1-MCP Review: Effects of Ethylene Inhibitor on Fruits and Vegetables

Postharvest Biology and Technology 28 (2003) 1 /25
www.elsevier.com/locate/postharvbio
Review
1-Methylcyclopropene: a review
Sylvia M. Blankenship *, John M. Dole
Department of Horticultural Science, North Carolina State University, Box 7609, Raleigh, NC 27695-7609, USA
Received 21 June 2002; accepted 30 November 2002
Abstract
Since the discovery of 1-methylcyclopropene (1-MCP) as an inhibitor of ethylene action, over 100 studies have
examined details of its action, application and effects on ethylene inhibition. This plant growth regulator is a tool that
can help scientists make major advances in understanding the role of ethylene in plants. 1-MCP prevents ethylene
effects in a broad range of fruits, vegetables and floriculture crops. Effective concentrations are low and range from 2.5
nl l1 to 1 ml l 1. Concentration interacts with temperature such that low concentrations of 1-MCP applied over longer
durations may be as effective as high concentrations. 1-MCP is most commonly applied at 68 /77 8F (20 /25 8C), but
can be used at lower temperatures in some commodities. Generally, treatment durations of 12 /24 h were sufficient to
achieve a full response. A variety of factors may need to be considered when using 1-MCP including cultivar,
developmental stage, time from harvest to treatment, and multiple applications. Depending on the species being treated,
1-MCP may have a variety of effects on respiration, ethylene production, volatile production, chlorophyll degradation
and other color changes, protein and membrane changes, softening, disorders and diseases, acidity and sugars. This
review compiles what is known about the technological uses for 1-MCP, defines where discrepancies exist between
reports, and aims to define areas requiring further study.
# 2003 Elsevier Science B.V. All rights reserved.
Keywords: Postharvest physiology; Storage; Plant growth regulators; Senescence; Ethylene; Preservative
1. Introduction
A new tool, 1-methylcyclopropene (1-MCP), has
been added to the list of options for extending the
shelf life and quality of plant products. Not only
does commercial use of 1-MCP promise to advance commercial agriculture, but also using 1-
* Corresponding author. Tel.: /1-919-515-1224; fax: /1919-515-7747.

E-mail address:
sylvia_blankenship@ncsu.edu (S.M.
Blankenship).
MCP in research programs promises to advance

our understanding and provide new insights into
plant ethylene responses. Watkins (2002) has
summarized the effects of 1-MCP on fruit and
how it relates to ethylene physiology.
At standard temperature and pressure, 1-MCP
is a gas with a molecular weight of 54 and a
formula of C4H6. 1-MCP is thought to occupy
ethylene receptors such that ethylene cannot bind
and elicit action. Sisler and Serek (1997) proposed
a model of how 1-MCP reacts with the ethylene
receptor. The affinity of 1-MCP for the receptor is
approximately 10 times greater than that of
0925-5214/03/$ - see front matter # 2003 Elsevier Science B.V. All rights reserved.
doi:10.1016/S0925-5214(02)00246-6
S.M. Blankenship, J.M. Dole / Postharvest Biology and Technology 28 (2003) 1 /25
ethylene. Compared with ethylene, 1-MCP is

active at much lower concentrations. 1-MCP also
influences ethylene biosynthesis in some species
through feedback inhibition.
The background work for the discovery of 1MCP as an ethylene inhibitor came out of the
laboratories of Edward Sisler and Sylvia Blankenship, Departments of Biochemistry and Horticultural Science, respectively, North Carolina State
University. These workers jointly hold the patent
on the use of cyclopropenes to inhibit ethylene
action (Sisler and Blankenship, 1996). Early work
by Sisler and Blankenship on diazocyclopentadiene (DACP), a known ethylene inhibitor, led to
the hypothesis that cyclopropenes were likely to be
the light-activated product of DACP inhibiting
ethylene action; however, this was never conclusively proven. Cyclopropenes are breakdown products of DACP (Johnson, 1987). 1-MCP was
found to be a good derivative for practical use as
it is less volatile than cyclopropene itself. 3-MCP is
an effective inhibitor, but a higher concentration is
required compared with 1-MCP (Sisler et al.,
1999). For example, 40-fold higher concentrations
of 3-MCP than 1-MCP were required to effectively
prevent the damaging effects of exogenous ethylene on campanula (Campanula carpatica ) Dark
Blue and kalanchoe (Kalanchoe blossfeldiana )
(Sisler et al., 1999). Synthesis of 1-MCP can be
accomplished using the methods of Magid et al.
(1971) or Fisher and Applequist (1965).
Sisler et al. (1996a) noted that up to 15%
methylenecyclopropane (MCP) could be produced
during the production of 1-MCP and that MCP
had a completely different effect than 1-MCP:
MCP induced senescence of carnation (Dianthus
caryophyllus) (1.5 /3 nl l 1). Although up to 15%
MCP was obtained when using sodium amide to
catalyze 1-MCP production, using lithium amide
during 1-MCP production created much lower
volumes of MCP per unit volume. Because 1-MCP
(Kd 1.9 nl l 1) is effective at such low concentrations, however, these contamination levels by
MCP (Kd 22.9 nl l 1) should have little if any
measurable effect.
Other compounds such as carbon dioxide, silver
thiosulfate
(STS),
aminoethyoxyvinylglycine
(AVG), 2,5-norbornadiene (2,5-NBD) and
DACP are also effective ethylene inhibitors. Analogs of 1-MCP have been found to be effective
(Goren et al., 2001; Sisler et al., 2001). Sisler and
Serek (1999) compare inhibitory efficacy of some
of these compounds. Serek et al. (1994a) noted
that DACP and STS delayed exogenous ethyleneinduced leaf and flower bud abscission of potted
rose (Rosa ) Victory Parade. DACP and STS also
increased the display life of plants not treated with
exogenous ethylene.
2. Commercialization of 1-MCP
Commercialization of 1-MCP for ornamental
crops was first undertaken by Floralife, Inc.
(Walterboro, SC) for use on ornamental crops.
Floralife formulated a a-cyclodextrin powder that
releases 1-MCP when mixed with water (H.
Warner, personal communication). The product
was approved by the United States Environmental
Protection Agency (EPA) in 1999 for ornamentals
and is sold under the trade name EthylBloc .
Commercial application of 1-MCP to edible crops
was undertaken by AgroFresh, Inc., a subsidiary
of Rohm and Haas (Spring House, PA), under the
trade name SmartFresh . Both EthylBloc and
SmartFresh are approved for use in the United
States. In addition to the fumigant application
method, liquid spray application is being researched and may be added to the label in the
United States (Hamrick, 2001).
At standard temperature and pressure 1-MCP is
released from EthylBloc powder in approximately 20 /30 min. Complete release may take
longer at lower temperatures. The concentration
of 1-MCP gas in a sealed container with plant
material declines with time. Only about a third of
the initial amount of 1-MCP remains in the
container after 24 h at 5 8C. Up to 10% carbon
dioxide in the same container atmosphere does not
inhibit binding effectiveness of 1-MCP on apples
(Malus ). Because of the greater affinity of 1-MCP
for the ethylene receptor, approximately 100 ml l1
or greater concentration of ethylene is required to
compete effectively for the receptor in apples
(Blankenship, unpublished).
Diffusion of 1-MCP out of plant material is

rapid. No 1-MCP was detected in the core area of
individual apples approximately 8 h after application (Blankenship, unpublished). Measurable 1MCP was not detected in sealed polyethylene bags
15 days after introducing 1 ml l 1 of 1-MCP to
bananas (Musa ) (Jiang et al., 1999a). 1-MCP
appears to readily pass through plastic bags and
fiberboard boxes. 1-MCP analysis can be conducted using gas chromatography with isobutylene
often being used as a standard. Glass Porapak Q
columns work well to analyze both 1-MCP and
ethylene while alumina columns will give an
inactivated form of 1-MCP.
The safety, toxicity and environmental profiles
of 1-MCP in regard to humans, animals and the
environment are extremely favorable (Environmental Protection Agency, 2002). The compound
is used at low rates, has a non-toxic mode of action
and is chemically similar to naturally occurring
substances. The rat inhalation LC50 is greater than
2.5 mg L (or 1.126 ppm v/v active ingredient in
air). In tests for acute toxicity of 1-MCP no death
or clinical signs of systemic toxicology were seen
(Environmental Protection Agency, 2002).
3. Conditions for use

3.1. Temperature
In most studies 1-MCP has been applied at
temperatures ranging from 68 to 77 8F (20 /
25 8C) (Tables 1 and 2). Lower temperatures
have been used but a relationship exists between
concentration, time and temperature, and applications at low temperatures are not effective on some
crops. In coriander (Coriandrum sativum ) 1-MCP
was not effective at low temperatures (5 /10 8C),
presumably due to low sensitivity to ethylene
(Jiang et al., 2002a). Application of 1-MCP (5 or
20 nl l1) at 36 8F (2 8C) was not effective on
penstemon (Penstemon) Firebird; however, application at 68 8F (20 8C) resulted in complete
protection from exogenous ethylene (Serek et al.,
1995a). In broccoli (Brassica oleracea) 1-MCP
application produced better results at 68 8F
(20 8C) than at 41 8F (5 8C) but an effect
occurred at both temperatures (Ku and Wills,

1999). Able et al. (2002a) also found that 68 8F
(20 8C) was best for broccoli treatment. In apples
a given concentration of 1-MCP had less effect on
firmness as storage temperature was lowered (Mir
et al., 2001) and it was hypothesized that lower
temperatures might lower the affinity of the
binding site for 1-MCP. A relationship was noted
between treatment time and temperature; apples at
37 8F (3 8C) required 9 h treatment, whereas only
6 h was needed at higher temperatures to delay
ripening (DeEll et al., 2002).
3.2. Active concentrations
Effective concentrations of 1-MCP vary widely
and with commodity; similarly, they vary with
respect to time, temperature and method of
application (Tables 1 and 2). Inconsistencies in
EthylBloc powder that occurred during the early
years of production may account for some discrepancies. The minimum concentration required
was 2.5 nl l1 on carnation, while in apples some
studies show 1 ml l 1 was required to block
ethylene action (Jiang and Joyce, 2002; Sisler et
al., 1996a; Fan et al., 1999a). Concentrations of
between 1 (Fan and Mattheis, 2000a; Ku and
Wills, 1999) and 12 ml l 1 (Able et al., 2002a) were
effective in blocking ethylene action in broccoli. In
tomato (Lycopersicon esculentum ) 7 nl l1 of 1MCP blocked the green to red color change for 8
days (Sisler et al., 1996b). Higher concentrations
of 1-MCP (0.1 /100 ml l 1) for short durations (2
h) were effective on green tomatoes (Wills and Ku,
2002), with at least 20 ml l 1 required to increase
postharvest life of ripe tomatoes. While 5 and 50 nl
l1 had no effect on unripe bananas, 500 nl l1
delayed ripening (Harris et al., 2000), which is in
contrast to Sisler et al. (1996b) who reported that
only 0.7 nl l 1 was effective. In apples 1 ml l 1 1MCP was effective in delaying ripening (Fan et al.,
1999a; Rupasinghe et al., 2000b). Watkins et al.
(2000) found that the response of McIntosh and
Law Rome apples to 1-MCP was more dependent on concentration than the response of Delicious or Empire. A 1-MCP concentration of 100
nl l 1 inhibited ethylene by 50% in Anna apples,
but did not inhibit volatile production, while 1 ml
Table 1
Summary of 1-MCP treatment conditions and effects on fruits and vegetables
Concentration
Treatment
temperature
(8C)
Treatment
time
Effects
Reference
Ananas comosus (pineapple)
0.1 ml l 1
20
18 h
Selvarajah et al., 2001
Annona squamosa /Annona cheimola (custard apple)

Arabidopsis
25 ml l 1
20
14 h
Controlled internal browning; slowed ascorbic acid, ethylene, and

soluble solids declines
Increased decay, delayed ripening
2.2 nl l 1
Hall et al., 2000
Brassica oleracea (broccoli)
1, 12 ml l 1
5, 10, 20
6, 12, 16 h
B. rapa var. chinensis (pak choy)
12 ml l 1
10
16 h
B. juncea var. foliosa (Chinese mus- 12 ml l 1

tard)
10
16 h
12 ml l 1
10
16 h
B. rapa var. nipposinica (mibuna and 12 ml l 1

mizuna)
B. rapa var. rosularis (tatsoi)
12 ml l 1
10
16 h
Allowed normal seedling growth

with exogenous ethylene present
Delayed yellowing, decreased respiration, decay, extended shelf life
Increased shelf life if exposed to
exogenous ethylene, reduced yellowing
Delayed yellowing, increased shelf
life if exposed to exogenous ethylene
life if exposed to exogenous ethylene
Increased shelf life
10
16 h
Able et al., in press
Carica papaya (papaya)
25 ml l 1
20
14 h
Chrysanthemum coronarium (garland chrysanthemum)

