全 文 ：武汉植物学研究 2001, 19 (2) : 135～ 142
J ourna l of W uhan B otan ica l Resea rch
3 种藓类植物水分含量与光合作用、
呼吸作用和水势的关系
刘应迪1Ξ 朱杰英1　陈　军1　曹　同2
(1. 湖南省吉首大学生命科学和化学学院, 湖南吉首　416000;
2. 中国科学院沈阳应用生态研究所, 沈阳　110015)
摘　要: 对湿地匍灯藓〔P lag iom nium acu tum (L indb. ) T. Kop. 〕、大羽藓〔T hu id ium cym bi2
f olium (Dozy &M o lk. ) Dozy &M o lk. 〕和垂藓〔Ch ry soclad iun retrorsum (M itt. ) F leisch. 〕
的水分含量与光合作用、呼吸作用和水势的关系进行了初步研究 (1999 年 5 月 20 日到 6 月
10 日)。在这 3 种藓类植物中, 其水分含量与光合作用速率 (Pn)的关系可以分为 2 种类型: 一
种类型如大羽藓和垂藓, 在藓体水分含量 20%～ 70% 时, Pn 随着水分含量增加而增加, 但是
在 80%～ 95% 时, Pn 随水分含量增加而下降, 光合最适水分含量约 70%～ 80% ; 另一种出现
在湿地匍灯藓, 水分含量 20%～ 80% 时, Pn 随着水分含量增加而增加, 在 80%～ 95% 时, Pn
维持一个较高的水平, 光合最适水分含量为 80%～ 90%。在一个大的水分含量范围内 (60%
～ 95% ) , 暗呼吸 (R d) 保持相对稳定, 但是在水分含量较低时 (20%～ 70% ) , R d 随着水分含
量的下降而下降。在藓体水分含量与水势之间的关系方面, 3 种藓类植物相似, 水分含量与水
势对数之间的回归曲线为 S 形曲线。
关键词: 藓类; 光合作用; 呼吸作用; 水分含量; 水势
　　中图分类号: Q 945　文献标识码: A 　文章编号: 10002470X (2001) 0220135208
Rela tion sh ips of W ater Con ten t to Photosyn thes is, Resp ira tion
and W ater Poten tia l in Three Spec ies of M osses
L IU Y ing2D i1Ξ , ZHU J ie2Y ing1, CH EN Jun1, CAO Tong2
(1. Colleg e of L if e S ciences and Chem istry , J ishou U n iversity , J ishou, H unan　416000, Ch ina;
2. Institu te of A pp lied E cology , T he Ch inese A cad emy of S ciences, Shenyang　110015, Ch ina)
Abstract: T he rela t ion sh ip s of w ater con ten t to pho to syn thesis, resp ira t ion
and w ater po ten t ia l in th ree species of mo sses, P lag iom n ium acu tum (L indb. )
T. Kop. , T hu id ium cym bif olium (Dozy &M o lk. ) Dozy &M o lk. , and Ch ry so2Ξ 收稿日期: 2000210231, 修回日期: 2001203204。作者简介: 刘应迪 (1958- ) , 男, 博士, 副教授, 从事植物学、植物生理学和植物生态学的教学和研究工作。本文是博士论文的一部分。通讯联系人。A utho r fo r co rrespondence.
clad iun retrorsum (M it t. ) F leisch. w ere studied du ring M ay 20 to June 10,
1999. Among these th ree mo sses, there are tw o types of w ater respon se mod2
els fo r net pho to syn thet ic ra te (Pn). O ne appeared in T hu id ium cym bif olium
and Ch ry soclad ium retrorsum , in w h ich Pn increased w ith the w ater con ten t
from 20% to 70% , bu t decreased from 80% to 95%. T he op t im um w ater con2
ten t w as 70%～ 80%. A no ther w as p resen ted in P lg iom n ium acu tum , in w h ich
net Pn increased w ith w ater con ten t increasing from 20% to 80% , and a h igh
level of Pn m ain ta ined from 80% to 95% of w ater con ten t. T he op t im um w ater
con ten t w as 80%～ 90%. In a w ide range of w ater con ten t from 60 (70) % to
95% , the R d rem ained rela t ively stab le. Bu t in the range of low er w ater con2
ten ts from 20% to 70% , the R d decreased w ith decreasing w ater con ten ts.
