全 文 ：广 西 植 物 Guihaia 22(2)：160— 166 2002年 3月
膜对采后园艺作物衰老的影响
蒋跃明 ，傅家瑞。，徐礼根。
t 1 中国科学院华南植物研究所．广东广州 5i065o；2 中山大学生命科学学院，
广东广州 5i 0275；3 浙江大学生命科学学院．浙江抗州 31001 2)
摘 要：植物衰老是由内在和外部环境因素所诱导和引起的一种主动过程 ，其中膜 的劣变是这种过程早期
的表现形式和基本特征 简要评述了膜系统对采后固艺作物的衰老作用 有证据表明膜系统破坏是采后园艺
作物衰老的关键事件。膜脂生物化学和生物物理性质的变化、酶促和非酶促的过氧化作用导致 了膜特性的改
变和功能上的失调(如离子泄漏和细胞去区域化)。这样 ，反过来叉进一步引起细胞代谢异常 ．并加速衰老进
程。目前．膜系统对采后园艺作物衰老的作用过程正逐步得到阐明。
关键词：固艺作物；膜 栗后 ；衰老
中国分类号：$31l 文献标识码 ：A 文章编号 1000—3142(2002)02—0160—07
M embrane effects in postharvest senescence
of horticul tural crops
JIANG Yue—ruing ，FU Jia—rui ，XU Li—gen。
L 1 South Chitin l,~sd~ute of Bomnyt Chinese Academy of sc Lcts．Guangzhou 510650-China 2．s ktmt【_f Ltjc．Zhongahan
Cnlverai~2,．Guangzhou 510275．China；3 Sc&~l LL／~．Zkediang University，Hangzhou 316012，China／
Abstract：Senescence is an active progress initiated by internal and envi ronment triggers，and rmmbrane deterio
ration is n early and fundamental feature of this process． Advances 【Ⅱ merFlbrRile effects in senescence 0f
postharvest horticultural crops are reviewed Considerable evidence for senescence of postharvest horticultural
crops supports membrane damage gis the key event．Bioehemieai and biophysical changes in membrane lipids and
eazymatie and non enzymatic lipid peroxidation lead to altered membrane properties and defects．such as ion
leakage and cellular decompartmentation-which r in turn—result in abnormality in cellular metabolism and accel
crated death A cleare r picture of the aequence of occurrences in effect~of~[1et：nbranes on postha rvest senescence
。f horticultural crops is FlOW emerging
Key words：horzicultural crops}membranes；postharve~t}sene cnc
1 Introduction
Aging or senescence of higher p[anls is viewed as
a both genetically and environmentally regulated pro
cess intimately associated wilh physiotogical aetivi—
ties．Postharvest horticultura1 scientists are jnterested
in determining the mechanisms of senescence in fresh
produce in order to find treatments ihal wilt atlow
shelf life to be prolongcd．1l has been suggested that
deterioralion of fruits， vegetab[es，and flowers by
收稿 日期 2000—05—08
Author introduction JIANG Yue ming(1 063一)．Male-bora in Laaxi city．Zhejiang province
working in postharvest bio{ogy aad techno{ogy oI horzi cultura{crops
维普资讯 http://www.cqvip.com
2期 蒋跃明等：膜对采后园艺作物衰老的影响 1 6l
natural senescence or aging due to physiological ab—
normalities．share a CO~lnlon mechanism： The fo_
C／IS of this proces is thought to be alterations in the
physica】and chenfica1 characteristics of rrlerlbranes，
which， 】n turn， lead to abnormality in cellular
nletaboisnl and accelerated death．
SeveraI reviews on senescence and membrane de
lerioration in plants ～ and on membrane effects【n
postharvest physiology： are available．In this review
x
．ve shall present，based on select reports，the state of
current research on membrane effects jn senescence of
postharvest horticultural crops．
2 Properties of plant membranes
Celular membranes are selective，dynamic barri—
ers that play an essential role in regulating b[ochemi
cal and physiological events．A detailed description of
plant nlembrane composition as wel as their physical
properties was reviewed by Boroehov and Wood
son ～
， Stanley ，I eshem ，and Paliyath and Drol
lard ．Present knowledge of plant membranes[denti—
ties them as fluid b[1ayers of phospholip[ds contaming
embedded proteins and sterols Funct[onal uleu1
hranes are fluid since phospholip[ds Cal move rather
freely in lateral dimensions，but the presence of pro—
reins and sterols influences membrane fluidity． The
conlpos[tion of const[tuent membrane Iip[ds can affect
the fluidity of membranes， with unsaturated fatty
add—containing lip[ds being more fluid than saturated
lip[ds．The concept of fluidity is of great importance
to both menlbrane function and deterioration．During