Coriandrum sativum (coriander)
12 ml l 1
10
16 h
50 nl l 1
20
24 h
Cucumis melo (melon) plants
0.1 ml l 1
10 min
Citrullus lanatus (watermelon)
5 ml l 1
20
18 h

life
Increased blemishes slightly, delayed ripening
life if exogenous ethylene
Chlorophyll and protein maintained, senescence delayed
Inhibited ethylene pathway transcripts, wounding response not effected
Reduced internal breakdown
Citrus paradisi (grapefruit)
25
16 h, twice
Citrus spp. (orange)
100 nl l 1 (fruit); 50 ml
l 1 (petioles)
25
6, 12 h
B. rapa var. parachinensis (choy

sum)
Hofman et al., 2001
Fan and Mattheis, 2000; Ku and

Wills, 1999; Able et al., 2002a
Able et al., 2002a
Hofman et al., 2001

Jiang et al., 2002b
Bouquin et al., 1997
Mao and Huber (personal communication)

Prevented degreening and ethylene Mullins et al., 2000
production
Increased decay and chilling injury, Porat et al., 1999; Sisler et al., 1999
blocked ethylene induced degreening, inhibited abscission
Plant
Table 1 (Continued )
Plant
Concentration
Treatment
temperature
(8C)
Treatment
time
1 ml l 1
20
4h
Effects
Nicotiana attenuata (tobacco) plants 0.04 /0.43 g of 0.43%

Growing tem- 6 h
EthylBloc in 27/40/ perature
22.5 cm chamber; 500
mg in a 18.5 L chamber
Fan and Mattheis, 2000
Nakano et al., 2001, 2002

Ku et al., 1999; Jiang et al., 2001;
Tian et al., 2000
Pereira-Netto, 2001
Fan and Mattheis, 2000; Wills et al.,
2002
Sisler et al., 1996b; Nakatsuka et al.,
1997, 1998; Hoeberichts et al., 2002;
Wills and Ku, 2002
Ciardi et al., 2001
Watkins et al., 2000; DeEll et al.,

2002; Baritelle et al., 2001; Fan and
Mattheis, 1999a,b; Fan et al.,
1999a,b; Fan and Mattheis, 2001;
Jiang and Joyce, 2002; Rupasinghe
et al., 2000a,b, 2001; Lurie et al.,
2002; Mir et al., 2001
Hofman et al., 2001; Jiang and
Joyce, 2000
Harris et al., 2000; Jiang et al.,
1999a,b; Golding et al., 1998; Sisler
et al., 1999; Serek et al., 1995b; Sisler
and Serek, 1999
Prevented ethylene response to in- Winz and Baldwin, 2001; Kahl et al.,
jury and nicotine response allowing 2000
insects to feed on plant
Prevented acidity loss, respiration

increase, and inhibited isocourmarin increase with exogenous ethylene
Diospyros kaki
300 nl l 1
20
3h
Inhibits fruit softening
Fragaria /ananassa (strawberry)
5 /15; 250 /500 nl l 1; 2 20
2 /18 h
Increased shelf life, maintained
ml l 1
firmness and color, lowered phenolic content, increased decay at
high 1-MCP concentrations, lowered ethylene production, effects on
respiration vary with maturity
Hanconia speciosa
90 ml l 1
31
Continuous Induced lateral shoot development
of in vitro-grown shoots
Lactuca sativa (lettuce)
1; 0.1 ml l 1
6
4h
Inhibited ethylene induced russet
spotting, postharvest life increased
Lycopersicon esculentum fruits (to- 5 /7; 10 /20, 150 nl l 1; 20
2 /24 h
Postharvest life increased, respiramato)
20 ml l 1
tion rate reduced, titratable acidity
loss reduced, ACC synthase and
ACC oxidase transcripts reduced,
delayed ethylene production
L. esculentum plants (tomato)
4 mg EthylBloc powder Growing tem- Overnight
Disease response as regulated by
per L
perature
ethylene sensitivity
0.6 /2 ml l 1
Malus domestica (apple) Anna,
0, 5, 10, 15,
Maintained firmness, reduced
7 /20 h
Fuji, Golden Delicious, Red De20 /25
scald, maintained titratable acidity,
licious, Rome, Gala, McIntosh,
reduced volatiles, slowed loss of
Granny Smith, Ginger Gold, Jochlorophyll and starch, inhibited
nagold, Empire, Law Rome
ethylene, reduced respiration and
decay, prevented/delayed disorders
such as scald
Mangifera indica (mango)
25; 100 ml l 1
20
14 h
Increased shelf life, increased stem
rots
Musa sp. , AAA group, Cavendish 5 /500 nl l 1; 0.1 ml l 1, 20 /24
6 /24 h
Delayed ripening and peel color
subgroup (banana)
45
change
Daucus carota (carrot)
Reference
Concentration
Treatment
temperature
(8C)
Treatment
time
Effects
Reference
Persea americana (avocado)
50 /300 nl l 1; 0.45, 25

ml l 1
3, 5, 20, 22
6 /48 h
Jeong et al., 2001; Hofman et al.,

2001; Pesis et al., 2001; Feng et al.,
2000
Pisum sativa seedlings (pea)
40 nl l 1
24 h
Inhibited ripening, maintained

firmness, reduced cell wall degrading enzymes, delayed and reduced
respiration, ethylene production
and weight loss; retained green peel
color, reduced mesocarp discoloration and decay development
Inhibited growth
Prunus armeniaca (apricot)
1 ml l 1
20
4, 20 h
P. persica (peach and nectarine)
20 nl l 1; 0.1 ml l 1; 0.5 20 /24

ml l 1
4, 18, 24 h
P. salicina (plum)
1, 13, 26 or 39 ml l 1
20
6, 20, 24 h
Pyrus communis (pear)
2, 4 ml l 1
16 h
Vigna radiata leaves (mung bean)
60 nl l 1
Ambient
24 h
Sisler and Serek, 1997; Sisler et al.,

1999
Maintained firmness and titratable Dong et al., 2002; Fan et al., 2000a
acidity, decreased internal flesh
browning and decay, inhibited
ethylene production in some conditions, reduced respiration, delayed volatile production
Maintained firmness and acidity;
Fan et al., 2002; Dong et al., 2001a;
delayed ethylene production and
Mathooko et al., 2001
reduced respiration; increased internal browning; reduced juice,
ethylene associated enzymes and
cell wall degrading enzymes
Reduced ethylene and respiration, Dong et al., 2001b, 2002; Abdi et al.,
maintained firmness, slowed color 1998
change, decreased internal flesh
browning
Reduced chilling induced ethylene, Lelievre et al., 1997; Baritelle et al.,
slowed softening
2001
Inhibited abscission
Sisler et al., 1999
Plant
Table 2
Summary of 1-MCP treatment conditions and effects on floricultural crops
Concentration
Treatment temperature (8C)
Treatment
time (h)
Effects
Reference
Alstroemeria spp.
20 nl l 1
20
Antirrhinum majus
20 nl l 1
20
Begonia /elatior Najada and

Rosa
5 or 20 nl l 1
20
Serek et al.,
1995a
Serek et al.,
1995a
Serek et al.,
1994b, 1995b
Begonia /tuberhybrida Non-Stop 5 or 20 nl l 1
20
Prevented decrease in vase life due to exogenous ethylene

(1 ml l 1)
Prevented decrease in vase life due to exogenous ethylene
(1 ml l 1)
Delayed exogenous ethylene-induced (1.0 ml l 1) abscission of buds, flowers, and leaves, and senescence and
increased postharvest life in a non-ethylene environment
Prevented exogenous ethylene-induced (10 ml l 1) fresh
weight loss and flower abscission
Increased individual flower longevity in the presence of
exogenous (3 ml l 1) ethylene and in a non-ethylene
environment
In the presence of exogenous ethylene (0.5 ml l 1),
increased display life and flower longevity. Without
exogenous ethylene, 1-MCP had no effect on Campanula
Dark Blue or on display life of Campanula Blue Clips
but did increase flower longevity of Campanula Blue
Clips. STS increased plant display life and flower
longevity greater than 1-MCP for both species regardless
of the presence of ethylene
Increased vase life in a non-ethylene environment but
only in combination with 5% sucrose
Neither 1-MCP or exogenous ethylene (10 ml l 1)
application affected senescence
Prevented effects of exogenous ethylene (10 ml l 1)
application including flower abscission and reduced
vaselife and increased vaselife without the presence of
exogenous ethylene
application including flower abscission and reduced vase
life but did not affect normal senescence in a nonethylene environment
Prevented the effects of exogenous ethylene (2 ml l 1)
application including bud and flower abscission. Partially prevented bud, flower, and leaf abscission from
dehydration and storage
Boronia heterophylla
10 nl l 1
20
12
Campanula carpatica Dark Blue
20, 50 or 100 nl
l 1
20
C. carpatica Dark Blue and Blue

Clips
20, 50 or 100 nl
l 1
21
C. medium Champion Pink
800 nl l 1
22
Cassinia adunca
10 nl l 1
20
12
Ceratopetalum gummiferum
10 nl l 1
20
12
Chamelaucium uncinatum
10 nl l 1
20
12
C. uncinatum
200 nl l 1
21
6 or 13
Serek et al.,
1994b, 1995b
Macnish et
al., 1999
Sisler et al.,
1999
Serek and
Sisler, 2001
Bosma and
Dole, 2002
Macnish et
al., 2000
Macnish et
al., 2000
Macnish et
al., 2000
Plant
Serek et al.,
1995c
Plant
Concentration
Treatment
time (h)
Effects
Reference
Consolido ambigua
20 nl l 1
20
Dendranthema grandiflorum Coral

Charm
Dianthus barbatus
200 nl l 1
20
20 nl l 1
20
D. caryophyllus Sandra
0.6, 1.7, 3.3, 5.8 or 20

20 nl l 1
Serek et
1995a
Serek et
1998
Serek et
1995a
Serek et
1995a,b
D. caryophyllus White Sim or

Sandra
5 nl l 1
Not specified, prob- 12

ably room temperature
Epipremnum pinnatum
200 nl l 1
20
Eriostemon scabe
10 nl l 1
20
12
Grevillea Kay Williams
10 nl l 1
20
12
Grevillea Misty Pink
10 nl l 1
20
12
Gypsophila paniculata Perfecta,

Gilboa, and Golan
200 nl l 1
20
24
Hibiscus rosa-sinensis numerous cul- 200 nl l 1

tivars
H. rosa-sinensis
200 nl l 1
20
20
Ixora coccinea Big Red
100 nl l 1
20
Kalanchoe blossfeldiana Tropicana
5 or 20 nl l 1
20
20
Prevented vase life decrease due to exogenous ethylene (1

ml l 1)
Reduced rooting of both stored and unstored unrooted
cuttings
ml l 1)
Prevented exogenous ethylene (0.4 ml l 1) induced
senescence and increase vase life even without exogenous
ethylene application
Prevented exogenous ethylene (1.0 ml l 1) induced
senescence and increase vase life even without exogenous
ethylene application. Older flowers required higher
concentrations to prevent damage
Decreased leaf yellowing and drop of cuttings after
storage of cuttings at 23 8C for 4 days
Delayed senescence of open flowers, but not floral buds,
exposed to ethylene (0.7 ml l 1) while STS delayed
senescence of both open flowers and unopen buds
Had no effect on open flowers unless flowers were held in
1-MCP continuously in a non-ethylene environment
Retarded storage-induced leaf yellowing of unrooted
cuttings but reduced subsequent rooting
Blocked chilling induced leaf abscission in both an
exogenous ethylene (5 ml l 1) and a non-ethylene
environment
Had no effect on postharvest life
al.,
al.,
al.,
Sisler et al.,
1996
Muller et al.,
1997
Macnish et
al., 2000
Macnish et
al., 2000
Macnish et
al., 2000
Newman et
al., 1998
Reid et al.,
2002
Serek et al.,
1998
Michaeli et
al., 1999
Serek et al.,
1994b, 1995b
Serek and
Reid, 2000
Kalanchoe Alexandria, Debbie,

200 nl l 1
Caroline, Jaqueline, Nadia, Pale
Jaqueline, and Simone
al.,
Plant
Concentration
K. blossfeldiana
Effects
Reference
Leptospermum petersonii
0.5, 2.5, 5 or 10 nl 20
l 1
10 nl l 1
20
Sisler et al.,
1999
Macnish et
al., 2000
L. scoparium
10 nl l 1
20
12
Lilium Mona Lisa
500 nl l 1
25
18
Lilium Stargazer
500 nl l 1
25
18
Lilium hybrids Cordelia and Elite 150 nl l 1

