin in the culmof Phyllostachys edulis persisted
for up to eight years . The pits and plasmodesmata per-
sisted in the fiber of 8-year-old culms of P . edulis,
and a large number of vesicles also remained in pits .
In general , vesicles in cytoplasmoriginated from endo-
plasmic reticula and dictyosomes, in which includes
many cytoplasm and organelles that can disintegrate
651 云 南 植 物 研 究 30 卷
them, but the vesicles in pits played a role in trans-
porting thematerial fromthe disintegrated protoplast to
its surrounding cells as transfer vesicle . Therefore, the
persistence of plasmodesmata and vesicles could guar-
antee intercellular linkage between the fibers and their
adjacent cells . Generally, a mature fiber is considered
as a dead cell without protoplast, whereby bamboo fi-
bers can remain their protoplast for many years (Liese,
1998) . In addition, it was reported that the xylem fi-
bersof Tamarix aphylla kept alive for 20 years (Fahn,
1982) . Due to secondary growth of woody dicotyle-
dons, new secondary xylemwill form and become sap-
wood insteadof old secondary xylem . Fibers with living
protoplast are usually in sapwood, and have functions
in sustaining and storing ( Gu et al. , 1993 ) . While
those in heartwood, losing their protoplast, become
dead cells to have only function in sustaining ( Liese,
1998) . Differently, bamboo in its absenceof secondary
growth depends mainly on its primary vascular system
during the whole life of a culm . Once differentiating
from procambium, bamboo fibers as an important com-
ponent of primary vascular system will remain many
years . Thepersistence of nuclei and intercellular link-
age contributes to continual thickening of fiber second-
ary wall . The results showed that the fibers in this re-
search kept their living protoplast for a long time after
lignification and wall thickening .
According to our investigation, H+ -ATPase was
persistent on endomembrane system like plasma mem-
brane, plasmodesmata andvesicles for six years, which
helped to the nutrient absorbance of fiber from its sur-
rounding cells during secondary wall formation . Fur-
thermore, the persistence of H+ -ATPase in agglutinat-
ed chromatin showed that nucleus chromatin of fibers
remained alive for many years after PCD and thickening
of secondary wall . This indicated that the fibers can be
alive during the formation of secondary wall .
During secondary wall formation of the bamboo fi-
bers, Ca2 + -ATPase persisted in plasma membrane,
plasmodesmata, transfer vesicles and agglutinated chro-
matin and cleavageprotoplast . As aCa2 + pump, Ca2 + -
ATPase localized in plasma membrane played a role in
expellingCa
2 +
from cytoplasm, which assured continu-
ous thickening of secondary wall and the maintenance
of living function ( Jian et al. , 2000; Morten et al. ,
2004) . TheCa2 + -ATPase activity on plasmodesmata in
fibers contributed to the sustainingpermeability of sub-
stance, which correlated to the symplast transportation .
Additionally, Ca
2 +
-ATPase persisted in fibers, as a
functional protein, was related to the supply of energy
as well as ATPase during secondary wall formation .
Hereby, the persistence of Ca
2 +
-ATPase in plasma
membrane and plasmodesmata served to maintain the
physiological function of culmfiber .
APase localized on the plasma membrane, plas-
modesmata, plasmamembrane invagination and transfer
vesicles were detected for many years in the develop-
ment of fibers . The results implied that fiber canmain-
tain active transportation of cell wall materials and nu-
trient for fiber via vesicles and plasmodesmata . Addit-
ionally, the APase in the nucleus is considered as a
non-lysosome enzyme that may be associated with the
dephosphorylation processof thenuclear protein, influ-
encing the type and the quantity of the gene transcrip-
tion (Tian et al. , 1999) . The persistenceof APaseon
thenuclear chromatin indicated that fiber kept physio-
logical metabolism for a long time after PCD, which
can contribute to serve the demand of many proteins
and enzymes about secondary wall formation and PCD .
Thereby, the APase persisted in the substructureof fi-
ber werecorrelated to keeping fiber cell alive after PCD
occurred .
In conclusion, plasmodesmata, vesicles, and the
nuclei with agglutinated chromatin remained in bamboo
fibers for many years during secondary wall formation,
and some enzymes including H
+
-ATPase, Ca
2 +
-AT-
Pase, and APase were persistent in the substructure of
fibers . It obviously showed that the culmfiber of P . e-
dulis, different from that of many other woody plants,
is a special long-lived cell . The common roles of these
enzymes were closely correlated to the long-lived char-
acteristic and continual thickening of secondary wall
formation of fibers in P . edulis culms .
Acknowledgements : Theauthorsthank technician XihuaGan for
assistance in sample preparation and Dr . Jun Yang for helpful
suggestions and critical readingof this manuscript .
7512 期 GAN and DING: A Special Long-Lived Cell : the CulmFiber Cell of Phyllostachys edulis (Gramineae)
References:
Bail ?ey IW, 1953 . Evolution of the tracheary tissue of land plants [ J ] .
Amer J Bot, 40 : 4—8
Cash ikar AG, Kumaresan R , Rao NM , 1997 . Biochemical characteriza-
tion and subcellular localization of the red kidney bean purple acid
phosphatase [ J ] . Plant Physiol , 114: 907—915
Darl ?een AD, Stillman J I , George SE, 1989 . Acid phosphatase localiza-
tion in seedling tissues of the palms, Phoenix dactylifera and Wash-
ingtonia filifera , and its relevance to controls of germination [ J ] .