senescence，there is a progressive loss of membrane
integrity and changed properties of plant tnetn—
brant
3 Bulk lipid fluidity
it is nov, well documented for a variety of
senesc[ng 1issues，including flowers and fruits，that
the plasma and microsomal membranes sustain a de
crease in bulk lipid fluidity with advancing age．In
senescing tissues that show a climacteric like rise in
ethylene production，the decrease in membrane fluid[一
ty，which occurs abruptly，is coincident with or jUSt
prior to the rise in ethylene prod uction ．” e rune
tions of Nle~ibrane proteins are known to be sensitive
to lipid fluidity ，and thus a change in bulk lipid flu
idity sustained by plasma and mierosoma]membranes
during senescence 【s l[kely to have a deleterious im—
pact on the functions of membrane—associated en—
zymes and receptors．
4 Lipid composition
The UtOSt conspicuous change in the lipid eompo
sition of senescing plasma and microsomal membranes
is a dramat【c decline in membrane phospho[ip[d，
which becomes man[fest as a membrane stero1：phos
pholip[d ratio increased． The rise in sterol：
phospolipid ratio of senescing mierosomal and plasma
membranes reduces b[1ayer fluidity ． For some
senesc[ng membranes．there 【s very little change m
saturated unsaturated fatty add ratio ，but for oth
ers，including microsomal membraHes from senesc[ng
carnation flowers，there is a significant decline in the
1evel of unsaturat[on that can be attributed to nlenl—
branous l[poxygenase： Sterol content in senescing
membranes Cal either jncrease，as observed in flow—
ers ” and fruits ． or remain constant
Changes in phospholip[ds and sterol contents also
bring about increase【n membrane microv[scos[ty 【n
apples ’ ．tomato and broccoli
5 Phase changes
Increased proportions of gel phase lipid appear in
the membrane bilayers as senescence intensifies ．
Thus．senesc[ng membranes may contain a mixture of
liquid—crystalline and gel phase lipid domain， and
these coexist[ng domains render the membrane leaky
and contr[bute to the 1oss of intracellular compart
mentat[on that is a feature of senescence⋯．The or—
mation of gel phase lipid【n membranes during senes—
cence is more apparent from measurements of the
lipid phase transition temperature． The transition
维普资讯 http://www.cqvip.com
1 62 广 西 植 物 22卷
temperature rises progressively with advancing senes-
cence 9J．
6 Membrane protein
There is evidence that the progressive destabi
[ization of membrane bilayers accompanying senes—
cence contributes to loss of membrane protein rune
tion．Protein degradation is an important feature of
the dismantling of membranes，and there is a grow-
ing conviction that selective degradation of membrane
proteins occurs during senescence。 For exanl—
ple．the number of the receptor protein for ethylene，
which is thought to be associated with membranes，
decreases with advancing senescence ． It is also
conceh,ab[e that free radicals， which produced
through membrane—associated lipoxygenase， act di—
rectly on proteins⋯ ．W ith the onset of senescence．
the chemical composition changes of membrane[ipids
may result in lipid phase separations within the biIay—
er．Evidence is described indicating that lipid metab c>
lites are released from membranes as lipid protein
particles under normal conditions but accumulate in
the biIayers of senescing membranes coincident with
impairment of lipid protein particle formation ．
7 Permeability
As a result of senescence．there is an【ncreased
permeability，which，in turn，leads to alterations in
cdblar metabolism 。,a 7．Increased leakage correlates
with【ncreased membrane viscosity and decreased de—
gree of fatty acid unsaturalion，which is associated
with increased phase transition temperatures of mem—
brane tipids and a decline in fluidity in the senescence
of flowers ㈤． 1一M ethykyc【0pr0pene，an anti—
