The Use of 1-Methylcyclopropene (1-MCP) On Fruits and Vegetables

Biotechnology Advances 24 (2006) 389 – 409
www.elsevier.com/locate/biotechadv
Research review paper

The use of 1-methylcyclopropene (1-MCP) on fruits and vegetables
Chris B. Watkins ∗
Department of Horticulture, Cornell University, Ithaca, NY 14853, USA
Available online 10 March 2006
Abstract
The recent availability of the inhibitor of ethylene perception, 1-methylcyclopropene (1-MCP), has resulted in an explosion of
research on its effects on fruits and vegetables, both as a tool to further investigate the role of ethylene in ripening and senescence,
and as a commercial technology to improve maintenance of product quality. The commercialization of 1-MCP was followed by
rapid adoption by many apple industries around the world, and strengths and weaknesses of the new technology have been
identified. However, use of 1-MCP remains limited for other products, and therefore it is still necessary to speculate on its
commercial potential for most fruits and vegetables. In this review, the effects of 1-MCP on fruits and vegetables are considered
from two aspects. First, a selected number of fruit (apple, avocado, banana, pear, peaches and nectarines, plums and tomato) are
used to illustrate the range of responses to 1-MCP, and indicate possible benefits and limitations for commercialization of 1-MCP-
based technology. Second, an outline of general physiological and biochemical responses of fruits and vegetables to the chemical
is provided to illustrate the potential for use of 1-MCP to better understand the role of ethylene in ripening and senescence
processes.
© 2006 Elsevier Inc. All rights reserved.
Keywords: Fruit; Vegetables; Ripening; Senescence; Ethylene; Softening; Respiration; Flavor; Antioxidants
Contents
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 390
2. Responses of selected fruit and vegetables to 1-MCP . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 392
2.1. Apple [Malus sylvestris (L.) Mill. var. domestica (Borkh.) Mansf.] . . . . . . . . . . . . . . . . . . . . . . 392
2.1.1. Ripening physiology and quality. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 392
2.1.2. Commercial application of 1-MCP. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 393
2.2. Avocado. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 394
2.2.2. Factors affecting commercial application of 1-MCP . . . . . . . . . . . . . . . . . . . . . . . . . . . 394
2.3. Banana (Musa sp., AAA group, Cavendish subgroup) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 394
2.4. Pear (Pyrus communis L.). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 395
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doi:10.1016/j.biotechadv.2006.01.005
390 C.B. Watkins / Biotechnology Advances 24 (2006) 389–409
2.5. Peach and nectarine (Prunus persica L.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 396

2.5.1. Ripening physiology and quality . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 396
2.5.2. Factors affecting commercial application of 1-MCP . . . . . . . . . . . . . . . . . . . . . . . . . . 396
2.6. Plum (Prunus domestica L. and Prunus salicina L.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 396
2.7. Tomato (Solanum esculentum Mill) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 397
3. Physiological and biochemical responses of fruits and vegetables to 1-MCP . . . . . . . . . . . . . . . . . . . . . 398
3.1. Ethylene metabolism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 398
3.2. Respiration rate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 400
3.3. Pigment metabolism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 400
3.4. Cell wall metabolism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 400
3.5. Volatile compound metabolism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 401
3.6. Nutritional quality . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 402
3.7. Physiological storage disorders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 402
3.7.1. Apple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 402
3.7.2. Other fruits and vegetables . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 403
3.7.3. A physiological basis for disorder development in response to 1-MCP . . . . . . . . . . . . . . . 403
3.8. Pathological storage disorders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 404
4. Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 404
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 405
1. Introduction ones, as well as senescence of vegetative tissues, has