Changes in betw een w ater con ten ts and w ater po ten t ia ls w ith in these th ree
mo sses are sim ilar to each o ther. T he w ho le cu rve of w ater con ten ts to loga2
rithm of w ater po ten t ia ls appears to be S2fo rm ed.
Key words: M o sses; Pho to syn thesis; R esp ira t ion; W ater con ten t; W ater po2
ten t ia l
1　 In troduction
B ryohpytes face the sam e basic p rob lem s of life on the land as flow ering p lan ts.
W ater is essen t ia l fo r m etabo lism , bu t is lim ited and erra t ic supp ly in the aboveground
environm en t w here the leaves m u st be expo sed in o rder to pho to syn thesize [1 ].
B ryophytes and vascu lar p lan ts exemp lify tw o alternat ive pat tern s of adap ta t ion to these
condit ion s. V ascu lar p lan ts have evo lved roo ts and an eff icien t conduct ing system ,
b ringing w ater from the so il w here it is rela t ively p len t ifu l and con stan t supp ly.
B ryophytes have evo lved to u t ilize w ater w here and w hen it is availab le above ground;
som e are confined to mo ist hab ita ts, bu t m any can to lera te drying ou t, and som e are ex2
t rem ely desiccat ion2to lera te and h igh ly adap ted to a po ik ilohydric ex istence. T he studies
of w ater rela t ion s of som e b ryophytes w ere w ell docum en ted in resen t pub lica t ion s in the
w o rld [2 8 ] , bu t w ere neglected in Ch ina.
It is necessary to cu lt iva te the w in ter ho st mo sses fo r the gallnu t p roduct ion in Ch i2
na
[9 17 ]
, w h ich requ ires u s to understand the physio logy of mo sses. So ou r a t ten t ion has
also been paid to the physio logica l studies including ligh t and temperatu re respon se in
pho to syn thesis, d iu rnal cou rse of pho to syn thesis and dark resp ira t ion of mo sses[18 21 ].
T h is paper repo rts som e experim en ta l resu lts on the rela t ion sh ip s of w ater con ten t of
w in ter ho st mo sses w ith pho to syn thesis, dark resp ira t ion and w ater po ten t ia l.
2　M a ter ia ls and M ethods
2. 1　M a ter ia ls
T he th ree mo sses u sed in the experim en ts w ere co llected from D ehang scen ic spo t in
J ishou, w estern H unan, Ch ina, 28°15′N , 109°40′E, at an elevat ion of 300～ 350 m. T he
mo sses include th ree abundan t and common species. O ne is P lag iom n ium acu tum
(L indb. ) T. Kop. it u sually grow s on a th in layer of so il over roch s near st ream s o r on
631 武 汉 植 物 学 研 究　　　　　　　　　　　　　　　　第 19 卷　　
the h illsides of w ooded moun ta in s. Its hab ita t is shaded, w et, o r som etim es un shaded.
A no ther common species is T hu id ium cym bif olium (Dozy & M o lk. ) Dozy & M o lk. .
T h is mo ss often grow s on expo sed rock s beside roads o r near st ream s in w ooded moun2
ta in s. T he th ird, Ch ry soclad iun retrorsum (M it t. ) F leisch. It is a d ist inct ive mo ss that
grow s on ly in the hum id hab ita ts of w ooded moun ta in s, common ly hanging on cliffs o r
rock s faces. T he voucher specim en s are hou sed in the H erbarium of J ishou U n iversity.
2. 2　M ea surem en t of the wa ter con ten t and net photosyn thesis
T he C I2301PS (C ID In s. , U SA ) w as u sed to m easu re the mo ss net pho to syn thesis
(Pn). T he open system w as u sed in the experim en t. T he Pn in the ou tpu t of th is sys2
tem w as based on the amoun t of CO 2 per leaf area un it (Λmo lõm - 2õs- 1) , w h ich m u st be
converted. In th is paper, the Pn is exp ressed based on the amoun t of CO 2 per dry w eigh t
un it of m ateria ls (Λmo lõkg - 1 DWõs- 1).
