Plant Growth Regulation 32: 83–90, 2000.

© 2000 Kluwer Academic Publishers. Printed in the Netherlands.

83

Responses of strawberry fruit to 1-Methylcyclopropene (1-MCP) and

ethylene

M.S. Tian, S. Prakash, H.J. Elgar, H. Young, D.M. Burmeister & G.S. Ross
The Horticulture and Food Research Institute of New Zealand Ltd., Mt Albert Research Centre, Private Bag 92169,
Auckland, New Zealand

Received 21 September 1999; accepted in revised form 21 January 2000

Key words: ethylene production, Fragaria ananassa, ionic conductivity, 1-MCP, peroxidase, respiration, ripening
index, strawberry

Abstract
1-Methylcyclopropene (1-MCP), a competitive inhibitor of ethylene action, binds to the ethylene receptor to
regulate tissue responses to ethylene. In this work, we investigated the effects of 1-MCP and exogenous ethyl-
ene on ripening, respiration rate, ionic conductivity and peroxidase activity in strawberry fruit. Strawberry fruit
can ripen without exogenous ethylene treatment, but exogenous ethylene induces secondary ripening processes.
Results indicated that stimulation of respiration by ethylene was dose-dependent. Fruit colour development
and softening were slightly accelerated by ethylene, but changes in soluble solid content were not. 1-MCP
may/may not affect the respiratory rise induced by exogenous ethylene dependent on fruit maturity. Cyclo-
heximide (CHI) reduced the ethylene-induced respiratory increase. Combinations of 1-MCP and CHI reduced
respiration more than CHI alone. 1-MCP and CHI did not influence the primary respiratory change in non
ethylene-treated fruit. This indicates that ethylene induced respiratory increase may involve an ethylene receptor
in early harvested fruit, but not in later harvested fruit. Exogenous ethylene stimulated respiration by regulating
new respiratory enzyme(s) synthesis in strawberry fruit. Ethylene induced an ionic leakage increase, and this
was positively correlated to fruit water loss and peroxidase activity. These results suggest that non-climacteric
fruit, such as strawberry, may have different ethylene receptor(s) and/or ethylene receptor(s) may have different
regulatory functions. It may be the secondary effect of ethylene to stimulate respiration in strawberry. Non-
climacteric fruit ripening may be related to the development of active oxygen species (AOS) induced by postharvest
stress.

Abbreviations: ACC – 1-aminocyclopropane-1-carboxylic acid; AOS – active oxygen species; CHI – cyclohexim-
ide; 1-MCP – 1-methylcyclopentene

