全 文 ：武汉植物学研究 2003, 21( 6) : 537～543
Journal of Wuhan Botanical Research
光敏色素分子特性及其信号转导机制
张鸿明1 , 赵 实1, 高荣孚2 , 刘玉军2
( 1.华东师范大学生命科学学院, 上海 200062; 2.北京林业大学生物科学与技术学院, 北京 100083)
摘 要: 结合生物物理、分子遗传学和细胞生物学的方法已证实,光敏色素信号转导是一个空间分布的、非线形信号
传递链。尤其是最近又发现了不同种类的光敏色素分子及其它们在P r、P fr 光转换中产生的中间体,不仅说明了光敏
色素信号转导链是一个多维的信号网络, 而且这也暗示着光转换中产生的中间体也直接参与了早期的信号转导。在
此,综述了光敏色素分子光转换及其早期信号转导的若干新进展,讨论光敏色素原初光反应及其信号转导的机制。
关键词: 光敏色素; 光转换; 信号转导; 信号中间组分
中图分类号: Q945. 41 文献标识码: A 文章编号: 1000-470X( 2003) 06-0537-07
Molecular Properties of Phytochromes and
Their Signalling Mechanism
ZHANG Hong-M ing
1
, ZHAO Shi
1
, GAO Rong-Fu
2
, LIU Yu-Jun
2
( 1. L if e Sci ence Col leg e, East China N ormal Univ ersity , Sh ang hai 200062, China;
2. Coll ege of B iolog ical S ci ence & Biotechnology , B eij ing F ore st ry Unive rsity, Beijin g 100083, Ch ina)
Abstract: Through a combinat ion o f biophysical, molecular genet ic and cell biolo gical approa-
ches, it is compel ling ev idence that phytochr ome signal t ransduction is spat ially separated nonlin-
ear chain of ev ents. In par ticular , the discovery that different phytochr ome species and their in-
termediates during Pr, P fr pho to transformation show mult idimensional signal netw o rk, w hich
lead to the suggestion that these intermediates are direct ly involved in these early signal tr ansduc-
tion. Here, the primary photorseact ion of phy to chrome and the corr esponding signalling mecha-
nism will be discussed by review ing some o f recent prog ress in phytochrome photot ransformation
and its signal tr ansduct ion in ear ly stage.
Key words: Phy to chrome; Photot ransformat ion; Signall ing t ransduct ion; Signalling intermedi-
ates
As sessile organisms, plants are unable to
move act ively tow ards favo rable or aw ay f rom un-
favo rable env ironmental conditions like animals.
Therefo re, plant have evolved div erse photorecep-
to r systems for detect ing light intensity , quality,
and duration to adjust their life in fluct ing env iron-
mental condit ions. Phytochrome, as the primary
photoreceptor , ex ert their photoregulato ry ef fects
at all stages of the plant life cy cle, including
chloroplast movement , const ruct ion of the photo-
synthet ic apparatus, seed germination, de-et iola-
t ion and seedling establishment , shade avoidance
and the induction and t iming of flowering, thus de-
termining the st rategy of their opt imum light de-
Received date: 2003-08-26, Accepted date: 2003-09-22.
Fou ndat ion item : Su pported by th e Excellent Yon g T each ers Program of MOE.
Biography: ZHANG Hong -Ming ( 1978- ) , male, m as ter, wh ose research field is mainly ab ou t plant biology.
Correspondin g author.
velopment
[ 1]
, that is, pho tomorphorg enesis. The
photosignal percept ion and transduct ion is based
on the photoreversible photot ransformat ion of the
init ial red-light ( R ) absorbing fo rm ( Pr, max ≈
666 nm) into a physiolog ically act iv e far -r ed-light
( FR) absorbing form ( Pfr, max≈730 nm ) , so the
pr imary photoreact ion is the star t for us to learn
the photochrome signall ing mechanism.
Fo r that the past decade has seen dramat ic ad-
vances in our know ledge of phytochr omes and their
signal t ransduction pathw ays that leads to various
physiolog ical r esponses, there are many review s on
the cell biolog y of phy to chrome signalling
[ 1 4] , but
few sy stemat ical literatur e on phy to chrome pho to-
tr ansformat ion and the corresponding signall ing
mechanism. Here, we briefly review the most re-
cent prog ress on this topic that maybe pr ovide new
insights into leaning phy to chrome signal t ransduc-
tion.