ethylene compound，which inhibits ethylene—induced
celular senescence．results】n a delay【n an】ncreased
electrolyte leakage These data support what
membrane deleriora~ion leading to toss of intracelular
compartmentation is an inherent feature of senes—
cence．It is not unreasonable，therefore，to propose
that progressive destablization of the bilayer con—
tributes in a major way to membrahe leakiness in
senescm g tissue-
8 Frei5 radicals and peroxidation
Peroxidation of fatty acids with resulting free
radica1 format【on has been described as one of the ma—
Jor senescence processes of membranes ⋯ ．En—
hanced peroxidation，observed in a variety of senese
ing tissues，leads to gel phase formation and los in
membrane functionality ’‘”．Changes in membrane
[ipids resuhing in decreased fluidity wil affect the
functionality of the associated proteins as well ．
Various studies demonstrate that fruit ripening and
the vase life of flowers can be modulated by radical
scavengers Membranes could he expected to be
highly prone to free radical attack inasmuch as unsat—
urated fatty acids are major components of most
membrane lipid bi[ayers． The consequences of free
radical attack on membranes are numerous and jn—
elude the induction of lipid peroxidation and fatty acid
deester_[jcat 0n ‘ ．Of particular nterest【s the find
ings thai levels of superoxide radical in microsoma[
membranes of carnation flower increased during
senescence 。
． Additionaly，this increased produc—
tion of superoxide radical during natural aging can be
simulated by in vitro aging of isolated microsoma[
membranes from young tissue in buffer ．The for
marion of superoxide radical by senescing microsomal
membranes has been attributed to a membrane—assc~
ciated [ipoxygenase ．Baker et at。 ．reported that
specific inhibitors of[ipoxygenase significantly delay
the senescence of cut carnation ftowers． Thus， t
woutd appear that superoxide radical is formed as an
intermediate during the conversion of polyunsaturated
fatty acids to their conjugated hydr。pef0xyd【ene
derivatiyes．
9 Ethylene
it is now known thai ethylene plays an impor
tant role in regulating plant senescence． Ethylene
production is regulated by celular membrane ．One
维普资讯 http://www.cqvip.com
2期 蒋跃明等：膜对采后园艺作物衰老的影响 163
mechanism 】s a direct effect on the enzymes that were
associated with membranes． The other jnvo[ves the
cellular membrane as a transducer whereby de novo
synthesis of the enzymes in ethylene biosynthesis is
affected． M[crosomal membranes from plant tissues
have been found to be capable of catalysing the con—
version of 1-aminocyclopropane—-1—-carboxylic acid
(ACC)to ethylene ” ． The conversion of ACC to
ethylene by microsomal membranes depends on hy
droperox[des generated through membrane—associated
lipoxygenase ．Treatment of microsomes with ex—
ogenous phospholipase A2，which generates fatty acid
substrates for lipoxygenase，enhances the production
of hydroperoxides as well as the conversion of ACC to
etby[ene⋯ However，these results do not distin—
guish between a direct and an indirect effect of ethy
lene on the peroxidative pathway leading to changes
1n membrane fl_netion．
10 Ca
An increasing amount of evidence points to Ca。一
regulating plant senescence ”． Calcium or the
calcium—calmodul[n complex can activate lipid— degrad—-
ing enzymes．Furthermore，the Ca 一calmodulin com-
plex is required for protein phosphorylation Phos-
phorylation of proteins promoted by the Ca 一calmod—
ultn complex is reduced in senescent apples and toma—
toes，and this COHid alter the biochemical processes in
the cell and participate in the onset of senescence ．
As numerous authors have shown a relationship be
tween ethylene and the induction of senescence ⋯ ，
research to determine【f Ca 【s able to translate the
ethylene signal and initiates membrane lipid degrada·-