(Asiatic types)
20
L. longiflorum Lorena
150 nl l 1
20
Lupinus havardii Texas Sapphire
450 nl l 1
15
12
Increased post production life in the presence of

exogenous (3 ml l 1) ethylene
Prevented effects of exogenous ethylene (2 or 5 ml l 1)
application including rapid bud and flower abscission
but did not effect normal senescence in a non-ethylene
environment
Prevented effects of exogenous ethylene (2 or 5 ml l 1)
application including rapid bud and flower abscission
but did not effect normal senescence in a non-ethylene
environment
Prevented exogenous ethylene (10 ml l 1) induced vase
life decline but had no effect on vase life, days to bud
opening, or percentage of buds opening in a nonethylene environment
Prevented exogenous ethylene (10 ml l 1) induced vase
life decline but had no effect on vase life, days to bud
opening, or percentage of buds opening in a nonethylene environment
Reduced fresh weight loss and reduced flower abscission
Matthida incana
20 nl l 1
20
M. incana
500 nl l 1
20
Metrosideros collina
0, 1.5, 15, or 150

nl l 1
20
Ozothamnus diosmifolius
10 nl l 1
20
12
Pelargonium /hortorum
0.1 or 1.0 ml l 1
20
3, 6, 12, or
24
12
Macnish et
al., 2000
Celikel et al.,
2002
Celikel et al.,
2002
Elgar et al.,
1999
Elgar et al.,
1999
Picchioni et
al., 2002
Prevented vase life decrease due to exogenous ethylene (1 Serek et al.,
ml l 1)
1995a
Delayed exogenous ethylene-induced (1 ml l 1) abscis- Celikel and
sion of flowers and increased fresh weight and vase life in Reid, 2002
a non-ethylene environment. STS induced longer vase
life and greater fresh weight retention than 1-MCP
Sun et al.,
Delayed exogenous ethylene-induced (0.1 ml l 1) ab2000
scission of stamens by 1 /2 days. STS delayed stamen
abscission by 4 or more days
Macnish et
al., 2000
Delayed exogenous ethylene-induced (1.0 ml l 1) petal Jones et al.,
abscission and prevented petal abscission in a non2001
ethylene environment
Treatment
time (h)
10
Concentration
Treatment
time (h)
Effects
Reference
P. zonale Isabel
200 nl l 1
20
Serek et al.,
1998
P. peltatum Pink Blizzard
1 ml l 1
22
Penstemon Firebird
20 nl l 1
20
Petunia hybrida Pink Cascade
150 nl l 1
22
Phalaenopsis Herbert Hager
250 nl l 1
22
Phlox paniculata Rembrandt
25, 250, or 500 nl 229/1

l 1
Platysace lanceolata
10 nl l 1
20
12
Rosa Royal and Sunset
100 nl l 1
20
Rosa Vanilla and Bronze
200 nl l 1
20
R. hybrida Victory Parade
5 or 20 nl l 1
20
21
Retarded storage-induced leaf yellowing and did not

effect rooting of both stored and unstored unrooted
cuttings
Delayed exogenous ethylene-induced (1.5 ml l 1) abscission of petals but the effect diminished and the halflife of the treatment was 2, 3 and 6 days at 25, 20.7 and
12 8C, respectively
ml l 1)
Prevented an exogenous ethylene-induced (1 /12 ml l 1)
increase in electrolytic leakage, a decrease of membrane
proteins, and a decrease in lipid fluidity. In a nonethylene environment increased individual flower longevity, fresh weight, and total protein content compared
with untreated controls but had no effect on electrolytic
leakage, membrane proteins, or lipid fluidity
Prevented the pollination-induced increase in ethylene
production and the enhanced senescence of the flowers
Prevented exogenous ethylene (0.3, 1 or 3 ml l 1) induced
flower abscission and vase life decline but had no effect
on abscission, number of open flowers, or vase life in a
non-ethylene environment
Delayed exogenous ethylene-induced (0.5 ml l 1) abscission of leaves, buds, and flowers, but did not effect
leaf yellowing of plants placed in dark
Delayed but did not prevent ABA (0.1 mM) induced
flower senescence in Bronze and had no effect on flower
abscission in Vanilla. Did not have any effect on ABA
induced leaf abscission
Increased display life, with or without the presence of
exogenous ethylene (0.5 ml l 1), of Schlumbergera ; STS
increased plant display life and flower longevity greater
than 1-MCP regardless of the presence of ethylene
Schlumbergera truncata Dark Mar- 20, 50 or 100 nl

ie
l 1
Cameron and
Reid, 2001
Serek et al.,
1995a
Serek et al.,
1995d
Porat et al.,
1995a
Porat et al.,
1995a
Macnish et
al., 2000
Serek et al.,
1996
Muller et al.,
1999
Serek et al.,
1994b, 1995b
Serek and
Sisler, 2001
Plant
Macnish et
al., 2000
Macnish et
al., 2000
De Wild et
al., 2002
Macnish et
al., 2000
Macnish et
al., 2000
20
10 nl l 1
Zieria cytisoides
12
20
10 nl l 1
Verticordia nitens
12
20
1 nl l 1
Tulipa gesneriana Apeldoorn
16
20
10 nl l 1
Thryptomene calycina
12
Did not prevent the effects of exogenous ethylene (10 ml

l 1) application including flower abscission and reduced
vase life and did not affect normal senescence in a nonethylene environment
Prevented exogenous ethylene (0.3 Pa) induced gummosis of stored bulbs
20
10 nl l 1
Telopea Shady Lady
12

Concentration
Plant
Treatment
time (h)
Effects
Reference
11
l1 produced 70% volatile inhibition and 95%

ethylene inhibition (Lurie et al., 2002). Jiang and
Joyce (2002) found that while 1 ml l 1 was
sufficient to produce a decrease in ethylene, 10 ml
l1 further reduced ethylene production in intact
and cut apples. Application was especially effective
if used prior to coring and peeling of apples. A
concentration of 0.09 ml l1for 24 h was not
enough to elicit a response in avocado (Persea
americana ) softening while 0.45 ml l1 for 24 h
affected softening and associated enzyme activity
(Jeong et al., 2002). Feng et al. (2000) found that
30 nl l 1 or higher was sufficient to delay ripening
in avocado and Pesis et al. (2002) used 300 nl l1
to reduce mesocarp discoloration.
Serek et al. (1995a,b) tested a range of 1-MCP
concentrations from 0.6 to 20 nl l 1 on cut
carnation Sandra flowers and noted that 90% of
possible carnation longevity was obtained with a
10/20 nl l 1 concentration range. Similarly, 20 nl
l1 was optimum when compared with 1 or 5 nl
l1 on cut snapdragon (Antirrhinum majus ) and
penstemon Firebird stems. Sisler et al. (1996a)
tested concentrations of 1.25, 2.5, 5, and 10 nl l1
for 24 h in young carnation flowers (Stage II) and
noted that at least 2.5 nl l 1 was required for
prevent effects of exogenous ethylene while 5 nl
l1 was required in older flowers (Stage III). Low
concentrations of 1-MCP may be as effective as
higher concentrations if the low concentrations are
applied over longer durations. Sisler et al. (1996a)
noted that 250 /300 nl l 1 1-MCP for 5 min were
as effective on carnation as 0.5 nl l 1 for 24 h.
With potted plants, increasing the concentration
of 1-MCP from 0 to 5.8 nl l 1 [heimalis begonia
(Begonia /elatior hybrida) Najada] or to 10 nl
l1 (kalanchoe Tropicana and rose Victory
Parade) decreased the percentage of begonia bud
and flower drop, senesced kalanchoe flowers or
rose leaf drop (Serek et al., 1994b, 1995b). Minimum concentration varied with species. Increasing
1-MCP concentration from 20 to 100 nl l1
increased display life of holiday cactus (Schlumbergera ); however, only 20 nl l 1 was required to
obtain maximum display life of campanula in the
presence of exogenous ethylene (Serek and Sisler,
2001). Increasing 1-MCP concentration from 0 to
10 nl l 1 increased postproduction life of kalan-
12
choe in the presence of exogenous (3 ml l 1)

ethylene, but the maximum effective concentration
was not determined (Sisler et al., 1999). 1-MCP
will reverse the ethylene-mediated triple response
in Arabidopis seedlings, with the first reduction in
response documented at 2.2 nl l 1 and full effects
at 220 nl l 1 (Hall et al., 2000).
the Redchief strain of Delicious apples advancing

maturity slightly decreased the effect of 1-MCP
(Mir et al., 2001). Arabidopsis wild-type seedlings
grown in an atmosphere of air are much like ones
grown in an atmosphere containing 1-MCP (Hall et
al., 2000) indicating that the suppression of the
ethylene response by 1-MCP will not drastically
alter growth.
3.3. Treatment duration

3.5. Time from harvest to treatment
In most studies treatment duration ranged from
12 to 24 h, which was sufficient to achieve a full
response (Tables 1 and 2). An exposure of 6 h at 0.45
ul l 1 was not enough to induce respiratory or
ethylene production changes in avocado (Jeong et
al., 2002). A time/temperature relationship was
noted with broccoli (Ku and Wills, 1999) and
banana (Jiang et al., 1999b) such that higher
concentrations of 1-MCP were required for shorter
treatment times. Cultivar should also be considered: Empire apples required less treatment time
than Cortland to achieve the same effect at the
same 1-MCP concentration (DeEll et al., 2002).
3.4. Developmental stage and plant maturity
Plant developmental stage must be considered
when applying 1-MCP as effects vary with plant
maturity. 1-MCP effects in apricots (Prunus armeniaca ) decreased with advanced fruit development
(Fan et al., 2000a). A leafy brassica was found to
respond less to 1-MCP than a floral brassica,
possibly due to different ages of leaves and inherent
differences between flowers and leaves (Able et al.,
2002a). Single flowers (i.e. carnations) may be
completely protected from ethylene, but spike or
spray flowers [i.e. delphinium (Delphinium ) and
gypsophila (Gypsophilia paniculata )] may be only
partially protected, presumably due to different
tissue ages (Sisler et al., 1996a; Newman et al.,
1998). Banana maturity was a major factor in the
response of the fruit to 1-MCP (Harris et al., 2000).
Because commercial lots of bananas can have a
range of maturity, 1-MCP treatment would likely
result in mixed lots of fruit. Stage of ripeness in
pears was found to be significant in 1-MCP
treatment effects when testing pears for tissue
mechanical properties (Baritelle et al., 2001). In
The importance of time from harvest to 1-MCP

treatment varies with the crop species; generally
the more perishable the crop, the more quickly
after harvest 1-MCP should be applied. 1-MCP
must be applied as soon as possible to broccoli and
pak choy (Brassica rapa ) (Able et al., 2002a). In
bananas treated with ethylene, fruit had to be
treated with 1-MCP within 24 h to delay ripening
(Jiang et al., 1999b). Ethylene production, softening, and internal browning in ripening apricots
and plums was inhibited when fruit were treated
with 1-MCP after storage, but not before storage
(Dong et al., 2002). A similar study was done on
kiwifruit (Actinidia deliciosa ) with 1-MCP treatment before or after storage (Kim et al., 2001).
3.6. Multiple applications
Two applications of 100 nl l1 were found to be
more beneficial in reducing ethylene-induced mesocarp discoloration in avocado than just one
application (Pesis et al., 2002). Multiple applications of 1-MCP during storage of Redchief
apples were more effective at higher temperatures
than at 32 8F (0 8C) (Mir et al., 2001). However,
multiple applications of 1-MCP to broccoli and
pak choy had no more experimental effect than
one application (Able et al., 2002a).
4. Protection from endogenous vs. exogenous

ethylene
1-MCP will protect plant products from both
endogenous and exogenous sources of ethylene.
Studies of 1-MCP have been conducted in the
presence of applied ethylene and/or in the presence
of endogenous ethylene. Responses to the two

types of tests vary by commodity. Some crops will
benefit from 1-MCP regardless of the presence of
exogenous ethylene. Other crops show little benefit
from 1-MCP application unless exogenous ethylene is present.
If exogenous ethylene was present, 1-MCP
protected both broccoli and pak choy from detrimental ethylene effects. In contrast, in the absence
of exogenous ethylene, 1-MCP had little effect in
extending shelf life of pak choy but did extend the
shelf life of broccoli by 20% (Able et al., 2002a).
Water soaking of tissue that occurs when exogenous ethylene is applied to watermelon (Citrillus
lanatus) is prevented by 1-MCP application even
though watermelon produces little endogenous
ethylene (Mao and Huber, personal communication). In six leafy Asian vegetables, 1-MCP delayed senescence as measured by leaf yellowing
and browning margins in the presence of exogenous ethylene; however, 1-MCP increased shelf life
of only two types in the absence of exogenous
ethylene (Able et al., in press). When strawberry
fruits were treated with 1-MCP and then exposed
to an exogenous application of ethylene, the effects
were dependent on fruit maturity and 1-MCP
concentration (Ku et al., 1999; Tian et al., 2000).
However, 1-MCP was also found to extend
strawberry postharvest life in the absence of
exogenous ethylene (Jiang et al., 2001). 1-MCP
delayed softening in mango (Mangifera indica)
when the fruit were treated with ethephon (Jiang
and Joyce, 2000).
1-MCP prevented damage from exogenous ethylene in numerous cut flower species. This included prevention of rapid desiccation and flower
abscission of stock (Celikel and Reid, 2002); rapid
bud and flower abscission of oriental lily (Lilium )
Mona Lisa and Stargazer (Celikel et al., 2002);
vase life decline of asiatic lilies (Lilium ) Cordelia
and Elite and Easter lily (Lilium longiflorum )
Lorena (Elgar et al., 1999); flower abscission and
reduced vase life of several Australian native cut
flowers (Macnish et al., 2000); fresh weight loss
and flower abscission of boronia (Boronia heterophylla) (Macnish et al., 1999); senescence of open
gypsophila flowers (Newman et al., 1998); flower
abscission and reduced vase life of garden phlox
13
(Phlox paniculata ) Rembrandt (Porat et al.,