1 Phytochrome Structure and Function
The major achiev ement of recent years is the
discovery o f a small fam ily of genes encoding for
several phytochrome apoproteins, e. g . , phyA -
phyE in A rabidop sis thaliana. ( Shar rock and
Quail , 1989 ) , Ar abidop sis phyB and p hyD
po lypept ides are approx . 80% identical and are
somewhat mor e related to phyE than they ar e to ei-
ther phyA or phyC ( approx . 50% ident ity ) .
Counterparts of p hyA, phyB and o ther PHY genes
are present in most , if no t all, higher plants
[ 5] .
All of the higher plant phy tochromes share
the same basic st ructure, consist ing of tw o struc-
tural domains: a photosenso ry: g lobular N-term i-
nal chromophore-binding domain w hich is suff i-
cient for light absorption and photoreversibil ity
( - 70 kDa ) , and a r egulatory , conformationally
mor e extended C-terminal domain ( - 55 kDa )
w hich encompasses tw o histidine kinase related do-
mains ( HRKD) and two mot if s w ith homology to
PAS ( PER-ARNT -SIM ) domains
[ 6, 7]
. PAS do-
mains are sensible to environmental signals such as
light condit ions, o xygen levels, and redox po ten-
tial. T hey may also mediate protein-protein inte-
r act ions
[ 8, 9]
. The amino-terminal half o f phy-
tochr omes can be considered as a light-sensing do-
main whilst the carboxy l-term inal half can be re-
garded as the regulatory domain[ 10] .
In the 1980s, spectrophotometric studies in-
dicated that there ar e at least tw o dist inct pools of
phytochromes, Type I ( l ig ht labile ) and type II
( l ig ht stable) . T ype I phytochrome is synthesized
as Pr in darkness and decay rapidly in the light as a
labile Pfr form . In contrast , type II phytochrome
is stable in the Pfr form and is present at relat ively
constant lev els both in the light and in darkness.
Now it is accepted that phyA corr esponds to the
major light-liable pigments ( type I ) , w her eas the
m inor light-stable pigments ( type II ) comprise
phyB and the other phytochr omes. Ther e is certain
overlapping betw een the absorpt ion spect ra of Pr
and Pfr , w hich is impo rtant fo r phyA act ivity be-
cause phyA is light dependent and r equires selec-
t ive recognit ion and ubiquit inat ion of P fr [ 11, 12] .
PhyA part icipates in seed germination, responsible
in a w ay of the very low f luence r esponse ( VLFR,
10- 4 to 10
- 1molm - 2s- 1 ) or the high-irradiance
r esponse ( HIR, > 1 000 mo lm- 2 ) in seedling de-
et iolat ion, including inhibition of hypocotyl elon-
gation, the expansion of cotyledons, changes in
gene expr ession and the synthesis of anthocyanin,
etc. On the o ther hand, phyB and the other phy-
tochr omes control the pho to rev ersible ef fects of
the so-called low-fluence responses ( LFR, 1 to
103 molm - 2) [ 11, 12] , w hoes distinguish feature is
it s conformity to the Bunsen-Roscoe Recipro city
Law , w hich states that a response should be de-
pendent only on the to tal amount of photons re-
ceived irrespective o f the durat ion of the expo-
sure[ 13] . Phy to chrome B is also considered to be
the main phytochrome r esponsible for the shade
avoidance response ( elongated gr ow th habit , re-
duced leaf area , incr eased apical dominance and
early flowering
[ 14]
) . The phytochr ome fam ily
members play different roles in the pho to regulation
processes, and phy to chromes also show redundan-
cy o f funct ion. Clearly, phytochromes also interact
and coact w ith other photor eceptors
[ 15]
.
538 武 汉 植 物 学 研 究 第 21 卷
2 Phytochrome Phototransformation and
Phytochrome Heterogeneity
T he photochem ist ry of phytochrome act ivat ion
is based upon the cis-tr ans isomerizat ion of the
chromopho re. T his conf igurat ional t ransit ion leads
to the fo rmat ion of a signal ling state of suf ficient
stabil ity to communicate the presence of photons to
a downst ream signal t ransduct ion partner . There-
fo re, phy tochrome pho to tr ansformat ion can be de-
fined as the start of the primary pho to react ion,
w hich is involv ed in the start o f signal t ransduc-
tion.