tion has been undertaken ．To date，the‘‘trigger’
through which calcium is released into the cytosol and
seneS亡ence is in【tiated js stil unclear．
Triggers
／ ⋯ 一 ’
Formation ofgel
Hydroc~bons and fai ry aldeh de
(Phospho)lipases
Free国【 acids
{
mdicals {Lipuxygena~s
＼ l
Feroxidized fatly acids
Fig．1 Generalized scheme to aceount for membrane deter[oration in po㈣tha est horticultural
c『。Ds due to seneseence． Adapted in part from Marangonia t a1． and Thompson t“ ’
1 1 Nonbilayer lipid
There is evidence for the induction of nonbdayer
lipid configurations followmg perox[dat[on． Indeed，
many of the products of peroxidafion are not cylindr[
tally shaped because they have bulky side groups(e
g．， bydroperoxy， cyclic peroxide， and epox[de
groups)，and，thus，could promote the conversion to
nonbdayer configurations． I．ip[d peroxidation has
been shown to increase the rate of transNlayer lipid
维普资讯 http://www.cqvip.com
l 64 广 西 植 物 22卷
migration： Moreover，trace amount of hydroper—
oxide derivatives of fatty acids may act as fusogens，
and oxidized di—and tri—enoic fatty acids serve as Ca
ionophores in model bilayers⋯ Thus，it is highly
conceivable that hpid peroxidation，which occurs dur
ing senescence，promotes the formation of nonbilayer
lipid configurations in sensescing membranes．
Recently，a modern approach to the study of
senescence is related to gene expression of enzymes
lnvolved ln lipid metabolism⋯ ． These authors
found that membrane associated enzymes concerning
lipid biosynthetic potential decreased during series—
cence and that second messenger synthesis continued，
maintaining the tissue responsiveness to the environ
ment and hormones． However，more work in this
area is greatly needed．
1 2 Conc]uding remarks
In light of information available，there appear to
be djstlf1guishable mechanisms of senescence for
membranes．These processes produce de-stabilization
and loss of functiona[itv of membrane via biochemieal
and biophysiea J changes in lipids and／or proleins．In
aH cases，attack byenzymes such as phospho[ipase
and[ipoxygenase as wel as free radieal formation has
been reported Specifically，the enhanced free radi
cats initiates lipid peroxidatioa and estabi[ization of
the membrane bilayer，which， n turn，leads to ieak
age，an increased proteolytic act v ty，and a general一
【zed】OSS of membrarte function Thus，senescence of
microsomal members appears to be Lipid driven A
pattern of membrane deterioration accompanying a
series of events during senescence of postharvest her—
ticuhura[cops has been summarized schematically in
Figure 1．
As membranes have an important efect on
postharvest scnescence of horticuhural crops，it is de—
sirable for food and horticultura】scientists to find
treatments that wil allow shelf lire to be prolonged．
The prospects of postharvest life extension of fresh
produce by regulating membrane functions appear to
be promising，either by exogenous chemical applica
tion or genetic manipulation．
References：
：1]MeKersie B D，Senaratna T，Walker M A．et a／．De
terioration of n]e~]branes during aging in p Lants：evi
dence for free radica[mediation[A] In Nooden I D．
I．eopo[d A C eds，Senescence and Aging in Plants
M] Academic Press．San Diego，California：1988．
441 464．
(23 Stan Ley D W Biological membr．ane deterioration and
associated qua Lity Losses in food tissue[J] Crit Rev
Fred Sci N“ ，1991，30：487 553
[3]Thompson J E．Froese C D，Hong Y，eg al Mem—
brahe deterioration du ring senescence[J]．Can BoL，
1997．75：867 879．
[4]TEompson J E．The motecutar basis for membrane de
teriorafion during sencscenc~[A] In Nooden I_D，
Leopold A C eds-~．nesceltce and ,Aging in P Lants
[M] Academic Press，San Diego，California：1 988．
51 83．
[53 Borochov A ，W oodson W R Physiology and biochem—
istry of flower petal senescence[J] H Rev，1 989，
11：15 43
：6] I．eshem Y Y．Plant Membranes：A Biophysica[Ap