1995b), flower or petal abscission and floret
wilting of carnation Sandra, alstroemeria (Alstroemeria ), snapdragon, larkspur (Consolida ambigua ), sweet william (Dianthus barbatus ), stock
(Matthiola incana ), and penstemon Firebird
(Serek et al., 1995a); bud and flower abscission
of geraldton wax (Chamelaucium uncinatum )
(Serek et al., 1995c); stamen abscission of metrosideros (Metrosideros collina ) (Sun et al., 2000);
and enhanced the longevity and fresh flower
weight in petunia (Petunia hybrida ) Pink Cascade
(Serek et al., 1995d).
Similarly, 1-MCP prevented damage from exogenous ethylene on many potted plants species
including prevention of flower abscission on
heimalis begonia Najada and Rosa and tuberous begonia (Begonia /tuberhybrida ) NonStop, senescence of kalanchoe Tropicana and
bud and leaf abscission of rose Victory Parade
(Serek et al., 1994b, 1995b). While 1-MCP and
STS delayed effects of ethylene-induced senescence
in heimalis begonia Rosa, kalanchoe Tropicana
and rose Victory Parade by 5/12 days compared
with untreated plants, plants not receiving ethylene lasted 26/35 days, indicating that 1-MCP and
STS did not completely eliminate the effect of
exogenous ethylene (Serek et al., 1994b, 1995b). In
addition, only rose showed an increase in display
life when treated with either 1-MCP or STS
compared with untreated plants. 1-MCP also
prevented petal abscission of ivy geranium (Pelargonium peltatum ) Pink Blizzard (Cameron and
Reid, 2001); petal abscission of zonal geranium
(Pelargonium /hortorum ) (Jones et al., 2001); leaf
abscission and chlorosis of croton (Codiaeum
variegatum var. pictum Aucubaefolia) (Muller
et al., 1998); leaf, bud, and flower abscission of
rose Royal and Sunset (Serek et al., 1996),
reduction in display life and flower longevity of
campanula Dark Blue and Blue Clips and
holiday cactus Dark Marie (Serek and Sisler,
2001; Sisler et al., 1999) and reduction of post
production life of kalanchoe (Sisler et al., 1999).
In some cases 1-MCP prevented some of the
damage resulting from exogenous ethylene when
the two compounds were applied together but
completely prevented ethylene damage when the 1-
14
MCP was applied prior to the ethylene. This

indicates that 1-MCP ideally should be applied
prior to any possible ethylene exposures (Serek et
al., 1995c). In addition to preventing ethylene
damage, 1-MCP was also partially effective in
reducing the bud, flower, and leaf abscission that
occurs from dehydration during storage of cut
geraldton wax stems (Serek et al., 1995c). Stems
treated with 1-MCP had 36, 47 and 48% less
abscission of bud, flower, and leaves than untreated stems.
With most cut and potted flower species 1-MCP
had no effect on normal senescence in the absence
of exogenous ethylene (Celikel et al., 2002; Elgar et
al., 1999; Macnish et al., 2000; Porat et al., 1995b;
Serek and Reid, 2000; Serek and Sisler, 2001;
Serek et al., 1994b, 1995b). With hibiscus (Hibiscus
rosa-sinensis ) 1-MCP was only effective in extending the life of open flowers when they were held in
1-MCP continuously. However, vase life was
increased with the application of 1-MCP in a
small number of cut flower species without exposure to exogenous ethylene including Canterbury bells (Campanula medium ) Champion
(Bosma and Dole, 2002), stock (Celikel and
Reid, 2002), ceratopetalum (Ceratopetalum gummiferum ) (Macnish et al., 2000), petunia Pink
Cascade (Serek et al., 1995d), and carnation
(Serek et al., 1995a,b). In addition, 1-MCP increased the postharvest life of several potted plants
including rose Victory Parade (Serek et al.,
1994b, 1995b) and campanula Dark Blue (Sisler
et al., 1999). With zonal geraniums 1-MCP decreased petal abscission (Jones et al., 2001). 1MCP had no effect on senescence of some cut
flower species regardless of the presence of exogenous ethylene. This included several Australian
native cut flowers (Macnish et al., 2000). 1-MCP
also did not prevent the effects of exogenous
ethylene application on flower abscission and
reduced vase life on telopea (Telopea ) Shady
Lady (Macnish et al., 2000).
5. Regeneration of sites or sites available

The assumption has been made that 1-MCP
binds permanently to receptors present at the time
of treatment and any return of ethylene sensitivity

is due to appearance of new sites. While this may
be true, there is little direct supporting data. Plant
tissues vary greatly in their ability to regenerate
sites. In avocado, ripening was delayed by about 2
weeks after 1-MCP treatment, then fruit ripened
normally (Feng et al., 2000). Pesis et al. (2002)
found that two applications of 300 nl l 1 made 10
days apart, was enough to prevent softening in
avocado, whereas one application would allow
fruit to soften normally. Binding sites appear to
develop after treatment of broccoli as continuous
application is more effective than a one-time
treatment, but this was not true in pak choy
(Able et al., 2002a). Ripening is delayed in
tomatoes by 5/10 days (Hoeberichts et al., 2002;
Wills and Ku, 2002; Sisler et al., 1996b) with one
application of 1-MCP but fruit must be retreated
for continued effect.
With bananas, factors such as cultivar, maturity
of fruit, prior exposure to ethylene, or growing
conditions appear to greatly influence the results
of using 1-MCP and the ability of the fruit to
eventually ripen. Bananas treated with 1-MCP and
kept in polyethylene bags eventually ripen, indicating that bananas might make new receptors in
this situation (Jiang et al., 1999a). However,
bananas treated with 1-MCP may also stay green
or will ripen with an uneven color (Harris et al.,
2000). Lack of ripening probably indicates that
receptors were not regenerated. It has been suggested that heat treatment in bananas will enhance
synthesis of ethylene receptors (Jiang et al., 2002a).
Bananas attached to the plant and treated with 1MCP eventually ripened normally, while harvested
treated fruit did not ripen properly (Blankenship,
unpublished data). The role of abscisic acid (ABA)
in ripening bananas appears to be mediated
through ethylene, as 1-MCP-treated bananas do
not show the stimulation of ripening caused by
ABA (Jiang et al., 2000). Jiang et al. (1999b)
suggested that 1-MCP exhibits non-competitive
binding with the receptor in banana. Sisler et al.
(1996b) stated that 1-MCP is bound on ethylene
receptors but probably competes with endogenous
ethylene for binding sites. The inconsistencies in
banana ripening after 1-MCP treatment, possibly
due to receptor site availability, may limit com-
mercial use of 1-MCP in the banana industry

unless suitable protocols are devised.
Partial responses in some apples cultivars either
seem to suggest these cultivars are able to regenerate sites or that binding is incomplete. McIntosh apples, for instance, may need higher 1-MCP
concentrations, perhaps because this cultivar produces large amounts of ethylene (Watkins et al.,
2000).
Interestingly, plant display life and flower longevity of campanula Dark Blue and Blue Clips
and holiday cactus Dark Marie were greater with
STS than with 1-MCP (Serek and Sisler, 2001). In
addition, senescence of gypsophila flowers open at
the time of treatment was delayed by both 1-MCP
and STS but only STS delayed senescence of floral
buds not opened at the time of treatment (Newman et al., 1998). The authors theorized that while
both 1-MCP and STS block most ethylene receptors after pretreatment, new sites are formed after
further plant development. These new receptors
are not blocked by 1-MCP but are blocked by
STS, which apparently remains active in the plant
tissue longer than 1-MCP. Celikel and Reid (2002)
also noted that flowers treated with STS had
longer vase life than 1-MCP-treated flowers.
However, treatment with 5 nl l1 1-MCP for 6 h
allowed carnation flowers to be stored for 10 days
at room temperature and still prevented the effects
of exogenous ethylene (1.0 ml l 1) (Sisler et al.,
1996a). 1-MCP appeared to be permanently bound
in carnation tissue (Sisler et al., 1996a). Cameron
and Reid (2001) found that the half-life of 1-MCP
treatment was 2, 3, or 6 days at 88, 69, or 54 8F
(25, 20.7 or 12 8C) indicating the new binding sites
were apparently being synthesized. Receptor regeneration may provide explanation for some
differences in response to the various ethylene
receptor blockers.
6. Physiological responses
6.1. Ethylene production
1-MCP lowered ethylene production in strawberry (Jiang et al., 2001), slowed ethylene production in apricots and plums (Dong et al., 2002), and
15
inhibited ethylene production in Fuji (Fan and

Mattheis, 1999a,b), Red Delicious and Granny
Smith apples (Fan et al., 1999a,b). When fruit
were treated with 1-MCP the ethylene climacteric
was delayed by 6 days and reduced in magnitude
over 50% in avocado (Jeong et al., 2002) and it was
also reduced in plums (Abdi et al., 1998) and
apricots (Fan et al., 2000a) treated with 1-MCP.
Reduction of ethylene production in kiwifruit was
dependent on temperature (Kim et al., 2001). 1MCP (5 nl l 1) reduced by 2/3 the climacteric rise
in ethylene production in carnation flowers but did
not prevent endogenous ethylene production in the
flowers (Sisler et al., 1996a). 1-MCP prevented the
pollination-induced increase in ethylene production and the enhanced senescence of moth orchid
(Phalaenopsis ) Herbert Hager flowers (Porat et
al., 1995a). Serek et al. (1995b) determined the 1MCP binding constant (Kd) to be 2.1 nl l 1 on cut
carnations Sandra and 8.1 nl l 1 on potted rose
Victory Parade. Sisler et al. (1996a) determined a
lower Kd of 0.5 nl l1 for carnation. Citrus leaves
and leaf explants and cut metrosideros flowers
treated with 1-MCP showed a concentration
dependent increase in ethylene production (Zhong
et al., 2001; Sun et al., 2000).
In contrast, in pineapple (Arianas comosus) the
decline in ethylene production was slowed in 1MCP-treated fruit compared with a control,
resulting in 1-MCP fruit producing more ethylene
than the control (Selvarajah et al., 2001). Ethylene
production was increased by 1-MCP in Penicillium
digitatum infected grapefruit (Mullins et al., 2000)
through uninhibited expression of the stress-associated ACC synthase genes; however these fruit
remained green. Ethylene production was increased by 1-MCP treatment in coriander (Jiang
et al., 2002b).
The ethylene biosynthetic enzymes ACC oxidase, ACC synthase and associated mRNA accumulation were inhibited by 1-MCP (Dong et al.,
2001a; Mathooko et al., 2001; Owino et al., 2002;
Shiomi et al., 1999). In addition, the use of 1-MCP
in peaches showed that CO2 does not regulate
ACC synthase during ripening (Mathooko et al.,
2001). The inhibition of accumulation of ACC
synthase and ACC oxidase mRNAs by 1-MCP
indicates that a strong positive feedback regulation
16
exists for ethylene production in tomato even at