The Pr Pfr conversion is a low-energ y stor -
ing photoreact ion based on photobio logical criter ia,
of w hich primary pro cesses are a Z, E isomer iza-
tion around the C15 = C16-methine bridge of the
tet rapyrro lic chromophore pro ceeding w ithin seve-
ral tens ps w ith part icipat ion of the sing let ex ited
state
[ 16, 17]
, and at almost the same time the struc-
ture of apopro tein is conformationally restruc-
tured. A number of spect rophoto-metrically ident i-
fiable intermediates have been detected ( Fig . 1) .
Lum i-R ( f< 100 s, ca. 700 nm) is the f irst inter -
mediate stable at low temper ature accompanying
the t ransformat ion process, w hich decays on a mi-
crosecond t ime scale to meta-Ra. Interestingly , pi-
cosecond kinetic measurements on phyA reveal a
decay of the Pr excited states ( Pr
* ) w ithin about
15-40 ps to prelum i-R which in its turn decays w ith
a delay ( up to 100 ps) to the Lumi-R ( state 1, pho-
toequilibrium betw een Pr and lumi-R) w here the Pr
→Pfr conversion is t ransitorily stopped [ 17, 18] . It is
w o rthy to note that the decay kinet ics of the lumi-
R are significant ly dif ferent between phyA and
phyB
[ 19]
, and the quantum yield for the formation
o f lum i-R is higher than Pr→Pfr react ion, so the
changes in the Pr→lum i-R is complex [ 20] . How ev-
er, some scho lars concluded that ther e must be the
ex istence of the heterog eneity o f the em itt ing
species produced from the Pr ground state popula-
t ions, several pools of lum i-R and the excited
states of the sever al pools
[ 17, 21, 22]
, because interac-
t ion of the chromorpho re w ith the apoprotein obvi-
ously af fects the fo rmat ion o f g round state lumi-R,
and whatever phy sical o r chemical process may be
involv ed therein. Furthermor e, it is suggested that
Z-E/ E-Z isomerizat ion takes place during the Pr→
Fig. 1 Simplif ied energy level diagram of the phytochrome photocycle ref lecting
539 第 6 期 张鸿明等: 光敏色素分子特性及其信号转导机制(英)
lumi-R, and lumi-R may act physio logical ly as
funct ional intermediate, w hereas C15 single bond
rotat ion occurs in one o f the subsequent steps. For
instance, Pr′( o ne of the tw o phenomenolog ical Pr
types ) is more ef f icient in the Pr→lum i-R pho to-
conversion. So, it is possible that the sho rt-lived lu-
mi-R state could affect the concentr at ion of the
physiolog ically act ive Pfr by chang ing the photocy-
cle of the Pr→Pfr , or possibly , the following pro-
tein conformational changes [ 23] .
As w e all know , the physiolog ical effect of P fr
is related to the abso lute or relat ive concentrat ion
of P fr or the stabilizat ion of Pf r in plant t issues,
and Pfr should be subject to rapid changes due to
it s degradat ion and dark r eversion into Pr
[ 24]
. The
Pfr photor ev ersion r eact ion including the thermal
reversion in the dark is very important in maintai-
ning a physio logical balance in plant development .
Recent ly, the r everse Pfr photoreversion is repor -
ted to be sequentially followed by several other
meta intermediates such as the first intermediate,
lumi-F ( f≈320 ns, ca . 673 nm ) , and the second,
meta-F ( f≈265 s, ca. 660 nm ) [ 22] , and is des-
cr ibed as a bi-exponent ional process with tw o t ime
constants in the range between 150 - 650 fs and
2- 5 ps
[ 25, 26]
. N otably , Pr photot ransfo rmat ion
pathw ay does no t share any intermediate w ith the
Pfr photorever sion pathw ay ( Fig . 1) , w hich may
be also seen from the different spect ra of the inter -
conversion. It seems that al l these intermediates
have no obv ious physiological act ivity, but they
are necessar y for the interconversion betw een Pfr
and Pr upon irradiat ion and passing photons to a
downstream signal t ransduct ion par ticle. Compa-
ring w ith photosynthet ic pho tons t ransfer path-
w ay , We lit t le know how phy tochrome percept
photons and the primar y photosignal tr ansduct ion
pathw ays, but these intermediates may be just the
implicat ion. Recent studies f rom Furuyas lab sug-
gest that the FR-HIR requir es a short-lived inter -
mediate( Pr
+
) g enerated dur ing Pfr to Pr pho tore-
version. Most interest ing ly, this r esponse is FR/ R
reversible ( not R/ FR) , w hich is oppo site to w hat
is seen in the LFRs such as FR inhibit ion of light-
r egulated gene expression or let tuce seed germ ina-
t ion
[ 27]
.