1. Introduction receptor. Experiments with yeast cultures expressing

Ripening of climacteric fruit is accompanied by
autocatalytic increases in ethylene production and an
increase in the rate of respiration of the fruit. Exo-
genous application of ethylene can also induce these
irreversible events [1, 24]. Along with many other
ripening-related changes, these events are presumed
to be mediated by an ethylene signal transduction
cascade, initiated by the binding of ethylene to a
an Arabidopsis ethylene receptor have demonstrated
that ethylene binds to this receptor, which may be
positioned in the membrane as a dimer [2, 13]. Homo-
logues of this gene have now been identified in a
number of climacteric fruit, such as tomato [20], apple
[17] and muskmelon [7].
In contrast with climacteric fruit, ripening of non-
climacteric fruit does not involve an autocatalytic
ethylene rise. In these fruit, ethylene does stimulate
84
the respiration rate [6, 24], although the effect is
reversible and is dependent on continuously elevated
levels of endogenous or exogenous ethylene. Little
work has been done to investigate the mechanism of
these different respiratory responses to ethylene in the
two types of fruit. Although a number of ripening-
related genes from strawberry have been reported
[8, 21], to our knowledge no ethylene receptor has
been isolated from any non-climacteric fruit. An
ethylene signal transduction pathway is in operation
however, as evidenced by the effects on respiration
and other work showing ethylene-enhanced straw-
berry fruit softening and reduced storage life [4, 22,
23].
1-Methylcyclopropene (1-MCP) is a competitive
inhibitor of ethylene action which binds to the ethyl-
ene receptor to regulate tissue responses to ethylene
[9, 14]. 1-MCP has been used as a tool to investig-
ate ethylene action and tissue responses to ethylene
during climacteric fruit ripening and flower senes-
cence [15]. Application of 1-MCP delays ripening
of banana, apple and tomato, and flower senescence
[10, 11, 15, 16], presumably via its blocking effect
on the ethylene signal transduction pathway. A pho-
tolysed derivative of DACP, an other ethylene action
inhibitor used in earlier trials of this nature [18, 19].
In our laboratory, we studied the effects of DACP
on ethylene-induced respiratory rises in strawberry
[19]. This work indicates that ethylene biosynthesis in
nonclimacteric strawberry fruit may be regulated by
ethylene receptor(s) with inhibition of ethylene pro-
duction. DACP may release this inhibitory effect, and
resulting in increased ethylene production. Respira-
tion in strawberry fruit may not be regulated by an
ethylene receptor at most stages of maturity. Present
work did not investigate differences of the responses
to 1-MCP and ethylene in fruit harvested at early and
later seasons. It is possible that they have different
sensitivities and responses to ethylene.
Here we report further investigations on the role
of ethylene in strawberry ripening and respiration. We
used 1-MCP as a tool to investigate whether exogen-
ous ethylene binds to the receptor to induce the res-
piratory rise and to affect ripening in strawberry fruit.
We also used cycloheximide to determine whether the
ethylene effects are the result of new protein syn-
thesis, and measured changes in ionic conductivity
and peroxidase activity in ethylene-treated strawberry
fruit as markers for the progression of ripening and as
indicators of tissue stress.
2. Materials and methods
2.1 Plant material
Strawberry fruit (Fragaria ananassa Duch. cv. Pajaro)
were harvested at the pink stage during the summer
seasons of 1996 to 1998 from a local grower property
in Auckland, New Zealand. Fruit were harvested only
in the late season (November-December) in 1996 and
1997, and both early (October) and late during 1998.
Fruit were transported to the Mt. Albert Research
Centre. To compare rates of fresh weight loss and
relative ionic conductivity for fruit of various harvest
maturities, fruit were harvested at the green, white,
pink and red stages.
2.2 1-MCP treatments
Fruit were placed in 10 L sealed jars, and treated with
1-MCP (2 µl L−1 ) for 18 h before being vented to air.
2.3 Inhibitors of protein synthesis
After 1-MCP treatment, fruit were dipped in a 0.4 M
mannitol solution containing cycloheximide (50 µg
ml−1 ) for 0.5 h, and then dried on paper.
2.4 Ethylene/propylene treatments
In an experiment investigating ethylene dose effect
on respiration rate, each replicate of four fruits was
placed in a 600 ml jar (n = 4), and flushed with an
air flow of 50 ml min−1 containing various concen-
trations of exogenous ethylene, at levels of 0 (air),
0.5, 1, 5, 20, 50 or 100 µl L−1 . In other experi-
ments, fruit were treated with 50 µl L−1 ethylene at
a continuous flow rate of 50 ml min−1 (with air as the
control).
2.5 Colour, firmness and soluble solids content
Fruit skin colour was measured with a Minolta CR300
reflectance Chroma Meter (Minolta Camera Co. Ltd.,
Osaka, Japan). The ‘a’ and ‘b’ values on the ‘Lab’
mode were used as indices of colour changes from
pink to red. Colour was measured at two opposite
points around the equator on each fruit. Fruit firmness
(N) was assessed using an Imada penetrometer (PSM-
10, Imada Co. Ltd., Toyohashi, Japan) fitted with a flat
round 13.16 mm diameter head. The head was pushed
into the strawberry flesh through the skin to the depth