Canada J Bot, 67 : 1103—1110
Fahn :A, 1982 . Plant Anatomy (3rd ed) [M ] . Pergamon Press
Fert ?eN , Moustacas AM , Nari J et al. , 2003 . Characterization and kinet-
ic properties of a soya-bean cell-wall phosphatase [ J ] . European J
Biochemistry, 211: 297—304
Gan !XH , Ding YL , 2005 . Development anatomy of the fiber in Phyl-
lostachys edulis culm [ J ] . Bamboo Sci Cult, 19 (1 ) : 16—22
Gros ?ser D, Liese W, 1974 . Verteilung der Leitbündel und Zellarten in
Spro?achsen verschiedener Bambusarten [ J ] . Holz Roh-Werkstoff,
32 : 473—482
Gu A 8G ( 谷安根 ) , Lu JM ( 陆静梅 ) , Wang LJ (王立军 ) , 1993 . The
evolvement morphology of vascular plants ( 维管 植 物演 化 形态
学 ) [M ] . Changchun: J ilin Scientific and Technologic Press, 135
He X 8Q ( 贺新强 ) , Wang YQ ( 王幼群 ) , Hu YX ( 胡玉熹 ) et al. ,
2000 . Ultrastructural study of secondary wall formation in the stem
fibre of Phyllostachys pubescens [ J ] . Acta Bot Sin ( 植物学报 ) ,
42 : 1009—1013
Ibra ?him H , Pertl H , Pittertschatscher K et al. , 2002 . Releaseof an acid
phosphatase activity during lily pollen tubegrowth involvescomponents
of the secretory pathway [ J ] . Protoplasma, 219: 176—183
J ian ?LC , Ji HL , Li PH et al. , 2000 . An eletron microscopic-cytochemi-
cal localization of plasma membrane Ca2 + -ATPase activity in poplar
apical bud cells during the induction of dormancy by short-day photo-
periods [ J ] . Cell Res, 10 : 103—114
Lies ?e W, Weiner G, 1997 . Modificationsof Bamboo CulmStructures Due
to Ageing and Wounding [ A ] . In: Chapman GP ed, The Bamboos
[M ] . London: The Linnean Society Symposium Academic Press,
313—322
Lies ?e W, 1998 . The Anatomy of Bamboo Culms [M ] . International Net-
work for Bamboo and Rattan Technical Report, 130—135
Liu ?L ( 刘林 ) , Fan SG ( 范树国 ) , Zhang ZJ ( 张再君 ) et al. , 2000 .
Ultrastructural localization of adenosine triphosphatase activity inwater
melon stigmapapilla cells during secretion [ J ] . Chin Bull Bot ( 植物
学通报 ) , 17 (2 ) : 179—184
Mort ?en S, Shawn MR , Lone B et al. , 2004 . A plant plasma membrane
Ca2 + pump is required for normal pollen tubegrowth and fertilization
[ J ] . Proceedings of the National Academy Sciences USA, 101 (25 ) :
9502—9507
Murp ?hy RJ , Alvin KL , 1997 . Fibre Maturation in Bamboos [ A ] . In:
Chapman GP ed, The Bamboos [M ] . London: The Linnean Society
SymposiumAcademic Press, 293—303
Murp ?hy RJ , Sulaiman O, Alvin KL , 1997 . UltrastructureAspects of Cell
Wall Organization in Bamboos [ A ] . In: Chapman GP ed, The Bam-
boos [ M ] . London: The Linnean Society Symposium Academic
Press, 305—312
Para ?meswaran N , Liese W, 1976 . On the fine structure of bamboo fibers
[ J ] . Wood Sci Tech, 10 : 231—246
Tian ?GW ( 田国伟 ) , Shen JH (申家恒 ) , 1994 . The ultracytochemical
localization of acid phosphatase activity in wheat during fertilization
[ J ] . Acta Bot Sin (植物学报 ) , 36 : 251—256
Tian ?GW ( 田国伟 ) , Shen JH ( 申家恒 ) , You RL ( 尤瑞麟 ) , 1999 .
Ultracytochemical of localization of acid phosphatase in nucellar cells
wheat during degeneration [ J ] . Acta Bot Sin, 41 : 791—794
Wang ?YQ ( 王雅清 ) , Chai J J (柴晶晶 ) , Cui KM ( 崔克明 ) , 1999 .
Ultracytochemical localization of acid phosphatase during differentia-
tion and dedifferentiation of the secondary xylem in Eucommia ul-
moides trunk [ J ] . Acta Bot Sin (植物学报 ) , 41 : 1155—1159
Wang ?YQ ( 王雅清 ) , Kalima-N′Koma MWANGE , Cui KM (崔克明 ) ,
2000 . Ultracytochemical localization of ATPase during the second-
ary xylem differentiation and dedifferentiation in Eucommia ulmoides
Trunk [ J ] . Acta Bot Sin (植物学报 ) , 42 ( 5) : 455—460
Xiong WY ( 熊文愈 ) , Ding ZF ( 丁祖福 ) , Li Y F ( 李又芬 ) , 1980a .
Intercalary meristem and internodal elongation ofamboo plants [ J ] .
Sci Silvae Sin (林业科学 ) , 16 ( 2) : 81—89
Xion ?gWY ( 熊文愈 ) , Qiao SY ( 乔士义 ) , Li YF ( 李又芬 ) , 1980b .
Anatomical structure of culms in Phyllostachys pubescens [ J ] . Acta
Bot Sin (植物学报 ) , 22 ( 4) : 343—348
Yan ?XL , Liao H , Trull MC et al. , 2001 . Induction of a major leaf acid
phosphatase does not confer adaptation to low phosphorus availability
in common bean [ J ] . Plant Physiol , 125: 1901—1911
Zhou ?FC ( 周芳纯 ) , 1998 . Cultivation and utilization of bamboo ( 竹林
培育与利用 ) [ R] . Nanjing: The editorial committee of bamboo
research, Nanjing Forestry University, 59—68
851 云 南 植 物 研 究 30 卷