W hen m easu ring the w ater rela t ion of Pn, the Pho to syn thet ic act ive radia t ion
(PA R ) on the assim ila t ion cham ber w as con tro lled w ith in 800～ 900 Λmo lõm - 2õs- 1, and
the temperatu re of the samp les, w ith in 24～ 26℃. T he dark resp ira t ion (R d) w as m ea2
su red in darkness. T he m ateria ls w ere selected and cleaned befo re m easu ring. A t first,
the samp les w ere w et ted and w eighed. T hey w ere then p laced in the cham ber and m ea2
su red rap id ly ( in 3～ 5 m in. ). T hen the samp les w ere removed from the cham ber and
p laced in sun ligh t o r under electric lamp fo r evapo ra t ion of the w ater in the samp les.
A bou t 10 m inu tes la ter, the samp les w ere again p laced in the cham ber and m easu red,
and so on. In terval t im e betw een the tw o m easu rem en ts depended on the evapo ra t ion
ra te of w ater in the samp les. U sually 5～ 7 m easu rem en ts w ere needed fo r a single sam 2
p le from the satu ra ted to abou t 15%～ 25% w ater con ten t. T he resu lts repo rted in th is
paper include m easu ring values of f ive para llel samp les fo r one species of mo sses, and a
single m easu ring value is an average of five readings in the experim en t. A fter the m ea2
su rem en t, the m ateria ls w ere dried at 80℃ fo r 8 hou rs and then w eighed.
2. 3　M ea surem en t of rela tion of wa ter con ten t to wa ter poten tia l
T he m ethod fo r m easu ring rela t ion sh ip betw een w ater con ten t and w ater po ten t ia l
w as app rox im ately the sam e as Bu sby & W h itf ield [4 ] and D ilk s & P rocto r [3 ]. Sa tu ra ted
salt so lu t ion s in clo sed con ta iners (a t certa in temperatu re) p rovide p recise rela t ive hu2
m idity that can be tran sfo rm ed in to w ater po ten t ia l values. A range of rela t ive hum idity
percen t of values (15, 32. 3, 44, 54, 75. 5, 81, 86, 90, 93, 95, 98) w as cho sen and salt
so lu t ion s that p rovide these hum idity w ere selected acco rd ing to W ylie [22 ] , W east [23 ] and
D ilk s & P rocto r [3 ].
M o ss samp les w ere su spended over the sa tu ra ted salt so lu t ion s in clo sed flask s.
T he temperatu re in the flask s w ere m ain ta ined at 20℃± 0. 5℃. T he samp les w ere
w eighed at an in terval of tw o days un t il a con stan t w eigh t w as reached. T he w ater con2
ten ts w ere determ ined from the equ ilib rium w eigh t and dry w eigh t (80℃ fo r 8 h). M ea2
731　第 2 期　　　　　　刘应迪等: 3 种藓类植物水分含量与光合作用、呼吸作用和水势的关系 (英)
su rem en ts w ere m ade on bo th w et and dry m ateria ls. W ater po ten t ia l near zero
(0. 1 m Pa) w as ob ta ined by su spending the m ateria ls over d ist illed w ater in the flask s.
3　Results
Op tim al tempera tu res and PA R levels have been estab lished fo r these th ree
mo sses in ou r p reviou s studies[21, 24 ]. A cco rd ing to these resu lts, m easu rem en ts of
influence of w ater con ten t on net pho to syn thet ic ra te w ere m ade at PA R level of 800～
900 Λmo lõm - 2õs- 1 and a samp le temperatu re of 24～ 26℃. T he resu lts ob ta ined under
the sam e condit ion bu t w ith the cham ber covered w ith dark fab ric rep resen t dark resp i2
ra t ion ra te. T he experim en ts w ere carried ou t in the labo ra to ry from M ay 20 to June 10,
1999. T he resu lts are show n in F igs. 1 and 2.
3. 1　W a ter con ten t and net photosyn thesis (Pn )
Among these th ree mo sses, there are tw o types of w ater respon se models fo r net
pho to syn thesis ra te (Pn ). O ne appeared in T hu id ium cym bif olium and Ch ry soclad ium
retrorsum , in w h ich Pn increased w ith the w ater con ten t from 20% to 70 (80) % , bu t
F ig11 　Relationsh ip s of net pho to syn thesis ( Pn,
so lid circles) and dark resp iration (Rd, open circles)
to w ater con ten t of the mo sses P lag iom n ium acu2
tum (L indb. ) T. Kop. (A ) , T hu id ium cym bif olium
(Dozy &M ilk. )Dozy &M ilk. (B) , and Ch ry soclad i2
um retrorsum (M itt. ) F leish. (C)
　
decreased from 80% to 95%. T he op t im um
w ater con ten t w as 70%～ 80% (F ig. 1: B , C).