Recent success in investig at ion is connected
w ith the discover y of heter ogeneity of phytochrome
in the cell observed using the low-temperature
( 85 k) fluorescence spect roscopy as w ell other ex-
per imental approaches
[ 26, 28, 29] . Tw o Pr types were
dist inguished physiolog ically , Pr′-major longer
w aveleng th ( 687/ 673 nm, emission/ absorption
maxima ) and Pr″-minor, shorter w avelength ( 682/
668 nm ) . It w as show that Pr′and Pr″correspond
to type I and type II, respect ively
[ 3 0, 31]
. Subse-
quent ly , Sineshchekov obtained the experimental
evidence that phyA is heterogeneous and compr ises
tw o spect roscopically distinct species, namely , the
l ight-labile phyA′( belong s to Pr′) and the light-
stable phyA″( belongs to Pr″) [ 28, 32] . PhyB w as la-
ter show n to have char acter ist ics close to those of
phyA″and belongs to the same phenomenolog ical
Pr″type. Schmidt also found that the ex istence of
Pr isofo rms by investig at ion of thermochromy tem-
per ature-induced abso rpt ion spectra changes[ 33] .
Al though the exact str uctural differ ence betw een
the tw o Pr′types is not known, it is assumed that
they could mediate different photor esponses.
Based on his long-run invest igat ion, Sineshchekov
concluded that phyA′should be responsible fo r de-
et iolat ion w hile the invarible phyB and phyA″could
funct ional through the plant life cy cle[ 31] . Nonethe-
less, the results w as just achieved by spect roscopy
analysis, not biochemical and genetical method,
l it t le is current ly known regarding Pr′o r Pr″sig-
nalling pathw ays, st ill less the downst ream com-
ponents. So to say, t ransduction of the light-
generated signal and ident ificat ion of the various
phytochrome signal t ransduct ion pathw ays are st ill
a major field in the com ing future.
3 The Cell Biology of Phytochrome Sig-
nalling Transduction
The central quest ion of the biochemical tr an-
sact ion that const itutes signal t ransfer from the
photoact ivated phytochrome molecule to it s prima-
ry signalling intermediates has int rigued re-
540 武 汉 植 物 学 研 究 第 21 卷
searchers in the f ield for many years
[ 34]
.
3. 1 Phytochromes as light-regulated kinase
The car boxy term inal domain of polypept ides
contains a region w ith sequence similarity to
pr okar yot ic tw o component hist idine kinases,
w hich suggested a biochem ical mechanism fo r phy-
to chrome signaling . T he Lagarias laboratory re-
cent ly has pro vided convincing evidence that phy-
to chrome funct ions as a photoreceptor kinase ( an
unusual Ser / Thr kinase w ith tw o His kinase-like
domains)
[ 35]
. The data showed that PKS1, as dif-
ferent ial tr ansphosphorylat ion o f a protein sub-
st rate, w as pho spho rylated on serine and threonine
residues, and that conversion of phyA to the act ive
Pfr fo rm enhanced the level of PKS1 phosphory la-
tion by tw o mor e fo ld relative to that observ ed for
Pr . Par k et al. ( 2000) has speculated that phos-
phory lation of phy to chrome may contribute to the
Pr to Pfr confo rmat ion change which may imply
that phosphory lat ion pat terns may be involved in
the retention and r elease of phytochromes[ 36] . The
pr otein subst rate, PKS1, may play a role in this
retent ion mechanism result ing in signal t ransduc-
t ion.
3. 2 The primary signaling intermediates
Early pharmacolog ical studies using micro in-
ject ion of a tomato phy to chrome mutant have iden-
t ified heterot rimeric G pr oteins, cGMP and Ca, as
second messengers in phytochrome signaling
[ 37, 38] .