of the head (5 mm). Soluble solids content (%) was
measured using a digital 0–20% refractometer (Atago
model PR-1; Atago Co. Ltd. Tokyo).
2.6 Respiration and ethylene production
Fruit were sealed in five 600 ml jars (four fruit per
jar) for 1 h. The respiration (ml.kg−1.h−1 ) and ethyl-
ene production (µl.kg−1.h−1 ) rates were measured by
withdrawing two 1 ml samples from the head space.
These were analysed immediately for CO2 using an
infra red detection cell (Servomex 01514/701 infra-
red transducer; Servomex PLC, Crowborough, East
Sussex, UK), and for ethylene using a gas chroma-
tograph (Hewlett-Packard HP5890 Series II gas chro-
matograph; 1.5 m × 6 mm alumina F1 column, injec-
tion temperature 160 ◦ C, detector temperature 200 ◦ C,
column temperature 130 ◦ C).
2.7 Ionic leakage
Six fruit discs were placed in a 45 ml test tube (n = 4)
containing 20 ml distilled water and incubated at 25 ◦ C
for 0.5 h. Thereafter, fruit tissue was homogenized
twice at top speed for 10 sec. The electronic con-
ductivity was determined with a conductivity meter
before (as initial conductivity) and after (as final con-
ductivity) homogenisation. The ionic leakage of the
tissue was calculated as: ionic leakage (%) = initial
conductivity/(final conductivity × fresh weight).
2.8 Peroxidase activity
Samples of fresh tissue (10 g) were ground with 20 ml
of 0.2 M sodium phosphate buffer (pH 6.5) con-
taining 4% (w/v) insoluble polyvinlpolypyrrolidone
(PVPP) and 1% Triton X-100 in an ultrahomogen-
izer at high speed for 3 min with stop intervals each
30 sec. The suspension was centrifuged at 8000 g at
4 ◦ C for 15 min, and the supernatant was used to assay
enzymatic activity. Enzymatic activity was assayed by
adding 200 µl crude enzymatic extract to the 2.7 ml
reaction mixture containing 0.05 M sodium phosphate
buffer (pH 6.5), 200 µl 1% ρ-phenylenediamine as
a H-donor, and 100 µl 1.5% (w/v) hydrogen perox-
ide as an oxidant. Enzymatic activity was determined
by the increase in absorbance at 485 nm (A485) after
incubation for 3 min at 22 ◦ C. The enzymatic activity
unit was defined as the change in absorbance.g−1 fresh
weight.min−1.
85
Figure 1. Respiration rate (ml.kg−1 .h−1 ) of strawberry fruit har-
vested at the pink stage and treated with various concentrations of
ethylene.
3. Results
3.1 Ethylene-induced respiration rise
Exogenous ethylene stimulated respiration in pink
strawberry (Figure 1). The increase of respiratory rate
depended on ethylene concentrations and treatment
duration. Fruit respiration was enhanced by treatment
with 20 µl L−1 ethylene for one day. It was signific-
antly increased in fruit treated with ethylene at less
than 1 µl L−1 for 2 days. The higher the ethylene
concentration, the greater the increase in respiration
rate.
3.2 Ethylene- or propylene-induced respiration rise
in 1-MCP treated fruit
Continuous ethylene (100 µl L−1 ) treatment stimu-
lated respiration in early-harvested strawberry fruit
within 2 days (Figure 2A). Thereafter, respiration
decreased to a similar level as the control fruit (air
treatment) by Day 3. 1-MCP treatment inhibited the
ethylene-induced respiration rise in early-harvested
fruit (Figure 2C). However, in late-harvested fruit,
1-MCP neither significantly changed the respiratory
rate, nor influenced the ethylene-induced respiratory
rise (Figure 2E & G).
3.3 Effects of cycloheximide on respiration in
strawberry fruit at the pink stage
Cycloheximide (CHI), a translation inhibitor, was used
to investigate whether the ethylene-induced respirat-
ory rise in early-harvested fruit was regulated at the
translation level. Dipping fruit in 1 mM CHI for
86

Figure 2. Respiration rate (ml.kg−1 .h−1 ) of (A–D) early- and (E–H) late-harvested pink strawberry fruit in ethylene (100 µl L−1 ) after being
treated with 1-MCP (2 µl L−1 ) and CHI(1 mM). A and E, –MC, –CHI; B and F, –MCP, +CHI; C and G, +MCP, –CHI; D and H, +MCP, +CHI.

Table 1. Colour (a/b ratio), firmness (Kg) and soluble solids

0.5 h before ethylene treatment did not change the content (%) of strawberry fruit harvested at the pink stage and
respiratory rate in the control fruit, but significantly continuously treated with various concentrations of ethylene
reduced the ethylene-induced respiratory rise at 1 and between 0 and 100 µl L−1
2 days after treatments (Figure 2B & F). Combina-
tions of 1-MCP and CHI inhibited the respiration rise Days after Ethylene Colour Firmness Soluble solids
treatment (µl L−1 ) (a/b ratio) (Kg) content (%)
at Day 2 more than CHI treatment alone (Figure 2D
& H). Inhibition of respiration by CHI in the con- 0 At harvest 0.67c1 2.37a 6.28a
trol fruit (air) and 1-MCP-treated fruit was greater in 3 0 1.53b 1.99b 6.46a
early-harvested fruit than late-harvested fruit. 0.5 1.50b 1.78c 6.46a
1 1.59b 1.76c 6.24a
3.4 Colour, firmness and soluble solids content in 20 1.54b 1.80c 6.24a
40 1.72a 1.75c 6.42a
1-MCP, CHI and ethylene-treated fruit
100 1.62ab 1.72c 6.44a