A no ther w as p resen ted in P lg iom n ium acu tum ,
in w h ich Pn increased w ith w ater con ten t in2
creasing from 20% to 80% , and a h igh level of
Pn m ain ta ined from 80% to 95% of w ater con2
ten t. T he op t im um w ater con ten t w as 80%～
90% (F ig. 1: A ).
T he rela t ion sh ip betw een w ater con ten t
and Pn has been w ell docum en ted in variou s
mo sses
[3 6, 25, 26 ]
. A cco rd ing to St ao lfelt [25 ] and
D ilk s & P rocto r [3 ] , w ater respon se cu rves fo r
Pn of m any mo sses grow ing in dry hab ita t ex2
h ib ited an op t im um , w ith the ra te declin ing a2
gain at h igh w ater con ten t. O n ly in species
grow ing in w et hab ita t such as H ookeria lu2
cens, d id the ra te of Pn increase con t inually a t
the h igh level of w ater con ten t. In ou r experi2
m en t, the w ater respon se model fo r Pn show n
by T hu id ium cym bif olium and Ch ry soclad ium
retrorsum is app rox im ately the sam e as that
w h ich appeared in mo sses grow ing in dry hab i2
ta ts described by Stao lfelt [25 ] and D ilk s & P roc2
831 武 汉 植 物 学 研 究　　　　　　　　　　　　　　　　第 19 卷　　
to r [3 ]. Bu t in P lag iom n ium acu tum , the model is sim ilar to that of mo sses grow ing in
w et hab ita ts.
3. 2　W a ter con ten t and dark resp ira tion (Rd)
A lthough there w ere differen t models of w ater respon se fo r Pn, the w ater respon se
model fo r dark resp ira t ion ra te (R d) in the th ree mo sses w as sim ilar. In a w ide range of
w ater con ten t from 70% to 95% the R d m ain ta ined rela t ively stab le. Bu t in the range of
low er w ater con ten t from 20% to 60 (70) % , the R d decreased w ith the w ater con ten t
decreasing.
Among these th ree mo sses, the w ater respon se of R d in the range of low er w ater
con ten t is rough ly synch ronou s w ith that of Pn fo r T hu id ium cym bif olium (F ig. 1: B )
and Ch ry soclad ium retrorsum (F ig. 1: C ) bu t beh ind fo r that of P lag iom n ium acu tum
(F ig. 1: A ). T h is show s that pho to syn thesis is mo re sen sit ive to desiccat ion than resp i2
ra t ion fo r mo sses grow ing in w et hab ita ts, such as P lag iom n ium acu tum . In mo sses
grow ing dry hab ita ts, such as T hu id ium cym bif olium and Ch ry soclad ium retrorsum ,
how ever, the respon se to desiccat ion fo r Pn and R d is rough ly the sam e.
3. 3　W a ter con ten t and wa ter poten tia l
R elat ion sh ip s betw een w ater con ten t and w ater po ten t ia l fo r these th ree mo sses are
p resen ted in F ig. 2. T he differences among species are m ino r, and there are no sign if i2
can t d ifferences betw een deso rp t ion cu rves (ob ta ined from w et samp les) (F ig. 2: A ) and
ab so rp t ion cu rves (from dry samp les) (F ig. 2: B ) excep t fo r the data m easu red from