Genet ic screens have ident ified tw o classes o f sig-
naling components, those act ing down-st ream of a
sing le photoreceptor ( e. g . PIF3) and tho se act ing
downst ream of mult iple pho to receptors ( see Fig .
2) . T his presumably reflects the fact that light
signals per ceiv ed by dif ferent pho to receptors must
be integ rated. The lat ter class includes bo th posi-
t ively act ing factor s ( i. e. HY5) and a larg e gr oup
o f negat ive regulator s of pho tomorphogenesis
( DET / COP/ FUS) . How do these various compo-
nents implicated in photosignal transduct ion func-
t ion? The answ er remain unkown although many
components have now been cloned. How ever, it is
notable that the cloned factors lo calised to nucle-
us, w hich suggest that ear ly light signall ing events
are nuclear-localised.
Fig. 2 A simplified model for phytochrome-mediated light signaling( Quail, 2002a)
541 第 6 期 张鸿明等: 光敏色素分子特性及其信号转导机制(英)
4 Discussion and Perspectives
Investig ations w ith A rabidop sis mutants null
fo r the individual phy tochrome and signalling com-
ponent since 1994, significant progr ess has been
made in unr aveling these components in the signal
t ransduct ion pathw ays. Coupled the phy tochrome
dynam ic photot ransfo rmat ion with follow ing signal
t ransduct ion, it is known that phy tochrome
molecule funct ion as unique binary optical stor ag e
devices w ho se biochemical output is contr olled by
the stored informat ion. Although a detailed kinet ic
scheme has not yet been elucidated, these data may
pr ovide some information for phytochrome mult idi-
mensional signal pathw ays. Fo r instance, Pr′or
Pfr
*
can be also an another signalling intermediate
and exert a signif icant ro le through the develop-
ment . To fur ther study the signal pathw ays.
Based on all these ex ist ing literatures, it is
tempt ing to ant icipate that the intermediates have
the effects on the later signal t ransduct ion or that
the bio act iv ity of Pf r ( or Pf r-x ) is light-dependent
because o f light-induced nuclear t ranslocat ion.
Conformat ional changes of the intermediates inf lu-
ence the subcellular localizat ion, stability , as w ell
as the protein kinase act ivity o f phytochr ome. It is
also proposed that the intermediates and the ab-
sor bt ion of pho tons by photoreceptors change their
conformat ions, result ing in phosphory lation of the
receptor s, w hich enventually t rigg ers signal tr ans-
duct ion and phy siolo gical responses. T her efore, it
seems that the urgent task is to f ind the factors af-
fect ing the stabilizat ion of P fr or Pf r-x form by in-
hibition of the dark rever sion of Pf r ( or Pf r-x ) to
Pr , and determine w hether this step is light-depen-
dent or l ight-independent . M eanwhile, some so-
phist icated experimental approaches may be ap-
plied to determine w hich step during pho to tr ans-
fo rmat ion f rom Pr to Pfr a signal is t ransduced
fr om the photor eceptor in the cell. How ever, the
dif ficulty is that ther e are dif ferent phytochromes
and their corr esponding heter ogeneous subpopula-
tions including the intermediates dur ing the pr i-
mary pho to tr ansformat ion. Luckily, various phy-
tochr ome mutants and signal component nul l mu-
tants are available as model plant to exploit the
mechanism coupling w ith devising light condit ions.
So tomo rrow is light !
Ref erences:
[ 1 ] Smith H. Phy tochromes and light signal per ception
by plant s—— an emerging synt hesis [ J] . Natur e,
2000, 407: 585 591.
[ 2 ] M ller S G , Ingles P J, Whitelam G C. T he cell
biolog y o f phyt ochrome signalling [ J] . New Phy tol,
2002, 154: 553 564.
[ 3 ] Quail P H. Pho tosensor y per ception and signalling in
plant cells: new paradigms? [ J ] . Cur r Op in Cell
Biol, 2002a, 14: 180 188.
[ 4 ] Quail P H. Phy tochr ome photo sensor y signalling
net wo rks [ J] . N at Rev Mol Cell Biol, 2002b, 3:
85 93.