the turning-color stage. After treatment, recovery
of transcript levels to those of the untreated
control required 2/4 days (Nakatsuka et al.,
1997). As shown through the use of 1-MCP,
negative feedback regulation of ethylene production is mediated through the repression of certain
ethylene biosynthetic pathway genes expressed in
the early stages of ripening (system I) while other
genes are not effected by ethylene (Nakatsuka et
al., 1998). This 1-MCP mediated decrease in
transcript levels of ripening-related genes in tomato may account for its effect on ripening
(Hoeberichts et al., 2002).
In yeast the ETR1 and ERS1 encoded ethylene
receptors, showed equal sensitivity to 1-MCP
indicating that their affinity for ethylene is similar
(Hall et al., 2000). Hall et al. (2000) also offered
evidence that 1-MCP binding to a subset of
receptors might be enough to keep ethylene
responses turned off.
In melon (Cucumis melo) plants ethylene-induced ACC oxidase gene expression was undetectable after 1-MCP treatment; however, transcript
accumulation due to wounding was not effected by
1-MCP, indicating separate signal induction pathways (Bouquin et al., 1997).
When pears (Pyrus communis ) which had already received their chilling requirement for ripening were treated with 1-MCP, they failed to ripen
for at least 26 days when placed at warm
temperatures. ACC synthase, ACC oxidase expression and ethylene production were much
reduced in 1-MCP-treated pears (Lelievre et al.,
1997; De Wild et al., 1999). In contrast to the
above, ethylene production was increased by 1MCP treatment of bananas (Golding et al., 1998).
1-MCP had a greater affinity for the receptor than
ethylene as determined by the Km for 1-MCP in
banana softening (Jiang et al., 1999b).
6.2. Chlorophyll degradation and color changes
1-MCP prevented or delayed chlorophyll degradation and various types of color changes in a
wide range of crop species. Degreening of oranges
(Citrus sinensis ) was blocked by 1-MCP and
stimulated by exogenous ethylene (Porat et al.,
1999). However, degreening was not effected by 1MCP in Oroblanco, a pummelo-grapefruit hybrid (Porat et al., 2001). 1-MCP delayed skin color
change in avocado (Feng et al., 2000; Jeong et al.,
2002). 1-MCP delayed chlorophyll degradation in
coriander (Jiang et al., 2002b) and in leafy Asian
vegetables (Able et al., in press) when ethylene was
present. 1-MCP prevented broccoli yellowing both
with and without exogenous ethylene (Fan and
Mattheis, 2000a) and prevented rotting in the
presence of ethylene (Ku and Wills, 1999). The
effect on broccoli yellowing was not concentration
dependent.
1-MCP was effective in reducing leaf drop and
yellowing of pothos (Epipremnum pinnatum ) cuttings stored at 73 8F (23 8C) for 4 days (Muller et
al., 1997). Interestingly, STS at 2 mM injured the
cuttings sufficiently to reduce rooting percentage,
number of roots, total root length, and dry weight
compared with untreated cuttings or cuttings
treated with 1-MCP. 1-MCP, however, did not
influence rooting percentage, number of roots,
total root length, or dry weight compared with
untreated control cuttings. In contrast, neither 1MCP nor STS had any effect on leaf yellowing of
rose plants due to dark storage. Darkness-induced
leaf yellowing of rose may not be ethylene
influenced or the chemicals may not have penetrated the tissue sufficiently to produce a response.
1-MCP also retarded storage-induced leaf yellowing of geranium (Pelargonium zonale Isabel), and
hibiscus (H. rosa-sinensis ) unrooted cuttings but
had no effect on chrysanthemum (Dendranthema
grandiflorum Coral Charm) unrooted cuttings
(Serek et al., 1998). Unfortunately, 1-MCP tended
to reduce rooting of geranium, hibiscus and
chrysanthemum cuttings. Muller et al. (1998)
also noted that 1-MCP reduced foliar abscission
and chlorosis of croton cuttings and reduced
rooting.
With 1-MCP treatment bananas became yellow
before they softened (Sisler et al., 1996b). When
bananas treated with 1-MCP were subsequently
treated with propylene, degreening was delayed
and fruit had a patchy, uneven color (Golding et
al., 1998). Harris et al. (2000) also found that
bananas treated with 1-MCP did not have commercially acceptable color development.
Degreening of Fuji apples was inhibited by 1MCP (Fan and Mattheis, 1999b). While 1-MCPtreated Red Chief apples had a greener background color than untreated fruit, chlorophyll
fluorescence measurements indicated that loss of
chloroplast function was largely independent of
ethylene (Mir et al., 2001).
The chl a/b color ratio was reduced in 1-MCPtreated strawberries when compared with the
control (Tian et al., 2000). Jiang et al. (2001)
found that 1-MCP maintained fruit color in
strawberries; however, it inhibited phenylalanine
ammonia lyase activity and lowered anthocyanin
production.
In apricots 1-MCP-treated fruit were greener
and exhibited less color change than untreated
controls (Fan et al., 2000a). This was also found to
be true in peaches (Kluge and Jacomino, 2002).
Others researchers found that apricots and plum
color changes were not affected by 1-MCP (Dong
et al., 2002) and Abdi et al. (1998) concluded that
ethylene was not necessary for color development
in climacteric and suppressed climacteric plums.
There was no influence of 1-MCP on postharvest
color changes or stem browning in sweet cherries
(Prunus avium ) (Gong et al., 2002).
6.3. Protein and membrane effects
1-MCP retarded the decline in protein in senescing coriander leaves (Jiang et al., 2002b). 1-MCP
prevented an exogenous ethylene-induced (1 /12 ml
l 1) increase in electrolytic leakage, a decrease of
membrane proteins and a decrease in lipid fluidity
in petunia flowers. In a non-ethylene environment
1-MCP increased individual flower longevity, fresh
weight and total protein content compared with
untreated controls but had no effect on electrolytic
leakage, membrane proteins, or lipid fluidity
(Serek et al., 1995d).
6.4. Respiration
In general, 1-MCP reduced respiration rates or
delayed increases in respiration. In strawberry 1MCP inhibited the ethylene-induced respiratory
increase in early-harvested fruit, but not lateharvested fruit (Tian et al., 2000). Respiratory
17
increase in avocado was delayed by about 6 days

and reduced in magnitude by about 40% with 1MCP treatment. The respiratory climacteric in
plums was delayed by 1-MCP (Abdi et al., 1998;
Dong et al., 2002). Respiration rate was reduced in
apricots treated with 1-MCP (Fan et al., 2000a).
However, 1-MCP treatment had no effect on
nectarine (Prunus persica) (Dong et al., 2001a)
and apricot (Dong et al., 2002). The differing
results in apricots might be due to fruit maturity,
cultivar, or some other unknown factor. The
stimulation of respiration in sweet cherry by
exogenous ethylene occurred regardless of 1MCP treatment (Gong et al., 2002). In the
presence of exogenous ethylene the respiratory
rise in carrots (Daucus carota ) was prevented (Fan
and Mattheis, 2000b). Respiration was inhibited in
1-MCP-treated broccoli in a concentration dependent manner when tested in the range of 0 /1 ml l1
(Fan and Mattheis, 2000a). Respiration in tomato
was suppressed for 6/8 days while titratable
acidity (TA) was maintained throughout the life
of the fruit (Wills and Ku, 2002). Respiration was
suppressed in propylene-treated bananas subsequently gassed with 1-MCP (Golding et al., 1998).
Respiration was inhibited in Fuji (Fan and
Mattheis, 1999b), Granny Smith and Red Delicious (Fan et al., 1999a,b). However, in post
irradiation storage, Gala apples that had been
treated with 1-MCP and irradiated showed a
stimulation in respiration whereas there was no
effect on control fruit (Fan and Mattheis, 2001).
Interestingly, respiration rates in coriander treated
with 1-MCP were equal to or greater than the
control (Jiang et al., 2002b).
6.5. Volatiles
Aroma development is apparently an ethylene
dependent process as evidenced by reduced aroma
in 1-MCP-treated plums, and by the fact that
propylene application restored aroma (Abdi et al.,
1998). This would indicate that the propylene
either reacts with different receptors or receptors
became available. 1-MCP-treated Anna apples
retained more of the volatiles associated with
newly harvested apples and lesser amounts of
volatiles associated with ripening apples. Un-
18
treated apples developed a more ripe and fruity

volatile
composition
than
1-MCP-treated
apples. A taste panel preferred the 1-MCP-treated
fruit to the untreated fruit even though the
volatiles were more like a greener fruit (Lurie et
al., 2002). Volatile formation in apples is differentially inhibited by 1-MCP (Fan and Mattheis,
1999a). 1-MCP inhibited total alcohol and total
ester formation in Fuji (Fan and Mattheis,
1999a) and Gala apples (Fan and Mattheis,
2001), while hexanol production was not
affected in Fuji. Total volatiles were inhibited in
McIntosh and Delicious (Rupasinghe et al.,
2000b).
Apricots showed delayed development of volatile alcohols and esters after 1-MCP treatment
(Fan et al., 2000a). Melon volatile synthesis is
reduced by 1-MCP (Flores et al., 2002). Volatile
production in bananas was delayed and suppressed in propylene-treated bananas also treated
with 1-MCP (Golding et al., 1998). 1-MCP also
caused a quantitative change in the volatile
composition of bananas, an increase in alcohols,
and a decrease in their related esters (Golding et
al., 1999). Induction of volatile off flavors was
increased by ethylene and 1-MCP in oranges
(Porat et al., 1999), which may have resulted
from unseen stress or damage to the fruit. Interestingly, the effect of 1-MCP on volatiles of floral
crops has not been reported. Either 1-MCP does
not affect floral crop volatiles or changes occurred
but have not been documented.
6.6. Softening
While 1-MCP delayed softening of most fruits,
other crop species were not affected. 1-MCP
delayed softening in avocado by 4.4 days, custard
apple (Annona ) by 3.4 days, and mango by 5.1
days (Hofman et al., 2001). Papaya (Carica
papaya ) softening was delayed by 15.6 days with
1-MCP treatment (Hofman et al., 2001). Apple
tissue mechanical properties were found to change
less in 1-MCP-treated fruit than in untreated fruit
(Baritelle et al., 2001). Apples maintained firmness
after 1-MCP treatment, with data presented on
Delicious, Granny Smith, Fuji, Ginger Gold,
Gala, Idared, Jonagold, and McIntosh (Ru-
pasinghe et al., 2000b; Fan et al., 1999a; Watkins

et al., 2000; Mir et al., 2001). 1-MCP treatment
maintained apple firmness better than controlled
atmosphere (CA) storage (Mir et al., 2001).
Watkins et al. (2000) found that the combination
of 1-MCP and CA was better than either alone.
Firmness was maintained by 1-MCP in apricots
(Fan et al., 2000a), nectarines (Dong et al., 2001a),
peaches (Kluge and Jacomino, 2002), and plums
(Dong et al., 2002, 2001b; Skog et al., 2001). After
cold storage 1-MCP did not allow nectarines to
ripen normally compared with ethylene-treated
fruit, which softened normally (Dong et al.,
2001a). In avocado 1-MCP delayed ethyleneinduced fruit softening (Feng et al., 2000; Jeong
et al., 2002). Persimmon fruit softening was
delayed (Nakano et al., 2001, 2002).
More detailed examinations of fruit softening
showed that polygalacturonase (PG) and cellulose
activities were lowered by 1-MCP; however, activities of both enzymes were still present and
avocado fruit ripened and softened normally
(Feng et al., 2000). Jeong et al. (2002) found that
PG activity was completely suppressed for up to 10
days, indicating that softening can occur without
PG activity in avocado. Pectinmethylesterase activity was delayed in 1-MCP-treated fruit compared with the control but followed a similar
pattern (Jeong et al., 2002). Uronic acid content
in avocado decreased in the control but the 1MCP-treated fruit showed little change (Jeong et
al., 2002). In plums exo-PG and endo-glucanase
were lower in 1-MCP-treated fruit when compared
to the untreated fruits while pectin esterase and
endo-PG were similar (Dong et al., 2001b). By
using 1-MCP to block ethylene action, it was
found that endo-beta-1,4-glucanase was ethylene
induced during raspberry (Rubus idaeus ) fruit
abscission (Iannetta et al., 2000).
Orange softening was not affected by 1-MCP or
ethylene (Porat et al., 1999). No firmness difference was detected between ethylene only or 1MCP/ethylene-treated strawberry fruit (Tian et
al., 2000). 1-MCP at 2 ml l1 actually gave greater
firmness loss than no treatment (Tian et al., 2000);
however, Jiang et al. (2001) found that 1-MCP
maintained strawberry firmness.
19
6.7. Titratable acidity
6.9. Weight loss
The effect of 1-MCP on TA is mixed, with some

crops being affected and others not. 1-MCP
delayed the decline in ascorbic acid in pineapples
(Selvarajah et al., 2001), prevented ethylene-induced acidity loss in carrots (Fan and Mattheis,
2000b), completely inhibited TA loss in tomatoes
(Wills and Ku, 2002), delayed TA loss in plums
(Dong et al., 2002), and maintained TA in Red
Delicious, Granny Smith, Fuji, Jonagold,
Ginger Gold, and Gala apples (Fan et al.,
1999a,b). Watkins et al. (2000) found that TA of
Law Rome, Delicious, Empire, and McIntosh
was always higher in 1-MCP-treated fruit during
air storage, but effects were inconsistent in CA
storage. In contrast, 1-MCP did not affect TA in
apricots (Dong et al., 2002) or Red Chief apples
during storage at several temperatures (Mir et al.,
2001). 1-MCP did not affect titratable acid content
of Shamouti oranges (Porat et al., 1999).
While 1-MCP did not affect weight loss in

oranges (Porat et al., 1999), it delayed weight
loss in avocado (Jeong et al., 2002).
6.8. Sugars
Soluble solids were higher in 1-MCP-treated
pineapple (Selvarajah et al., 2001), papaya (Hofman et al., 2001), and apples (Fan et al., 1999a).
However, soluble solids were reduced in 1-MCPtreated strawberries regardless of the presence or
absence of exogenous ethylene (Tian et al., 2000).
Soluble solids were unaffected by 1-MCP in
oranges (Porat et al., 1999), apricots and plums
(Dong et al., 2002), custard apple and mango
(Hofman et al., 2001) and apples (Rupasinghe et
al., 2000b; DeEll et al., 2002). The contrasting
results with apples are notable and may be due to
different cultivars or other experimental conditions used. Watkins et al. (2000) found differences
in responses of apple cultivars with 1-MCP-treated
McIntosh and Law Rome fruit having lower
soluble solids and Delicious and Empire having
higher soluble solids than untreated fruit. They
also found that storage method, whether air or CA
also influenced the soluble solids, with CA storage
giving more inconsistent or weaker effects that air
storage.
7. Disorders, injury and disease