The a/b ratio of skin colour and soluble solids content
increased, and firmness decreased significantly after
harvest in the control fruit (air) at 20 ◦ C (Tables 1 &
2). Exogenous ethylene appeared to accelerate colour
development and softening, without affecting soluble
solids content. The enhancement of colour develop-
ment and softening depended on ethylene concentra-
tion and duration of exposure (Table 1). After treat-
ment for three days, the a/b ratios were significantly
greater in fruit treated with higher concentrations of
ethylene (>40 µl L−1 ) than with lower concentrations
(Table 1). This was mainly due to the increase in the
1 Different letters represented significant differences at the 5%
level (n = 16).
‘a’ value for these fruit. The decrease in fruit firmness
was greater in fruit treated with ethylene at concen-
trations over 0.5 µl L−1 . No significant difference in
soluble solid content was evident in fruit treated with
different concentrations of ethylene. Again, ethylene
at 50 µl L−1 significantly enhanced the a/b ratio and
accelerated softening in the control fruit (Table 2),
but did not change soluble solids content. 1-MCP,
 87
Table 2. Colour (a/b ratio), firmness (Kg) and soluble solids content (%) in strawberry
fruit harvested at the pink stage and treated with 1-MCP (2 µl L−1 ), cycloheximide
(CHI, 1 mM) and ethylene (50 µl L−1 )

Days after Treatment Colour Firmness Soluble solids

harvest (a/b ratio) (Kg) content (%)

0 Air –MCP/–CHI 0.48d1 2.97a 5.85b

4 Air –MCP/–CHI 1.43b 2.54b 6.18a
–MCP/+CHI 1.08c 2.32b 5.84b
+MCP/–CHI 1.11c 1.89c 5.85b
+MCP/+CHI 1.19bc 2.32b 5.63b
4 Ethylene –MCP/–CHI 1.64a 1.81c 6.11a
–MCP/+CHI 1.32b 2.31b 5.98ab
+MCP/–CHI 1.35b 2.00bc 5.50b
+MCP/+CHI 1.21bc 2.41b 5.60b

1 Different letters represented significant differences at the 5% level (n = 16).

ene (100 µl L−1 ) stimulated ionic leakage within 2

days of treatment. Thereafter, it reached the same level
as in control fruit after three days.

3.7 Peroxidase (PEO) activity

PEO activity decreased after harvest in control fruit

(Table 3). Treatment with ethylene (100 µl L−1 ) main-
tained PEO activity during the first three days after
harvest.

Figure 3. Changes in relative ion conductivity (%) in strawberry 4. Discussion

fruit at various ripening stages.