[ 5 ] Mathew s S, Shrr ock R . A Phy tochr ome gene diver-
sity [ J] . P lant Cell E nv ir on, 1997, 20: 666 671.
[ 6 ] Lagar ias D M , Wu S H , Lagar ias J C . A typical
phy tochr ome gene str uctur e in the g reen alga Meso-
taenium caldariorum[ J] . P lant Mol Biol , 1995, 29:
1 127 1 142.
[ 7 ] Kay S A . PAS, pr esent , and future: clues to the
or ig ins o f cir cadian clocks [ J] . Science, 1997, 276:
753 754.
[ 8 ] Quail P H. An emerging mo lecular map of the phy-
tochromes[ J] . Plant Cell Environ, 1997, 20: 657
665.
[ 9 ] T aylor B L , Zhulin I B. PAS domains: internal sen-
so rs of ox ygen, redox po tential, and light [ J] . Mi-
cr obiological Molecular Biolog ical Rev iew , 1999,
63: 479 506.
[ 10] Quail P H, Boy lan M T , Parks B M , et al. Phy-
tochromes: pho tosensor y per ception and signal
tr ansduction[ J] . S cience, 1995, 268: 675 680.
[ 11] Clough R C, Jo rdan-Beebe E T , Lohman, K N , et
al. Sequences w ith-in t he N- and C-t erminal
domains of phy tochrome A are r equir ed fo r PFR
ubiquitina tion and degr adat ion [ J] . Plant J , 1999,
17: 155 167.
[ 12] Hennig L , Bche C, Eichenberg K , et al . Dynamic
pr oper ties o f endogenous phyto chrome A in A ra-
bidopsis seedling s [ J ] . P lant Phy siol, 1999, 121:
571 577.
[ 13] Mancinelli A L . The physio log y o f phyto chr om e ac-
tion[ A ] . In: Kendrick R E, K ronenber g G H M eds.
Pho tomo rphogenesis in P lant s [ C ] . Do rdrecht, The
Netherlands: K luwer Academic Publishers, 1994.
542 武 汉 植 物 学 研 究 第 21 卷
211 269.
[ 14] Smith H, Whit elam G C. T he shade avoidance syn-
drome: multiple r esponses m ediat ed by multiple
phyto chromes [ J] . P lant Cell E nvir on, 1997, 20:
840 844.
[ 15] Casal J J. Phy to chr omes, cr yt ochromes, pho-
to tr opin: phot or ecepto r inter actions in plants [ J ] .
P hotochem photobiol, 2000, 71( 1) : 1 11.
[ 16] Schaffner K , Brasla vsky S E , Holzwar th A R.
Photophy sics and photo chemistry of phyto chr ome
[ A ] . I n : Volman D H , Hammond G S , Go llnik K
eds. Advances in Pho tochemistr y [ C] . New Yo rk:
W iley, 1990. 229 277.
[ 17] L ippitsh M E, Hermann G , Brunner H, et al. P i-
cosecond events in the photo transfo rmat ion of phy-
to chr ome-a t ime-resolved absorption study [ J ] . J
P hotochem Photobiol B : Biol, 1993, 18: 17 25.
[ 18] L ippitsh M E , Rieg ler H, Aussenegg F R, et al. P i-
cosecond absorption and fluo rescence studies on
lar ge phyto chr ome fr om rye [ J ] . B iochem Phy siol
Pf lanz , 1988, 183: 1 6.
[ 19] Sineshchekov V A . Phot obiophy sics and pho tobio-
chemistr y o f the heter og eneous phy tochrome system
[ J ] . Biochim B iophy s A cta, 1995, 1 228: 125
164.
[ 20] Andel F, Lagar ias J C, M athies R A . Resonance r a-
man analy sis o f chr omopho re str uctur e in the lumi-R
photoproduct o f phy tochr ome [ J ] . Biochemistry ,
1996, 35: 15 997 16 008.
[ 21] Holzwa rth A R , Venuti E, Braslav sky S E, et al.
T he phot otr ansfo rmation process in phy tochrome:
ultrafa st fluo rescence component and models fo r t he
int ital Pr → Pfr tr ansfo rm ation st eps in nativ e phy-
to chr ome [ J] . Biochim B iophy s A cta, 1992, 1 140:
59 68.