The effect of 1-MCP on various disorders and
diseases has been inconsistent with results being
species specific. In some cases 1-MCP treatment
alleviates disorders; for example, it reduced superficial scald development in apples (Fan et al.,
1999b) and prevented ethylene-induced gummosis
of tulip (Tulipa gesneriana ) bulbs (De Wild et al.,
2002). With some species, plants treated with 1MCP had increased incidence or severity of
diseases and disorders. In strawberries (Fragaria
/anaassa) 1-MCP concentrations greater than 15
nl l 1 are associated with increased decay of fruit
(Ku et al., 1999). Jiang et al. (2001) also found an
increase in disease with high 1-MCP concentrations on strawberries. 1-MCP may inhibit a
beneficial metabolic response or stimulate an
undesirable characteristic, possibly relating to a
natural defense mechanism. (Ku et al., 1999).
Lower phenolic content in 1-MCP-treated strawberries was thought to account for the increased
disease incidence (Jiang et al., 2001). Synthesis of
the phytoalexin, 6-methoxymellin, found in carrot,
was inhibited by 1-MCP (Fan et al., 2000b). Diaz
et al. (2002) found that treatment with 1-MCP
increased susceptibility of tomatoes to Botrytis
cinerea.
1-MCP is associated with higher incidences of
external blemishes in papaya and custard apple
(Hofman et al., 2001). Rot incidence was higher in
1-MCP-treated avocado, custard apple and papaya compared with untreated fruits (Hofman et
al., 2001). Mango had twice the amount of stem
rot with 1-MCP treatment than did an untreated
control (Hofman et al., 2001). Wooliness and
reddening of nectarines was more prominent in
1-MCP-treated fruit as compared with untreated
controls (Dong et al., 2002). A higher incidence of
internal browning occurred when apricots were
treated with 1-MCP before storage, but internal
20
browning was not increased if fruit were treated

after storage (Dong et al., 2002). However, Fan et
al. (2000a) reporting on the same crop did not
mention quality problems in apricots. Plum quality was not adversely affected by 1-MCP treatment
(Dong et al., 2002, 2001b). While both treated and
untreated peaches stored at 5 8C had internal
browning, the problem was more exaggerated in 1MCP-treated fruit. Internal browning was not
associated with 1-MCP treatment when peaches
were stored at 0 or 10 8C or later harvested fruit
was used (Fan et al., 2002). Water soaking in
Charentais cantaloupe melons was not effected by
1-MCP (du Chatenet et al., 2000).
Chilling injury in oranges was more severe in
ethylene-treated oranges than in oranges without
ethylene treatment. However, treatment with 1MCP did not prevent the ethylene effect, and
oranges showed more chilling injury than did the
controls (Porat et al., 1999). 1-MCP-treated oranges had higher rot incidence than control fruit
(Porat et al., 1999), and the authors postulated
that endogenous ethylene in oranges might be
necessary to maintain defense mechanisms. Chilling-induced and ethylene-enhanced leaf abscission
in ixora (Ixora coccinea ) plants was inhibited by 1MCP, indicating ethylene involvement in the
process (Michaeli et al., 1999).
However, 1-MCP treatments reduced or eliminated chilling-induced mesocarp discoloration and
decay development in avocado (Pesis et al., 2002)
and internal browning in pineapple (Selvarajah et
al., 2001), probably due to a difference in mechanism from previously listed disorders. Decay development was slowed by using 1-MCP in apricots
(Dong et al., 2002). Ethylene-induced russet spotting on the midribs of lettuce (Lactuca sativa) was
reduced by 1-MCP (Fan and Mattheis, 2000b;
Manleitner et al., 2001). Postharvest life of
shredded lettuce was also extended by 1-MCP
use (Wills et al., 2002). In the presence of
exogenous ethylene, 1-MCP-treated carrots
showed little accumulation of isocoumarin when
compared with the control, resulting in less bitter
carrots (Fan and Mattheis, 2000b).
Using 1-MCP allowed researchers to determine
that tobacco (Nicotiana attenuata ) plants produced ethylene in response to insect damage and
that ethylene suppressed the plants nicotine response, thereby making the insect able to feed on
the plant (Kahl et al., 2000; Winz and Baldwin,
2001). In addition, using 1-MCP and tomato wild
type and hypersensitive phenotypes, the hypersensitive response to pathogens was shown to be a
result of increased ethylene sensitivity (Ciardi et
al., 2001). In contrast, infection rate in grapefruit
(Citrus /paradisi ) was not slowed by 1-MCP
treatment nor was host susceptibility altered
(Mullins et al., 2000).
Mixed results occurred with 1-MCP application
on apples. Irradiated apples treated with 1-MCP
showed more internal damage than untreated
fruit. However, low temperatures reduced browning (Fan and Mattheis, 2001). Granny Smith
apples did not develop peel greasiness when
treated with 1-MCP (Fan et al., 1999b) and
Fuji apples developed less core flush than the
control when treated with 1-MCP (Fan et al.,
1999b). 1-MCP treatments reduced but did not
prevent decay in apples, especially at higher
temperatures (Mir et al., 2001).
1-MCP was found to suppress superficial scald
by 30% in McIntosh and by 90% in Delicious
apples during air storage (Rupasinghe et al.,
2000a). In Granny Smith apples 1-MCP suppressed a-farnesene, reduced the amount of its
oxidation products, conjugated trienes and 6methyl-5-hepten-2-one, and reduced scald (Fan et
al., 1999b). Superficial scald was also suppressed
in Delicious and Law Rome apples along with a
reduction in a-farnesene and conjugated triols
(Watkins et al., 2000). 1-MCP reduced 3-hydroxy-3-methylglutaryl coenzyme A reductase
(HMGR) activity by 30% and respiration by 60%
in apples, resulting in a 72% reduction in afarnesene (Rupasinghe et al., 2001). While E ,E a-farnesene synthase activity was no different in 1MCP-treated apples when compared with the
control, 1-MCP treatment greatly reduced afarnesene in apple skin (Rupasinghe et al.,
2000a). In Cortland apple a 3 h 1-MCP treatment
resulted in less than 30% scald while a 6 h
treatment gave less than 15% scald, with treatment
temperature having no effect on the level of scald
control at either timing (DeEll et al., 2002).
Porat et al. (1995a) used 1-MCP concentrations

of 25, 250 and 500 nl l 1 on garden phlox
Rembrandt and noted no visible phytotoxicity
at any rate and no reports of phytotoxicity have
been documented in ornamental crops.
8. In vitro culture
Not surprisingly, the role of ethylene in in vitro
culture has led researchers to consider the effects
of 1-MCP in vitro. 1-MCP induced lateral shoot
development of hanconia (Hanconia speciosa ) in
vitro-grown shoots due to inhibition of ethylene,
which normally induces apical dominance (Pereira-Netto, 2001). 1-MCP increased the number of
somatic embryos of geranium hypocotyls cultures
when applied during the initial 3 days of induction
(Hutchinson et al., 1998).
9. Summary and future research directions

Many questions remain concerning 1-MCP and
its reactions. The effect of 1-MCP treatment
temperature seems to be important in some crops
but not others, and could be due to differences in
receptor sensitivity. The range of effective concentrations is large (Tables 1 and 2) and there is no
explanation as to why this diversity exists. The
future availability of binding sites after treatment
varies between crops. Some crops can regenerate
sites fairly quickly (flowers, tomatoes), rendering
1-MCP treatment ineffective, while others seem to
have difficulty at times (bananas). With several
plant groups (stone fruits, bananas, and some
green vegetables), species and cultivar, growing
conditions, and maturity may influence the results
of 1-MCP treatment since differences in the 1MCP effects have been reported.
Use of 1-MCP is just starting to help unravel the
effects of ethylene on postharvest processes. 1MCP reduces production of aromatic compounds
in many fruits yet its effect on floral aromas is
unstudied. There are indications that in some
plants ethylene production is important in disease
resistance and development of physiological disorders. The role of ethylene in starch/sugar con-
21
versions is still not clear: some workers report 1MCP effects on conversions during ripening while
others do not.
The amount of ethylene necessary to stimulate
some ripening associated changes may not be the
same for every process, leading to the hypothesis
that multiple binding sites may exist. For example,
yellowing might be delayed but respiration is only
partially repressed. While both 1-MCP and STS
greatly increase the postharvest life of most potted
plants exposed to exogenous ethylene, compared
with untreated plants exposed to ethylene, the
postharvest life is much shorter in some species
than if the crops were never exposed to exogenous
ethylene (Serek et al., 1994b, 1995b). Once the
minimum concentration for effectiveness is
reached, increasing 1-MCP or STS concentration
has no affect on improving postharvest life of
ethylene-treated flowers. Thus, 1-MCP and STS
may not completely block ethylene action in some
species.
Similar situations exist in apples where the 1MCP response of some cultivars, such as McIntosh, is less than other cultivars such as Delicious, giving the appearance that there is
incomplete blocking of ethylene action. Ethylene
production of cultivars such as McIntosh has
traditionally been difficult to control.
With floral crops the comparison of 1-MCP
with STS remains of interest. Both compounds
appear to work equally well in preventing accelerated senescence due to exogenous ethylene, but
STS may be more effective in delaying senescence
than 1-MCP in the absence of exogenous ethylene.
We have to wonder if STS is more effective in
counteracting the effects of endogenous ethylene,
or if STS influences more physiological processes
than ethylene while the effects of 1-MCP are
limited only to ethylene?
Postharvest research on 1-MCP and ethylene
needs to continue in two directions. The first is to
learn how to effectively use 1-MCP to maximize
profits for agricultural businesses and provide a
quality consumer product. The second is to use 1MCP as a research tool to understand the role of
ethylene in plant development. Of course following
these paths will undoubtedly open others, to the
22
benefit of horticultural science, commercial agriculture and plant science.
References
Abdi, N., McGlasson, W.B., Holford, P., Williams, M.,
Mizrahi, Y., 1998. Responses of climacteric and suppressed-climacteric plums to treatment with propylene and
1-methylcyclopropene. Postharvest Biol. Technol. 14, 29 /
39.
Able, A.J., Wong, L.S., Prasad, A., OHare, T.J., 2002a. 1MCP is more effective on a floral brassica (Brassica oleracea
var. italica L. ) than a leafy brassica (Brassica rapa var.
chinensis ). Postharvest Biol. Technol. 26, 147 /155.
Able, A.J., Wong. L.S., Prasad, A., OHare, T.J., 2002b. The
effects of 1-methylcyclopropene on the shelf-life of minimally processed leafy Asian vegetables. Postharvest Biol.
Technol. (in press).
Baritelle, A.L., Hyde, G.M., Fellman, J.K., Varith, J., 2001.
Using 1-MCP to inhibit the influence of ripening on impact
properties of pear and apple tissue. Postharvest Biol.
Technol. 23, 153 /160.
Bosma, T., Dole, J.M., 2002. Postharvest handling of cut
Campanula medium flowers. HortScience 37, 954 /958.
Bouquin, T., Lasserre, E., Pradier, J., Pech, J.C., Balague, C.,
1997. Wound and ethylene induction of the ACC oxidase
melon gene CM-ACO1 occurs via two direct and independent transduction pathways. Plant Mol. Biol. 35, 1029 /
1035.
Cameron, A.C., Reid, M.S., 2001. 1-MCP blocks ethyleneinduced petal abscission of Pelargonium peltatum but the
effect is transient. Postharvest Biol. Technol. 22, 169 /177.
Celikel, F.G., Reid, M.S., 2002. Postharvest handling of stock
(Matthiola incana ). HortScience 37, 144 /147.
Celikel, F.G., Dodge, L.L., Reid, M.S., 2002. Efficacy of 1MCP (1-methylcyclopropene) and Promalin for extending
the post-harvest life of Oriental lilies (Lilium /Mona Lisa
and Stargazer). Scientia Hort. 93, 149 /155.
Ciardi, J.A., Tieman, D.M., Jones, J.B., Klee, H.J., 2001.
Reduced expression of the tomato ethylene receptor gene
LeETR4 enhances the hypersensitive response to Xanthomonas campestris pv. Vesicatoria . Mol. Plant-Microbe Inter.
14, 487 /495.
DeEll, J.R., Murr, D.P., Porteous, M.D., Rupasinghe, H.P.V.,
2002. Influence of temperature and duration of 1-methylcyclopropene (1-MCP) treatment on apple quality. Postharvest Biol. Technol. 24, 349 /353.
De Wild, H.P.J., Gude, H., Peppelenbos, H.W., 2002. Carbon
dioxide and ethylene interactions in tulip bulbs. Physiol.
Plant. 114, 320 /326.
De Wild, H.P.J., Woltering, E.J., Peppelenbos, H.W., 1999.
Carbon dioxide and 1-MCP inhibit ethylene production and
respiration of pear fruit by different mechanisms. J. Exp.
Bot. 50, 837 /844.
Diaz, J., ten Have, A., van Kan, J.A.L., 2002. The role of
ethylene and wound signaling in resistance of tomato to
Botrytis cinerea . Plant Physiol. 129, 1341 /1351.
Dong, L., Lurie, S., Zhou, H., 2002. Effect of 1-methylcyclopropene on ripening of Canino apricots and Royal Zee
plums. Postharvest Biol. Technol. 24, 135 /145.
Dong, L., Zhou, H., Sonego, L., Lers, A., Lurie, S., 2001a.
Ethylene involvement in the cold storage disorder of
Flavortop nectarine. Postharvest Biol. Technol. 23, 105 /
115.
Dong, L., Zhou-Hong, W., Sonego, L., Lers, A., Lurie, S.,
2001b. Ripening of Red Rosa plums: effect of ethylene and
1-methylcyclopropene. Aust. J. Plant Physiol. 28, 1039 /
1045.
du Chatenet, C., Latche, A., Olmos, E., Ranty, B., Charpenteau, M., Ranjeva, R., Pech, J.C., Graziana, A., 2000.
Spatial-resolved analysis of histological and biochemical
alterations induced by water-soaking in melon fruit. Physiol. Plant. 110, 248 /255.
Elgar, H.J., Woolf, A.B., Bieleski, L., 1999. Ethylene production by three lily species and their response to ethylene
exposure. Postharvest Biol. Technol. 16, 257 /267.
Environmental Protection Agency. Federal Register, July 26,
2002. Vol. 67, Number 144, pp. 48 796 /48 800.
Fan, X., Mattheis, J.P., 1999a. Impact of 1-methylcyclopropene
and methyl jasmonate on apple volatile production. J.
Agric. Food Chem. 47, 2847 /2853.
Fan, X., Mattheis, J.P., 1999b. Methyl jasmonate promotes
apple fruit degreening independently of ethylene action.
HortScience 34, 310 /312.
Fan, X., Mattheis, J.P., 2000a. Yellowing of broccoli in storage
is reduced by 1-methylcyclopropene. HortScience 35, 885 /
887.
Fan, X., Mattheis, J.P., 2000b. Reduction of ethylene-induced
physiological disorders of carrots and iceberg lettuce by 1methylcyclopropene. HortScience 35, 1312 /1314.
Fan, X., Mattheis, J.P., 2001. 1-Methylcyclopropene and
storage temperature influence responses of Gala apple
fruit to gamma irradiation. Postharvest Biol. Technol. 23,
143 /151.
Fan, X., Argenta, L., Mattheis, J.P., 2000a. Inhibition of
ethylene action by 1-methylcyclopropene prolongs storage
life of apricots. Postharvest Biol. Technol. 20, 135 /142.
Fan, X., Argenta, L., Mattheis, J.P., 2002. Interactive effects of
1-MCP and temperatures on Elberta peach quality.
Fan, X., Blankenship, S.M., Mattheis, J.P., 1999a. 1-Methylcyclopropene inhibits apple ripening. J. Am. Soc. Hort. Sci.
124, 690 /695.
Fan, X., Mattheis, J.P., Blankenship, S.M., 1999b. Development of apple superficial scald, soft scald, core flush, and
greasiness is reduced by MCP. J. Agric. Food Chem. 47,
3063 /3068.
Fan, X., Mattheis, J.P., Roberts, R.G., 2000b. Biosynthesis of
phytoalexin in carrot root requires ethylene action. Physiol.
Plant. 110, 450 /454.
Feng, X., Apelbaum, A., Sisler, E.C., Goren, R., 2000. Control
of ethylene responses in avocado fruit with 1-methylcyclopropene. Postharvest Biol. Technol. 20, 143 /150.
Fisher, F., Applequist, D.E., 1965. Synthesis of 1-methylcyclopropene. J. Org. Chem. 30, 2089 /2090.
Flores, F., El Yahyaoui, F., de Billerbeck, G., Romojaro, F.,
Latche, A., Bouzayen, M., Pech, J.C., Ambid, C., 2002.
Role of ethylene in the biosynthetic pathway of aliphatic
ester aroma volatiles in Charentais Cantaloupe melons. J.
Exp. Bot. 53, 201 /206.
Golding, J.B., Shearer, D., McGlasson, W.B., Wyllie, S.G.,
1999. Relationships between respiration, ethylene, and
aroma production in ripening bananas. J. Agric. Food
Chem. 47, 1646 /1651.
Golding, J.B., Shearer, D., Wyllie, S.G., McGlasson, W.B.,
1998. Application of 1-MCP and propylene to identify
ethylene-dependent ripening processes in mature banana
fruit. Postharvest Biol. Technol. 14, 87 /98.
Gong, Y., Fan, X., Mattheis, J.P., 2002. Responses of Bing
and Rainier sweet cherries to ethylene and 1-methylcyclopropene. J. Am. Soc. Hort. Sci. 127, 831 /835.
Goren, R., Feng, X., Apelbaum, A., Sisler, E.C., Ben-Aire, R.,
Philosoph-Hadas, S., 2001. Effect of two structural analogues of 1-methylcyclopropene on ethylene-induced ripening
of avocado fruits. Acta Hort. 552, 163 /166.
Hall, A.E., Findell, J.L., Schaller, G.E., Sisler, E.C., Bleecker,
A.B., 2000. Ethylene perception by the ERS1 Protein in
Arabidopsis . Plant Physiol. 123, 1449 /1457.
Hamrick, D., 2001. Ethylbloc goes liquid. GrowerTalks 65, 105.
Harris, D.R., Seberry, J.A., Wills, R.B.H., Spohr, L.J., 2000.
Effect of fruit maturity on efficiency of 1-methylcyclopropene to delay the ripening of banana. Postharvest Biol.
Technol. 20, 303 /308.
Hoeberichts, F.A., Van Der Plas, L.H.W., Woltering, E.J.,
2002. Ethylene perception is required for the expression of
tomato ripening-related genes and associated physiological
changes even at advanced stages of ripening. Postharvest
Biol. Technol. 26, 125 /133.
Hofman, P.J., Jobin-Decor, M., Meiburg, G.F., Macnish, A.J.,
Joyce, D.C., 2001. Ripening and quality responses of
avocado, custard apple, mango and papaya fruit to 1methylcyclopropene. Aust. J. Exp. Agric. 41, 567 /572.
Hutchinson, M.J., Murr, D., Krishnaraj, S., Senaratna, T.,
Saxena, P.K., 1998. Does ethylene play a role in thidiazuron-regulated somatic embryogenesis of geranium (Pelargonium /hortorum Bailey) hypocotyls cultures. In-vitro
Cell. Dev. Bio.-Plant 33, 136 /141.
Iannetta, P.P.M., Wyman, M., Neelam, A., Jones, C., Taylor,
M.A., Davies, H.V., Sexton, R., 2000. A causal role for
ethylene and endo-beta-1,4-glucanase in the abscission of
red-raspberry (Rubus idaeus ) drupelets. Physiol. Plant. 110,
535 /543.
Jeong, J., Huber, D.J., Sargent, S.A., 2002. Influence of 1methylcyclopropene (1-MCP) on ripening and cell-wall
matrix polysaccharides of avocado (Persea americana ) fruit.
Postharvest Biol. Technol. 25, 241 /364.
23
Jiang, Y., Joyce, D., 2000. Effects of 1-methylcyclopropene