Differences between ethylene-induced respiration

CHI, or a combination of these, depressed ethylene-
induced colour development and softening, but did not
influence soluble solids content.
3.5 Ionic leakage and fresh weight loss
Fresh weight of strawberry fruit harvested at the vari-
ous stages of maturity (green, white, pink and red)
decreased significantly one day after harvest (data was
not shown). Ionic leakage of fruit tissue increased with
maturity development and ripening (Figure 3). The
greater the maturity, the greater the increase in ionic
leakage during postharvest life.
3.6 Changes in ionic leakage in ethylene-treated pink
fruit
Ionic leakage increased continuously in strawberry
fruit for three days following harvest (Figure 4). Ethyl-
rise in climacteric and non-climacteric fruits have
been reported [1]. Ethylene irreversibly induces the
autocatalytic respiration rise in climacteric fruit, but
it increases the respiration rate reversibly in non-
climacteric fruit. The respiration rise induced by ethyl-
ene in non-climacteric fruit is dependent on ethylene
concentration suggesting that ethylene response of the
tissue may be linked to respiration [24]. It is not clear
why different respiration responses to ethylene occur
in both types of fruit.
In this work, different respiration responses to
ethylene were found in early- and late-harvested
strawberry fruit. 1-MCP inhibited, or did not affect,
the respiratory rise induced by exogenous ethylene
in early- and late-harvested strawberry fruit, respect-
ively. Colour development and fruit softening required
different treatment duration and concentrations of exo-
genous ethylene. These results may be explained by
the fact that fruit at different ripening stages may have
88
Figure 4. Changes in ion conductivity (%) in ethylene -treated (100 µl L−1 ) strawberry fruit harvested at the pink stage.
Table 3. Peroxidase activity (1A485 .g−1 FW.min−1 ) in
propylene–treated (0.5%) pink strawberry fruit (n = 4)
Days at 20 ◦ C Air Ethylene
1 1.66 1.66
2 1.47 1.59
3 0.47 1.47
different sensitivity thresholds to exogenous ethyl-
ene. Also, different ripening processes such as colour,
softening and soluble solids may all be regulated by
ethylene, but may have different threshold levels.
In late-harvested fruit, 1-MCP did not affect the
ethylene-induced respiration increase. This result can
be explained by the fact that ethylene may not neces-
sarily bind to the receptor to enhance respiration of
strawberry fruit. Similar results were observed with
another ethylene action inhibitor, DACP, which stim-
ulated ethylene biosynthesis without changing res-
piration rate in late-harvested strawberry fruit [19].
Different results were detected for DACP-inhibited
ethylene biosynthesis and respiration in climacteric
fruit, such as apple and tomato [16, 18]. A pos-
sible explanation is that there may either be different
ethylene receptor(s), or the same receptor has differ-
ent functions, in climacteric and non-climacteric fruits
[24].
1-MCP can depress respiration and delay ripening
of climacteric fruit [10]. The decreases in respiration
and ethylene production were not recovered by applic-
ation of exogenous ethylene [16]. It was hypothesised
that ethylene may bind to metal in the reactive or
catalytic centers of enzymes to stimulate their activ-
ities [15]. Ethylene binds to the receptor, then diffuses
rapidly from the receptor, whereas 1-MCP and other
inhibitors remain bound for a long period. While
inhibitors are bound, ethylene cannot bind [15]. There-
fore, responses to ethylene in climacteric fruit are
delayed.
1-MCP stimulated ethylene production, but did not
induce a respiratory rise, in late-harvested strawberry
fruit [19]. The ethylene-induced respiratory rise was
dose-dependent. Thus, this result can be explained
that the respiratory rise in strawberry fruit required a
greater concentration of ethylene than that induced by
1-MCP.
Cycloheximide (CHI), a protein synthesis inhib-
itor, depressed ethylene-induced respiration in straw-
berry. This indicated that de novo synthesis of respirat-
ory enzymes may be required for the ethylene-induced
respiratory rise. Combination of 1-MCP and CHI
reduced the ethylene-induced respiratory rise more
than CHI alone. Thus, there is an interaction between
1-MCP and ethylene in the regulation of synthesis
of respiratory enzymes, but the mechanism of this
interaction still remains unclear.
It has been reported that 1-MCP inhibits ethylene-
induced cellular senescence symptoms in ethylene-
sensitive flowers such as petunia [12]. Ionic con-
ductivity of strawberry fruit increased with stages of

maturity and ripening. This increase was correlated
with loss of fresh weight, and was also stimulated by
exogenous ethylene.
Free radicals and other active derivatives of oxy-
gen inactivate enzymes and damage important cellular
components [5]. To counteract the toxicity of act-
ive oxygen species, a highly efficient anti-oxidative
defence system, composed of non-enzymatic and
enzymatic constituents, is present in all plant cells.
The enzymatic antioxidative components include
superoxide dismutase and peroxidase. Two peroxidase
isoenzymes (58.1 and 65.5 kD) have been detected
in strawberry fruit [3]. Peroxidase activity, found
primarily in a membrane-bound form, decreased dur-
ing strawberry ripening. Present results have demon-
strated that peroxidase activity in pink strawberry fruit
was low at harvest and decreased during fruit ripening.
However, treatment with exogenous ethylene main-
tained peroxidase activity. It is possible that active
oxygen species, induced by postharvest stress such as
water loss, may be involved in non-climacteric fruit
ripening.
In this work, results suggest that non-climacteric
fruit such as strawberry may have different ethylene
receptor(s), and/or ethylene receptor(s) may have dif-
ferent regulatory functions. Exogenous ethylene stim-
ulated respiration by regulating de novo respiratory
enzyme synthesis in strawberry fruit. However, it may
not be necessary for ethylene to bind to the receptor
to stimulate respiration in strawberry. Non-climacteric
fruit ripening may be related to the development of
active oxygen species induced by postharvest stress.
Acknowledgements
The authors wish to thank Drs. IB Ferguson and
N Lallu from the Horticulture and Food Research
Institute of New Zealand for helpful discussions.
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