A. D eso rp tion (w etting) curves; B. A bso rp tion (drying) curves.
●P lag iom n ium acu tum ; ○T hu id ium cym bif olium ;
□Ch ry soclad ium retrorsum (M itt. ) F leish. (C)
F ig12　Relationsh ip s of w ater con ten t (% ) to w ater po ten tial
　
near 0 to - 217 m Pa of w a2
ter po ten t ia l. T he equ ilib ri2
um w ater con ten ts over d is2
t illed w ater w ere diff icu lt to
determ ine [4 ]. Becau se the
w ater po ten t ia l grad ien t be2
tw een mo sses and the satu2
ra ted atmo sphere w as
m inu te, the ra te of w ater
vapo r movem en t w as m inu te
as w ell. A lthough the sam 2
p les w ere m ain ta ined fo r
tw o w eek s on dist illed w ater and a satu ra ted so lu t ion of CuSO 4 ( - 217 m Pa of w ater
po ten t ia l) in ou r experim en t, it d id no t elim inate the hysteresis effect on w et t ing and
drying cu rves from near 0 to - 217 m Pa of w ater po ten t ia l as show n in F ig. 2. F rom
- 1413 to - 257 m Pa of w ater po ten t ia l, irregard less of w et t ing o r drying cu rves, the
equ ilib rium w ater con ten t of these th ree mo sses is qu ite sim ilar, and a linear rela t ion be2
tw een the w ater con ten ts and the logarithm of w ater po ten t ia l w as p resen ted. T he en t ire
931　第 2 期　　　　　　刘应迪等: 3 种藓类植物水分含量与光合作用、呼吸作用和水势的关系 (英)
cu rve of w ater con ten t to logarithm of w ater po ten t ia l appears to be S2fo rm ed (F ig. 2).
T he differences fo r rela t ion of w ater con ten t to w ater po ten t ia l among species of
mo sses have been p roven to be m inu te by m any w o rkers[3, 4 ]. D ilk s & P rocto r pu t fo r2
w ard a schem atic rela t ion of w ater con ten t to w ater po ten t ia l fo r a po ik ilohydric
mo ss
[1, 3 ]
. W e averaged the values m easu red from ou r th ree mo sses, bo th w et and dry
samp les, and also p lo t ted ou t a schem atic rela t ion of w ater con ten t to w ater po ten t ia l
(F ig. 3). Compared w ith D ilk s & P rocto r, ou r schem atic rela t ion demon stra ted a sim ilar
genera l w ater con ten t respon se model to w ater po ten t ia l, bu t included a lit t le descrip t ion
of ex ternal cap illa ry w ater associa ted w ith shoo ts of mo sses.
F ig13　Relationsh ip s of w ater con ten t to w ater po ten2
t ial fo r the mo sses P lag iom n ium acu tum , T hu id ium
cym bif olium and Ch ry soclad ium retrorsum. T he curve
w as fit ted acco rding to averages of data from bo th w et
and dry samp les of the th ree mo sses. F rom A to B, mo st
w ater is clo sely associated w ith su rfaces and is relatively
immobile. F rom B to C, increase of w ater con ten t is
m ain ly w ith in the cells. C is the po in t of incip ien t p las2
mo lysis; from C to D the w ater con ten t increases w ith
the bu ilt2up of w all p ressu re and the first appearnce of
ex ternal cap illary w ater
4　D iscussion
Stao lfelt (1937) po in ted ou t that the w ater respon se cu rve of pho to syn thesis fo r H y 2
locom ium sp lend ens exh ib ited an op t im um , w ith the ra te declin ing again at h igh w ater
con ten t, in con trast to the behavio r of resp ira t ion [25 ]. T h is is p robab ly a con sequence of
the m uch slow er diffu sion of gases th rough w ater than th rough air; and excess w ater
w ou ld be expected to rest rict gas exchange at the low concen tra t ion gradien ts availab le
fo r CO 2 up take bu t shou ld have neglig ib le effect on resp ira t ion. Bu t D ilk s & P rocto r [3 ]
p resen ted data that the Pn rate declin ing at h igh w ater con ten ts on ly appeared in mo sses
grow ing in dry hab ita ts. T h is m ean s that the Pn rate declin ing at h igh w ater con ten ts
p robab ly show ed the physio logica l characterist ics of the mo sses grow ing in dry hab ita ts,
and w as no t, o r w as no t comp letely, a con sequence of the slow er diffu sion of gases
th rough w ater than th rough air. O u r resu lts a lso suppo rt th is idea. T hu id ium cym bif oli2
um and Ch ry soclad ium retrorsum are bo th mo sses that grow on rock, and they are no t
on ly the desiccat ion2resistan t mo sses, bu t are a lso sen sit ive to excess w ater in their
hab ita ts. P lag iom n ium acu tum , how ever, grow ing on so il and in a w et hab ita t, is a
desiccat ion2sen sit ive mo ss, and can to lera te excess w ater in its hab ita t.