alone and in combination with polyethylene bags on the
postharvest life of mango fruit. Ann. Appl. Biol. 137, 321 /
327.
Jiang, Y., Joyce, D.C., 2002. 1-Methylcyclopropene treatment
effects on intact and fresh-cut apple. J. Hort. Sci. Biotech.
77, 19 /21.
Jiang, Y., Joyce, D.C., Macnish, A.J., 1999a. Extension of the
shelf life of banana fruit by 1-methylcyclopropene in
combination with polyethylene bags. Postharvest Biol.
Technol. 6, 187 /193.
Jiang, Y., Joyce, D.C., Macnish, A.J., 1999b. Responses of
banana fruit to treatment with 1-methylcyclopropene. Plant
Growth Regul. 28, 77 /82.
Jiang, Y., Joyce, D.C., Macnish, A.J., 2000. Effect of abscisic
acid on banana fruit ripening in relation to the role of
ethylene. J. Plant Growth Regul. 19, 106 /111.
Jiang, Y., Joyce, D.C., Terry, L.A., 2001. 1-methylcyclcopropene treatment affects strawberry fruit decay. Postharvest
Biol. Technol. 23, 227 /232.
Jiang, Y., Joyce, D.C., Macnish, A.J., Jiang, Y.M., 2002a.
Softening response of banana fruit treated with 1-methylcyclopropene to high temperature exposure. Plant Growth
Regul. 36, 7 /11.
Jiang, W., Sheng, Q., Zhou, X., Zhang, M., Liu, X., 2002b.
Regulation of coriander senescence by 1-methylcyclopropene and ethylene. Postharvest Biol. Technol. 26, 339 /345.
Johnson, J.W., 1987. Studies of the carbon (7) hydrogen (6) and
1,3-pentadiyne energy surfaces (cycloheptatetraene, diazocyclopentadiene). PhD Thesis Univ. of Calif. Los Angeles.
Jones, M.L., Kim, E.-S., Newman, S.E., 2001. Role of ethylene
and 1-MCP in flower development and petal abscission in
zonal geraniums. HortScience 36, 1305 /1309.
Kahl, J., Siemens, D.H., Aerts, R.J., Gabler, R., Kuhnemann,
F., Preston, C.A., Baldwin, I.T., 2000. Herbivore-induced
ethylene suppresses a direct defense but not a putative
indirect defense against an adapted herbivore. Planta 210,
336 /342.
Kim, H.O., Hewett, E.W., Lallu, N., 2001. Softening and
ethylene production of kiwifruit reduced with 1-methylcyclopropene. Acta Hort. 553, 167 /170.
Kluge, R.A., Jacomino, A.P., 2002. Shelf life of peaches treated
with 1-methylcyclopropene. Scientia-Agricola 59, 69 /72.
Ku, V.V.V., Wills, R.B.H., 1999. Effect of 1-methylcyclopropene on the storage life of broccoli. Postharvest Biol.
Technol. 17, 127 /132.
Ku, V.V.V., Wills, R.B.H., Ben-Yehoshua, S., 1999. 1-Methylcyclopropene can differentially affect the postharvest life of
strawberries exposed to ethylene. HortScience 34, 119 /120.
Lelievre, J.M., Tichit, L., Dao, P., Fillion, L., Nam, Y.W.,
Pech, J.C., Latche, A., 1997. Effects of chilling on the
expression of ethylene biosynthetic genes in Passe-Crassane
pear (Pyrus communis L. ) fruits. Plant Mol. Biol. 33, 847 /
855.
Lurie, S., Pre-Aymard, C., Ravid, U., Larkov, O., Fallik, E.,
2002. Effect of 1-methylcyclopropene on volatile emission
24
and aroma in Anna apples. J. Agric. Food Chem. 50, 4251 /

4256.
Macnish, A.J., Joyce, D.C., Hofman, P.J., Simons, D.H., 1999.
Involvement of ethylene in postharvest senescence of
Boronia heterophylla flowers. Aust. J. Exp. Agric. 39,
911 /913.
Macnish, A.J., Simons, D.H., Joyce, D.C., Faragher, J.D.,
Hofman, P.J., 2000. Responses of native Australian cut
flowers to treatment with 1-methylcyclopropene and ethylene. HortScience 35, 254 /255.
Magid, R.M., Clarke, T.C., Duncan, C.D., 1971. An efficient
and convenient synthesis of 1-methylcyclopropene. J. Org.
Chem. 36, 1320 /1321.
Manleitner, S.R., Noga, G., Cameron, A.C., 2001. Influence of
1-MCP on russet spotting in lettuce midribs. Acta Hort.
553, 321 /322.
Mathooko, F.M., Tsunashima, Y., Owino, W.Z.O., Kubo, Y.,
Inaba, A., 2001. Regulation of genes encoding ethylene
biosynthetic enzymes in peach (Prunus persica L. ) fruit by
carbon dioxide and 1-methylcyclopropene. Postharvest
Biol. Technol. 21, 265 /281.
Michaeli, R., Philosoph-Hadas, S., Riov, J., Meir, S., 1999.
Chilling-induced leaf abscission of Ixora coccinea plants. I.
Induction by oxidative stress via increased sensitivity to
ethylene. Physiol. Plant. 107, 166 /173.
Mir, N.A., Curell, E., Khan, N., Whitaker, M., Beaudry, R.M.,
2001. Harvest maturity, storage temperature, and 1-MCP
application frequency alter firmness retention and chlorophyll fluorescence of Redchief Delicious apples. J. Am.
Soc. Hort. Sci. 126, 618 /624.
Muller, R., Serek, M., Sisler, E.C., Andersen, A.S., 1997.
Poststorage quality and rooting ability of Epipremnum
pinnatum cuttings after treatment with ethylene action
inhibitors. 72, 445 /452.
Muller, R., Serek, M., Sisler, E.C., Anderson, A.S., 1998.
Ethylene involvement in leaf abscission, chlorosis, and
rooting of Codiaeum variegatum var. pictum (Lodd.) Muell.
Aucubaefolia. Gartenbauwissenschaft 63, 66 /71.
Muller, R., Stummann, B.M., Andersen, A.S., Serek, M., 1999.
Involvement of ABA in postharvest life of miniature potted
roses. Plant Growth Regul. 29, 143 /150.
Mullins, E.D., McCollum, T.G., McDonald, R.E., 2000.
Consequences on ethylene metabolism of inactivating the
ethylene receptor sites in diseased non-climacteric fruit.
Nakano, R., Harima, S., Ogura, E., Inoue, S., Kubo, Y., Inaba,
A., 2001. Involvement of stress-induced ethylene biosynthesis in fruit softening of Saijo persimmon. J. Jpn. Soc.
Hort. Sci. 70, 581 /585.
Nakano, R., Inoue, S., Kubo, Y., Inaba, A., 2002. Water stressinduced ethylene in the calyx triggers autocatalytic ethylene
production and fruit softening in Tonewase persimmon
grown in a heated plastic-house. Postharvest Biol. Technol.
25, 293 /300.
Nakatsuka, A., Shiomi, S., Kubo, Y., Inaba, A., 1997.
Expression and internal feedback regulation of ACC
synthase and ACC oxidase genes in ripening tomato fruit.