preach to Structure， Development and Senescence
EM：．Kluwer Academic Puhlishers，Dordrecht：1992
1 68
：73 Pa[iyath G．Droiliard M J．The mechanism of rtlet~1．
brane deterioration and disassembly during senescence
_J] Plon~Physiol Biochem，1 992，30 789 8 2
：8]Borochov A，Spieglatein H．Philosoph H S Ed~y[ene
and f Lower petat scncsc nc ： Lnterre【 _1】0nshlp with
membrane lipid catabolism[J]．Physml Plant．1 997，
100：606 610．
：9：Marangoni A G，Palma T，Stan!ey D W Membrane
effects in postharvest physiotogy[J] Postharvest B zoI
T c舶 o1．1996，7：1 93 21 7．
[10 j hzhaki H，Borochov A t Mayak S． Aged—related
changes in petal nlclR1bra：t2 from attached and de
tacked rose flowers[J] Plant Physiol，1 990．94：1
233 —1 236．
[11]Thompson J E，Mayak S，Shinitzky M ， al Accd
eration 0f Foembrane 8eneseence j13 cur earnatIon
维普资讯 http://www.cqvip.com
2期 蒋跃明等：膜对采后园艺作物衰老的影响 l 65
flowers by treatment with ethylene[J2．Plant Physi-
， i982，69：8j9 863
[1 2] Paliyath G， Thompson J E． Evidence for ear[y
changes in membrane structure during post—harvest
deve[opment of cut carnation(Dia~hus caryophyllus
I，)flowers[J]．New Phyto1．1990．114：j55
562．
：1 33 McCormac D J，Todd J F．Patiyath G．“ ．Modu
lation of hibLver fluidity affects Liptd catabotism in
microsoma[merebranes of tomato fruit[J]_Pla*~
Phystbl Bicx-hem ，1993，31：1 8
[i4]Jiang Y M，Chen F Efect of spermidine on regu[a
t[on senescence of【itch】fruit and lts re[ation to ethv一
[erieEJi．Chlnese J Bat，1995，5．22 2j
：l 53 Fobet M ，Lynch D V．Thompson J E Membrane de
terioration in senescing carnation flowers：Co ord[
hated effects of phosphotipid degradation and the ac
tion of membranous lipoxygenase[J3 Pla*~Physi-
o ， l987．85：2O4 211．
[16]Hudak K ，Yao K，Thompson J E．Retease of fluores
cent peroxidized lipids from membranes in senesdng
tissue by blebbing of lipid—protein pa rticles[J：．
HortScf l 995．30：209 2l 3．
：l 73 Faragher J D，Mayak S，Watchel E J Changes in
physical properties of cel membranes and their role
in senescence of rose flower petals[J] Acta H ，
1986，l81：371 375．
[18]Duxbury C I ．1．egge R T ，Pallyath G，el al Alter
ations in memb rane protein conformation in response
to senescence related changes[J]． Phytochem，
1 99 ．3O：63 68
Q g)I uric S Ben—Arie R．Microsomal membrane changes
during the ripeaing of apple fruit[J3 Plant Physio[，
l 983，73：636 638．
：2o3 Rartley T M ．Changes in stero]and phospholipid corn
position of apples during storage at low temperature
and low oxygen concentr~ tion-j ．i sct F∽d
gric，1 986，37：31 36．
[2i]Jtzhaki H ，Borochov A，Mayak S Petal senescence
of robe flowers：a comparison of changes in nletJl
brane related properties in cut and attached flowers
[J] Acta Mort，1 989 261：85 89
~
． 223 M akhlouf J，W iltemot C，Coutu re R， a1．Effecv of
low temperature and controlled atmosphere storage
on membrane lipid composition of brocco[i flower
buds[J[．Sci IIr)r￡，1 990，42．9 1 9．
：23]LeggeR I ，Cheng K H，I epock J R，et a1．Dtfferen
tia[effects of senescence on the molecular organiza—
t[on of membranes in ripening tomato fruit J]_
Plant Physiol·1986，81：954 959．
[24] Deschene A，Pat[yath G，Lougheed E C， et a1．
Memhrane deterioration during postharvest series—
cence of broccoti f Lorets，modulation by temperature
and controlled atmosphere storage ll1：．Postharvest
iol Techn ，1991．1：l 13
(Z52 Davidowiez Grzegorzewsha A，Podstotski A．Age—re
lated changes in tbe uttrastructure and membrane
properties of Brasstca napus I ．seeds[J]_Ann Bo；
(Lond棚 )．1 992，69：39 46．
[26：Hudak K，Thompson J E．Flotation of lipid protein
particles containing triacy[g[ycero[and phosphotipid
Dam the cytosol of carnation peta[s[J] Physiol
Plant，1996，98：8l 0 818
：27]Soudain P，Guclu J，Bureau D，et a1．Purlfication of
plasma membrane and protein composition changes
during ripening and senescence of cherry tomato[J]
．1 Plant Physiol 1992，139：257 264
[28]Sister E C，Serek M Jnhibkors of ethylene responses