emerged with the discovery and commercialization of
Ethylene is one of several plant growth regulators that the inhibitor of ethylene perception, 1-methylcyclo-
affect growth and developmental processes including propene (1-MCP).
ripening and senescence (Abeles et al., 1992). It is a 1-MCP is thought to interact with ethylene
simple hydrocarbon that can diffuse into and out of plant receptors and thereby prevent ethylene-dependent
tissues from both endogenous and exogenous (non-bio- responses (Sisler and Blankenship, 1996; Sisler and
logical and biological) sources (Saltveit, 1999) and has Serek, 1997, 2003). The use of cyclopropenes to
been the subject of extensive research on its biosynthesis inhibit ethylene action was patented by Sisler and
and action (Lelievre et al., 1998; Saltveit, 1999; Blankenship (1996). A commercial breakthrough in 1-
Giovannoni, 2001; Watkins, 2002; Adams-Phillips et MCP application technology resulted from the formu-
al., 2004). lation of 1-MCP as a stable powder in which it is
Ethylene can profoundly affect quality of harvested complexed with γ-cyclodextrin, so that 1-MCP is
products. These effects can be beneficial or deleterious easily released as a gas when the powder is dissolved
depending on the product, its ripening stage, and its in water. 1-MCP was approved by the Environmental
desired use (Saltveit, 1999). Endogenous ethylene Protection Agency (EPA) in 1999 for use on
production is an essential part of ripening of ornamentals, and was marketed as EthylBloc® by
climacteric fruit and probably acts as rheostat for Floralife, Inc. (Walterboro, SC). AgroFresh, Inc., a
ethylene-dependent processes (Theologis, 1992). Ex- subsidiary of Rohm and Haas (Springhouse, PA),
ogenous ethylene application is routinely used to subsequently developed 1-MCP under the trade name
initiate uniform ripening for fruit such as banana. Most SmartFresh™ and have global use rights for edible
commonly, however, commercial strategies for horti- horticultural products. 1-MCP has a non-toxic mode of
cultural products are based on avoiding exposure to action, negligible residue and is active at very low
ethylene and/or attempting to minimize ethylene concentrations (E.P.A., 2002), and by 2005 food use
production and action during ripening, harvest, registration for the chemical had been obtained in
storage, transport and handling by temperature and Argentina, Australia, Austria, Brazil, Canada, Chile,
atmosphere control (Watkins, 2002). An exciting new Costa Rica, France, Guatemala and Honduras, Israel,
strategy for controlling ethylene production and thus Mexico, the Netherlands, New Zealand, South Africa,
ripening and senescence of fruit, especially climacteric Switzerland, Turkey, UK, and the US. Registered
Table 1
Climacteric and non-climacteric fruit and vegetables for which responses to 1-MCP have been investigated from references available at Watkins and Miller (2005a)
Fruit (climacteric) Fruit (non-climacteric) Vegetables
Apple [Malus sylvestris (L) Mill. var. Melon (Cucumis melo L.) Cherry (Prunus avium L.) Broccoli (Brassica oleracea L.)
C.B. Watkins / Biotechnology Advances 24 (2006) 389–409