[ 22] Chen E, Lapko V N , Lew is J W , et al. Mechanism
o f nativ e oat phyto chr ome pho tor ever sion: a t ime-
resolved abso rption investig ation[ J] . Biochemistry ,
1996, 35: 843 850.
[ 23] Heyn M P , Borucki B, O tto H. Chromopho re r eor i-
entation dur ing the photo cycle o f bact eriorhodopsin
experiment al methods and functional significance
[ J] . Biochimica B iophy sica A cta, 2000, 1 460: 60
74.
[ 24] Clugh R C , V ier stre R D. Phyto chr ome degr adation
[ J] . P lant Cell E nvir on, 1997, 20: 713 721.
[ 25] Bischoff M , Hermann G , Rentsch S, et al. F irst
steps in the phyt ochrome photo tr ansfo rma tion: a
com parative femtosecond st udy on the forward ( P r
→ Pfr ) and back reaction ( Pfr →Pr ) [ J] . B ioche-
mistry , 2001, 40: 181 186.
[ 26] Remberg A , L indner I , L amparter T , et al. Raman
spectr oscopic and light -induced kinet ic char acter iza-
tion of a recom binant phy tochr ome o f the cyanobac-
terium synechocy stis [ J ] . Biochemistry , 1997, 36:
13 389 13 395.
[ 27] Shinomura T K , Uchida M F . Elementar y r esponses
of pho toperception by phy tochr ome A for high ir r a-
diance r esponse of hypoco tyl elongation in A rabidop -
sis thaliana [ J ] . Plant Phy siol, 2000, 122:
147 156.
[ 28] Sineshchekov V A . Pho tobiophysics and phot obio-
chemist ry of t he het ero geneous phyto chr ome sy stem
[ J] . Biochim Biop hy s A cta, 1995, 1 228: 125 164.
[ 29] Bischo ff M , Hermann G , Rentsch S, et al. Fir st
steps in t he phy to chr ome pho tot ransformation: a
comparat ive femto second study on the fo rw ard ( Pr
→ Pfr ) and back reaction ( Pfr→ Pr ) [ J ] . Bioche-
mistry , 2001, 40: 181 186.
[ 30] Sineshchekov V A . Tw o spectr o scopically and pho-
tochemically dist inguishable phyto chr om es in etio lat -
ed seedling s o f monocots and dico ts[ J] . P hotochem .
Photobiol, 1994, 59: 77 86.
[ 31] Sineshchekov V A , Ogor odnikova O B, Weller J L .
Fluor escence and photo chem ical proper ties of phy-
tochromes A and B in etio lo ated pea seedling s[ J] . J
Photochem Photobiol, 1999, 49: 204 211.
[ 32] Sineshchekov V A . Evidence for the ex itense o f tw o
phy tochr ome A populations[ J] . J Photochim Photo-
biol B : Biology , 1995, 28: 53 55.
[ 33] Schmidt P , Gensch T , Remberg A , et al. T he com-
plex ity o f the Pr t o Pfr phot otr ansfo rm tion kinetics
is an intr insic proper ty of native phyto chr ome [ J] .
Photochem Photobiol, 1998, 68( 5) : 754 761.
[ 34] Quail P H. Phy to chr ome inter act ing factor s. Semin
[ J] . Cell D ev Biol , 2000, 11: 457 466.
[ 35] Fankhauser C, Yeh K C, Lagar ias J C, et al. PKS1,
a substr ate phospho r ylated by phyto chr ome that
modulat es light signaling in A rabidop sis [ J ] . Sci-
ence, 1999, 284: 1 539 1 541.
[ 36] Park C M , Bhoo S H, Song P S. Inter-domain
cro sstalk in the phyto chr om e molecules [ J] . S emi-
nar s in Cell and Develop mental B iology , 2000, 11:
449 456.
[ 37] Neuhaus G , Bow ler C, Kern R , et al. Calcium /
calmodulin-dependent and-independent phy tochrome
signal tr ansduction pathw ays [ J] . Cell, 1993, 73:
937 952.
[ 38] Bow ler C, Nuehaus N , Chua N H. Cy lic GM P and
calcium mediate phyt ochrome pho tot ransduction[ J] .
Cell, 1994, 77: 73 81.
543 第 6 期 张鸿明等: 光敏色素分子特性及其信号转导机制(英)