in plants at the receptor level：recent development
J¨] Physiol Plant．1997，100：577 j82
[29]Jtang Y M ，Fu J R． Ethylene regulation 。f fruit
ripening：molecular aspects[J]．Plant Grow Regu]，
2008，30： 1 93～ 200
：3。]Yao K，Paltyath G，Thompson J E．T ocalization of
peroxidized ][pids in nonsedimentable mic rovesic[es
of senescing bean cotyledons[J]．，Exp Bot，1 993，
44·l267 l274
[31]Borochov A，Cha M H ，Boss W F Plasma mem—
brahe lipid metabolism of petunia petals during
senescence L J．Physiol Plant，199,1．90：279
284
[323 Yamane K，Abiru S，Fijishige N．et a1．Export of
soluble sugars and increase in membrane permeabili
ty of gladiolus riote rs during senescence[J] ．，，
Sc~ IioFg，l 993，62：575一一58o．
：33j Palma T，Marangon[A G，Stan ey D W．Environ—
Inertia【stresses affect tot~1．ato mJcrosoma 2membrane
function differently than natural ripening and senes—
维普资讯 http://www.cqvip.com
l 66 广 西 植 物 22卷
cenceEJ]．Postharvest Biol Technol，1 995，6：257
273
【34 Serek M ，Tamar】G，S[s Ler E C，et a1．Inh Lb[tion of
ethylene induced celtutar senescence symptoms by
1 methy1cycl0pr。pene，a new inhibitor of ethy[ene
action
— J]．Physiol Plan~，1995，94：229‘-232．
【35：Vo[s[ne R，Vezina I P，W [1emot C．Induction of
senescence like deterioration of microsoma1 mem
branes from cauliflower by free radicals generated
during gamma irradiation[J：． Plant Physiol，
1 991，97：545 550．
136]Votsine R．Vezina _P．W [1emot C Modifieation of
phospholipid catabo[ism in microsoma[membranes of
r irradiated cauliflower(BrQssica oleracea I )[J]
Plant Physiol，1 993，102：21 3— 21 8．
[37：Ce[ike[F G，Van Doorn W G．Solute Leakage，lipid
perox[dafion， and protein degradation during the
senescence of Iris tepa[s[J]．Physiol Plant，1 995．
94-51 5 521
．383 Mayak S，I．egge R 1 一Thmnpson J E Superox[de
radical production by mierosomal membranes from
senescing carnation flowers：an effect on membrane
fluidity[J]．Phytochem，1983，22：l耵5—1 380
[39]Baker J E，Wang C Y，Terlizzi D E．De Lay of senes—
eence in carnations by pyrazon，phenidone analogs
and tiron[J] [[ortSct，1985，2O：121 122．
64．03 Kende H Ethy Lene biosynthesis[J] J4 M Rev,Plan*
Physiol Mol Bio1．1993，44—283— 307．
[41：Lynch D V，Sridhara S，Thompson J E L[poxyge
nase generated hydroperox[des account for the non—
physio Logica L{eatures of ethylene formation from 1
aminocyc[oprppane一1-carboxy[ic acid by microsoma[
membranes of carnations J]，Ptan*a．1985，164：
1 21 l2j
(42]PMiyath G，Thompson J E Effect of membrane
phospholipid catabolism on the release of Ca ’from
m[erosoma【membrane vesicles of carnation flower
petalslJ] Plant Physiol(I i re Sei Adv)，1993，12：
l57 165
[43：I ester G．Calcium alters senescence rate of posthar
vest muskmelon fruit disks__I]．Posthar~aest Biol
Technoi，l996，7：9l 96
(443 Poovaiah BW Calcium and senescence[A]In Nood
on I D—Leopold A C．eds，Senescence and Aging in
P!ants[M3 Academic Press，San Diego，Ca[iforni
1：1988．369 389
[45]Ftuhr R，Mutton A K．Ethylene biosynthesis and
perception[J~．cr Rev Plant Sci，1 996，15-479
523．
．463 Shaw J M ，Thompson T E．Efect of phospholipid
oxidation products on transbilayer movement of
phospholtpids in single lamelar vesicles[J]
i(
．~-gen1．1981，2l 920 927
[47]Serhan C，Anderson P，GoodFl~n E，et a1．Phos
phatidate and oxidized farty acids are ca Lcium
ionophores． Studies employing arsenazn III in [ipo
somes~J一．J Bid Chem，1981，256：2 736—2 741．
q8]Gast K，Zirwer D，I adhoff A M ， a1．Auto—oxida—
tion induced fusion。f lipid vesicbs~J] Blochem B
phys Acta一1 988，686 09 109
9j Ferric B J，Beaudoin Burkhart W ，Bowsher C G，
． The cloning of twn tomato lipoxygenase genes
and their differential expression during fruit ripening
2JZ，Plan*Physiol，1 994—106：1 09-l1 8，
(5O：Fukuchl M M ．Savin K，Cornish E，et al Senes
cence induced expression 0f a homo[ogue of
DE1 TA9 desaturase in rose peta[s[J]．Plan* Mol
Bioi，]995，29：627— 635
维普资讯 http://www.cqvip.com