Plant Cell Physiol. 38, 1103 /1110.
Nakatsuka, A., Murachi, S., Okunishi, H., Shiomi, S., Nakano,
R., Kubo, Y., Inaba, A., 1998. Differential expression and
internal feedback regulation of 1-aminocyclopropane-1carboxylate synthase, 1-aminocyclopropane-1-carboxylate
oxidase, and ethylene receptor genes in tomato fruit during
development and ripening. Plant Physiol. 118, 1295 /1305.
Newman, J.P., Dodge, L.L., Reid, M.S., 1998. Evaluation of
ethylene inhibitors for postharvest treatment of Gypsophila
paniculata L. HortTechnology 8, 58 /63.
Owino, W.O., Nakano, R., Kubo, Y., Inaba, A., 2002.
Differential regulation of genes encoding ethylene biosynthesis enzymes and ethylene response sensor ortholog during
ripening and in response to wounding in avocado. J. Am.
Soc. Hort. Sci. 127, 520 /527.
Pereira-Netto, A.B., 2001. Effect of inhibitors of ethylene
biosynthesis and signal transduction pathway on the multiplication of in vitro -grown Hancornia speciosa . Plant Cell
Tissue Organ Culture 66, 1 /7.
Pesis, E., Ackerman, M., Ben-Aire, R., Feygenberg, O., Feng,
X., Apelbaum, A., Goren, R., Prusky, D., 2002. Ethylene
involvement in chilling injury symptoms of avocado during
cold storage. Postharvest Biol. Technol. 24, 171 /181.
Picchioni, G.A., Valenzuela-Vzquez, M., Murray, L.W., 2002.
Calcium and 1-methylcyclopropene delay desiccation of
Lupinus havardii cut racemes. HortScience 37, 122 /125.
Porat, R., Halevy, A.H., Serek, M., Borochov, A., 1995a. An
increase in ethylene sensitivity following pollination is the
initial event triggering an increase in ethylene production
and enhanced senescence of Phalaenopsis orchid flowers.
Physiol. Plant. 93, 778 /784.
Porat, R., Shlomo, E., Serek, M., Sisler, E.C., Borochov, A.,
1995b. 1-Methylcyclopropene inhibits ethylene action in cut
phlox flowers. Postharvest Biol. Technol. 6, 313 /319.
Porat, R., Weiss, B., Cohen, L., Daus, A., Goren, R., Droby,
S., 1999. Effects of ethylene and 1-methylcyclopropene on
the postharvest qualities of Shamouti oranges. Postharvest
Biol. Technol. 15, 155 /163.
Porat, R., Feng, X., Huberman, M., Galili, D., Goren, R.,
Goldschmidt, E.E., Feng, X.Q., 2001. Gibberellic acid slows
postharvest degreening of Oroblanco citrus fruit.
Reid, M.S., Wollenweber, B., Serek, M., 2002. Carbon balance
and ethylene in the postharvest life of flowering hibiscus.
Rupasinghe, H.P.V., Almquist, K.C., Paliyath, G., Murr, D.P.,
2001. Cloning of hmg1 and hmg2 cDNAs encoding 3hydroxy-3-methylglutaryl coenzyme A reductase and their
expression and activity in relation to a-farnesene synthesis
in apple. Plant Physiol. Biochem. 39, 933 /947.
Rupasinghe, H.P.V., Murr, D.P., Palyath, G., DeEll, J.R.,
2000a. Suppression of a-farnesene synthesis in Delicious
apples by aminoethoxyvinylglycine (AVG) and 1-methylcyclopropene (1-MCP). Physiol. Mol. Biol. Plants 6, 195 /198.
Rupasinghe, H.P.V., Murr, D.P., Paliyath, G., Skog, L., 2000b.
Inhibitory effect of 1-MCP on ripening and superficial scald
development in McIntosh and Delicious apples. J. Hort.
Sci. Biotech. 75, 271 /276.
Selvarajah, S., Bauchot, A.D., John, P., 2001. Internal browning in cold-stored pineapples is suppressed by a postharvest
application of 1-methylcyclopropene. Postharvest Biol.
Technol. 23, 167 /170.
Serek, M., Prabucki, A., Sisler, E.C., Andersen, A.S., 1998.
Inhibitors of ethylene action affect final quality and rooting
of cuttings before and after storage. HortScience 33, 153 /
155.
Serek, M., Reid, M.S., 2000. Ethylene and postharvest performance of potted kalanchoe. Postharvest Biol. Technol. 18,
43 /48.
Serek, M., Sisler, E.C., 2001. Efficacy of inhibitors of ethylene
binding in improvement of the postharvest characteristics of
potted flowering plants. Postharvest Biol. Technol. 23, 161 /
166.
Serek, M., Reid, M.S., Sisler, E.C., 1994a. A volatile ethylene
inhibitor improves the postharvest life of potted roses. J.
Am. Soc. Hort. Sci. 119, 572 /577.
Serek, M., Sisler, E.C., Reid, M.S., 1994b. Novel gaseous
ethylene binding inhibitor prevents ethylene effects in potted
flowering plants. J. Am. Soc. Hort. Sci. 119, 1230 /1233.
Serek, M., Sisler, E.C., Reid, M.S., 1995a. Effects of 1-MCP on
the vase life and ethylene response of cut flowers. Plant
Serek, M., Sisler, E.C., Reid, M.S., 1995b. 1-Methylcyclopropene, a novel gaseous inhibitor of ethylene action, improves
the life of fruits, cut flowers and potted plants. Acta Hort.
394, 337 /345.
Serek, M., Sisler, E.C., Reid, M.S., 1996. Ethylene and the
postharvest performance of miniature roses. Acta Hort. 424,
145 /149.
Serek, M., Sisler, E.C., Tirosh, T., Mayak, S., 1995c. 1Methylcyclcopropene prevents bud, flower, and leaf abscission of geraldton waxflower. HortScience 30, 1310.
Serek, M., Tamari, G., Sisler, E.C., Borochov, A., 1995d.
Inhibition of ethylene-induced cellular senescence symptoms
by 1-methylcyclopropene, a new inhibitor of ethylene
action. Physiol. Plant. 94, 229 /232.
Sisler, E.C., Blankenship, S.M., May 21, 1996. Methods of
counteracting an ethylene response in plants. U.S. Patent
Number 5,518,988.
Sisler, E.C., Serek, M., 1997. Inhibitors of ethylene responses in
plants at the receptor level: recent developments. Physiol.
Plant. 100, 577 /582.
Sisler, E.C., Serek, M., 1999. Compounds controlling the
ethylene receptor. Bot. Bull. Acad. Sin. 40, 1 /7.
Sisler, E.C., Dupille, E., Serek, M., 1996a. Effect of 1methylcyclopropene and methylenecyclopropene on ethylene binding and ethylene action on cut carnations. Plant
25
Sisler, E.C., Serek, M., Dupille, E., 1996b. Comparison of

cyclopropene, 1-methylcyclopropene, and 3,3-dimethylcyclopropene as ethylene antagonists in plants. Plant Growth
Regul. 18, 164 /174.
Sisler, E.C., Serek, M., Dupille, E., Goren, R., 1999. Inhibition
of ethylene responses by 1-methylcyclopropene and 3methylcyclopropene. Plant Growth Regul. 27, 105 /111.
Sisler, E.C., Serek, M., Roh, K.A., Goren, R., 2001. The effect
of the chemical structure on the antagonism by cyclopropenes of ethylene responses in banana. Plant Growth Regul.
33, 107 /110.
Shiomi, S., Yamamoto, M., Nakamure, R., Inaba, A., 1999.
Expression of ACC synthase and ACC oxidase genes in
melons harvested at different stages of maturity. J. Jpn. Soc.
Hort. Sci. 68, 10 /17.
Skog, L.J., Schaefer, B.H., Smith, P.G., 2001. 1-Methylcyclopropene preserves the firmness of plums during postharvest
storage and ripening. Acta Hort. 553, 171 /172.
Sun, J., Jameson, P.E., Clemens, J., 2000. Stamen abscission
and water balance in Metrosideros flowers. Physiol. Plant.
110, 271 /278.
Tian, M.S., Prakash, S., Elgar, H.J., Young, H., Burmeister,
D.M., Ross, G.S., 2000. Responses of strawberry fruit to 1methylcyclopropene (1-MCP) and ethylene. Plant Growth
Regul. 32, 83 /90.
Watkins, C.B., 2002. Ethylene synthesis, mode of action,
consequences and control. In: Knee, M. (Ed.), Fruit Quality
and its Biological Basis. Sheffield Academic Press, pp. 180 /
224.
Watkins, C.B., Nock, J.F., Whitaker, B.D., 2000. Responses of
early, mid and late season apple cultivars to postharvest
application of 1-methylcyclopropene (1-MCP) under air and
controlled atmosphere storage conditions. Postharvest Biol.
Technol. 19, 17 /32.
Wills, R.B.H., Ku, V.V.V., 2002. Use of 1-MCP to extend the
time to ripen of green tomatoes and postharvest life of ripe
tomatoes. Postharvest Biol. Technol. 26, 85 /90.
Wills, R.B.H., Ku, V.V.V., Warton, M.A., 2002. Use of 1methylcyclopropene to extend the postharvest life of lettuce.
J. Sci. Food Agric. 82, 1253 /1255.
Winz, R.A., Baldwin, I.T., 2001. Molecular interactions
between the specialist herbivore Manduca sexta (lepidoptera, Sphingidae) and its natural host Nicotiana attenuata .
IV. Insect-induced ethylene reduces jasmonate-induced
nicotine accumulation by regulating putrescine N -methyltransferase transcripts. Plant Physiol. 125, 2189 /2202.
Zhong, G.Y., Huberman, M., Feng, X., Sisler, E.C., Holland,
D., Goren, R., Zhong, G.Y., Feng, X.Q., 2001. Effect of 1methylcyclopropene on ethylene-induced abscission in citrus. Physiol. Plant. 113, 134 /141.

1-MCP Review: Effects of Ethylene Inhibitor on Fruits and Vegetables

Transféré par

Informations du document

Titre original

Copyright

Formats disponibles

Partager ce document

Partager ou intégrer le document

Options de partage

Avez-vous trouvé ce document utile ?

Ce contenu est-il inapproprié ?

Droits d'auteur :

Formats disponibles

1-MCP Review: Effects of Ethylene Inhibitor on Fruits and Vegetables

Transféré par

Droits d'auteur :

Formats disponibles

Postharvest Biology and Technology 28 (2003) 1 /25

* Corresponding author. Tel.: /1-919-515-1224; fax: /1919-515-7747.

MCP in research programs promises to advance

ethylene. Compared with ethylene, 1-MCP is

Diffusion of 1-MCP out of plant material is

3. Conditions for use

occurred at both temperatures (Ku and Wills,

Ananas comosus (pineapple)

Selvarajah et al., 2001

Annona squamosa /Annona cheimola (custard apple)

Controlled internal browning; slowed ascorbic acid, ethylene, and

Hall et al., 2000

Brassica oleracea (broccoli)

B. rapa var. chinensis (pak choy)

B. juncea var. foliosa (Chinese mus- 12 ml l 1

B. rapa var. nipposinica (mibuna and 12 ml l 1

Allowed normal seedling growth

Able et al., in press

Carica papaya (papaya)

Chrysanthemum coronarium (garland chrysanthemum)

Cucumis melo (melon) plants

Citrullus lanatus (watermelon)

Delayed yellowing, increased shelf

Citrus paradisi (grapefruit)

Citrus spp. (orange)

B. rapa var. parachinensis (choy

Hofman et al., 2001

Fan and Mattheis, 2000; Ku and

Able et al., in press

Able et al., in press

Able et al., in press

Hofman et al., 2001

Mao and Huber (personal communication)

Nicotiana attenuata (tobacco) plants 0.04 /0.43 g of 0.43%

Fan and Mattheis, 2000

Nakano et al., 2001, 2002

Ciardi et al., 2001

Watkins et al., 2000; DeEll et al.,

Prevented acidity loss, respiration

Persea americana (avocado)

50 /300 nl l 1; 0.45, 25

Jeong et al., 2001; Hofman et al.,

Pisum sativa seedlings (pea)

Inhibited ripening, maintained

Prunus armeniaca (apricot)

P. persica (peach and nectarine)

20 nl l 1; 0.1 ml l 1; 0.5 20 /24

Pyrus communis (pear)

Vigna radiata leaves (mung bean)

Sisler and Serek, 1997; Sisler et al.,

Treatment temperature (8C)

Begonia /elatior Najada and

Begonia /tuberhybrida Non-Stop 5 or 20 nl l 1

Prevented decrease in vase life due to exogenous ethylene

Campanula carpatica Dark Blue

C. carpatica Dark Blue and Blue

C. medium Champion Pink

Treatment temperature (8C)

Dendranthema grandiflorum Coral

0.6, 1.7, 3.3, 5.8 or 20

D. caryophyllus White Sim or

Not specified, prob- 12

Grevillea Kay Williams

* Corresponding author. Tel.: /1-919-515-1224; fax: /1919-515-7747.

Annona squamosa /Annona cheimola (custard apple)

B. juncea var. foliosa (Chinese mus- 12 ml l 1

B. rapa var. nipposinica (mibuna and 12 ml l 1

50 /300 nl l 1; 0.45, 25

20 nl l 1; 0.1 ml l 1; 0.5 20 /24

Begonia /elatior Najada and

Begonia /tuberhybrida Non-Stop 5 or 20 nl l 1

Hibiscus rosa-sinensis numerous cul- 200 nl l 1

Lilium hybrids Cordelia and Elite 150 nl l 1

l1 produced 70% volatile inhibition and 95%

choe in the presence of exogenous (3 ml l 1)