R esp ira t ion, on o ther hand, show s lit t le change over a w ide range of w ater con ten t
fo r these th ree mo sses in ou r experim en t. A cco rd ing to the experim en t described by
D ilk s & P rocto r, po rt ion s of the cu rves over w h ich Pn and R d are steep ly rela ted to w a2
ter con ten t are no t con sp icuou sly separa ted fo r mo sses grow ing in dry hab ita t (F ig. 2) ,
041 武 汉 植 物 学 研 究　　　　　　　　　　　　　　　　第 19 卷　　
bu t are separa ted fo r mo sses grow ing in w et hab ita ts (F ig. 1) [3 ]. T h is is a con sequence of
a slow er increase w ith w ater con ten t increasing mo re fo r Pn than fo r R d fo r the mo sses
grow ing in w et hab ita ts. In ou r experim en t, Pn and R d of T hu id ium cym bif olium and
Ch ry soclad ium retrorsum increased rap id ly w ith increasing w ater con ten t, bo th reached
their m ax im um at 70% of w ater con ten t. T hese tw o mo sses behaved the sam e as mo sses
grow ing in dry hab ita ts described by D ilk s & P rocto r [3 ]. Fo r ano ther species, P lag iom 2
n ium acu tum , the Pn increased w ith w ater con ten t slow er than did R d, behaving the
sam e as mo sses grow ing in w et hab ita ts as described by D ilk s & P rocto r [3 ].
Fo r the rela t ion sh ip betw een w ater con ten t and w ater po ten t ia l, Bu sby &W h itfeld
po in ted ou t that d ifferences among species are m ino r [4 ]. T h is w as confirm ed in ou r ex2
perim en t. Bu t som e p rob lem s ex isted w ith th is m easu rem en t especia lly a t a w ater po ten2
t ia l near zero. In the experim en t described by Bu sby &W h itfeld [4 ] , w ater po ten t ia l near
zero w as ob ta ined by su spending dried m ateria l (- 2913 to - 281 m Pa) over d ist illed
w ater. Equ ilib rium w ater con ten ts, how ever, cou ld no t be accu ra tely determ ined be2
cau se the samp les w ere overgrow n by fungi befo re their w eigh ts stab ilized. T he data ob2
ta ined in the m easu rem en t described by Bu sby &W h itfeld [4 ] w ere p resum ab ly low er than
the real values. W e u sed w et and dry samp les to m easu re the equ ilib rium w ater con ten t
fo r w ater po ten t ia l near zero by su spending the samp les over d ist illed w ater (near zero)
and a satu ra ted so lu t ion of CuSO 4 (- 217 m Pa) fo r tw o w eek s. T he average of values
ob ta ined from w et and dry samp les m ay be clo se to the real value.
T he differences betw een mo sses and flow ering p lan ts is pu t in perspect ive by the
fact tha t Pn in the la t ter decreases as soon as w ater po ten t ia l drop s below - 011 to
- 013 m Pa. Indeed, in mo st f low ering p lan ts Pn m ay reach zero o r even becom e nega2
t ive ( i. e. , resp ira t ion is exceeding pho to syn thesis) w hen w ater po ten t ia l drop s below
- 112 to - 115 m Pa. T hu s, to su rvive, f low ering p lan ts and fern s m u st con tro l their
w ater con ten ts so that they rem ain w ith in the lim its set by their w ater po ten t ia l
values[4 ]. Bu t in mo sses, especia lly in mo sses grow ing in dry hab ita ts, Pn decreases
mo re slow ly w ith a decreasing w ater po ten t ia l (See F ig. 1 and F ig. 2). W hen w ater po2
ten t ia l dropped to - 110～ - 115 m Pa, the Pn decreased on ly a lit t le. W hen w ater po2
ten t ia l con t inued to drop below - 217 to - 918 m Pa, Pn decreased qu ick ly. W hen w ater
po ten t ia l dropped below - 918 to - 1413 m Pa, Pn app roached zero o r becam e negat ive.
T h is resu lt show s that the mo sses canno t con tro l their w ater con ten ts o r w ater po ten2
t ia ls w ith in a rela t ively stab le range, bu t they can viab ly respon se to the fluctuat ion of
w ater po ten t ia l in their hab ita t.
Acknowledgemen t: W e w ish to thank the Educational Comm ittee of H unan P rovince fo r the suppo rt a t
th is w o rk. W e also thank D r. Buck W. R. fo r reading th is m anuscrip t andM r. H uang J R &D uan SW
fo r tak ing part in the experim ents.
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