domestica (Borkh.) Mansf.]
Apricot (Prunus armeniaca L.) Mountain papaya (Vasconcellea pubescens) Clementine mandarin (Citrus reticulata L.) Carrot (Daucus carota L.)
Avocado (Persea americana Mill.) Nectarine (Prunus persica Lindl.) Cucumber (Cucumis sativus L.) Chinese cabbage (Brassica campestris
L. spp. pekinensis (Lour) Olsson)
Banana (Musa L.) Papaya (Carica papaya L.) Grape (Vitis vinifera L.) Chinese mustard (Brassica juncea var. foliosa)
Blueberry, highbush (Vaccinium Peach (Prunus persica L. Batsch) Grapefuit (Citrus paradisi Macf.) Choysum (Brassica rapa var. parachinensis)
corymbosum L.)
Chinese bayberry (Myrica rubra Pear (Pyrus communis L.) Lime (Citrus latifolia Tanaka) Chrysanthemum, garland (Chrysanthemum coronarium)
Siebold and Zuccarni)
Chinese jujube (Zizyphus jujube M.) Pear (Pyrus pyrifolia Nakai) Orange (Citrus sinensis L. Osbeck) Coriander (Coriandrum sativum L.)
Custard apple (Annona squamosa L.) Persimmon (Diospyros khaki L.) Pepper (Capsicum frutescens L.) Lettuce (Lactuca sativa L.)
Figs (Ficus carica L.) Plum (Prunus salicina L.; Pineapple (Ananas comosus L.) Mibuna (Brassica rapa var. nipposinica)
Prunus x domestica L.)
Guava (Psidium guajava L.) Tomato (Solanum esculentum Mill) Strawberry (Fragaria × ananassa Duch.) Mizuna (Brassica rapa var. nipposinica)
Kiwifruit (Actinidia deliciosa Watermelon (Citrullus lanatus) Pak choy (Brassica rapa)
(A. Chev) C.F. Liang et A.R.
Ferguson var. deliciosa
Lychee (Litchi chinensis) Parsley (Petroselinum crispum Mill.)
Mamey sapote (Pouteria sapote Potato (Solanum tuberosum)
(Jacq.) H.E. Moore and Stearn)
Mango (Mangifera indica L.) Tatsoi (Brassica rapa var. rosularis)
391
crops, which are specific to countries, include apple, 2. Responses of selected fruit and vegetables to 1-MCP
apricot, avocado, kiwifruit, mango, melon, nectarine,
papaya, peach, pear, pepper, persimmon, pineapple, 2.1. Apple [Malus sylvestris (L.) Mill. var. domestica
plantain, plum, squash, tomatoes and tulip bulbs. (Borkh.) Mansf.]
Registration for use on various fruit and vegetables is
expected soon for other countries. 2.1.1. Ripening physiology and quality
The impact of 1-MCP on postharvest science and 1-MCP dramatically inhibits ripening of apple fruit.
technology has been two-fold. First, it provides the The increases in ethylene production and internal eth-
potential to maintain fruit and vegetable quality after ylene concentrations (IECs) associated with the climac-
harvest. Second, 1-MCP provides a powerful tool to teric ripening stage are prevented or delayed by 1-MCP
gain insight into the fundamental processes that are treatment, the extent of inhibition being related to cul-
involved in ripening and senescence. A large lite- tivar, storage type and length of storage (Fan et al.,
rature on the responses of both whole and fresh cut 1999; Fan and Mattheis, 1999a; Rupasinghe et al., 2000;
fruit and vegetables (as well as for ornamental Watkins et al., 2000; Dauny and Joyce, 2002; Jiang and
products) to 1-MCP is developing, with 40 papers Joyce, 2002; Pre-Aymard et al., 2003; Saftner et al.,
published in 2004 and 73 papers in 2003, compared 2003; Defilippi et al., 2004; Arquiza et al., 2005; Bai et
with 36 in 2002, 19 in 2000, and a total of 16 published al., 2005; Kondo et al., 2005; Mattheis et al., 2005;
by 1998 (Watkins and Miller, 2005a). Recent reviews Moran and McManus, 2005; Pechous et al., 2005;
on the effects of 1-MCP on horticultural products Toivonen and Lu, 2005; Watkins and Nock, 2005).
include Blankenship and Dole (2003), Sisler and Serek Respiration rates in treated fruit have been less com-
(2003), Watkins (2002), Watkins and Miller (2003), monly reported but are also inhibited by 1-MCP (Fan et
Watkins and Ekman (2005) and Watkins and Miller al., 1999; Fan and Mattheis, 1999a, 2001; Jiang and
(2005b). A website bhttp://www.hort. cornell.edu/mcp/ Joyce, 2002; Pre-Aymard et al., 2003; Saftner et al.,
N that categorizes the physiological and biochemical 2003; Defilippi et al., 2004; Mattheis et al., 2005;
responses for each product as decreased or delayed, Toivonen and Lu, 2005). However, respiration rates are
increased, or unaffected, was initiated in 2001 and is not reduced by 1-MCP below pre-climacteric levels (Mir
regularly updated (Watkins and Miller, 2005a); re- and Beaudry, 2002).
search results are available for 35 fruit and 14 vege- Softening is prevented or delayed by 1-MCP, the
tables (Table 1). effects of treatment often closely associated with eth-
In this review, the effect of 1-MCP on fruit and ylene production (Fan et al., 1999; Rupasinghe et al.,
vegetables is considered from two aspects. The first 2000; Watkins et al., 2000; Mir et al., 2001; Dauny and
considers the effects on factors that influence product Joyce, 2002; Pre-Aymard et al., 2003; Saftner et al.,
quality using several fruit that have received the most 2003; Zanella, 2003; Defilippi et al., 2004; DeLong et
attention in the literature, and that highlight some of al., 2004; Jayanty et al., 2004; Bai et al., 2005;
the challenges that exist in commercialization of 1- Mattheis et al., 2005; Moran and McManus, 2005;
MCP-based technology. While the responses of a wide Toivonen and Lu, 2005). The components of texture
range of vegetables have been examined (Table 1); that are affected by 1-MCP have not been adequately
most of these investigations have required supplemen- investigated but tissue toughness is greater in 1-MCP-
tary treatment with exogenous ethylene to show effects treated fruit than untreated fruit (Baritelle et al., 2001).
of 1-MCP and future commercial utilization is Firmness retention can also be excellent in fruit kept at
uncertain. Therefore, individual vegetables are not high temperatures (20–24°C) after treatment (Fan et
considered in detail here. Commercial development of al., 1999; Mir et al., 2001), and interestingly, Toivonen
1-MCP has largely centered on apple fruit. While semi- and Lu (2005) found that effects of 1-MCP on an early
commercial trials have been carried out with a wide ripening summer apple cultivar were lost at storage
variety of other registered crops, much of this work is temperatures below 15 °C.
proprietary, and access to commercial information is Loss of greenness of the background or ground color
less easily obtained. The second aspect takes informa- of the fruit skin, usually considered as a negative attri-
tion from the literature to provide an overview about bute in commercial conditions, is inhibited (Fan and
the physiological and biochemical responses of fruit Mattheis, 1999a, 2001; Dauny and Joyce, 2002; Jiang
and vegetables to 1-MCP to illustrate its potential to and Joyce, 2002; Pre-Aymard et al., 2003; Saftner et al.,
better understand the role of ethylene in ripening and 2003; Zanella, 2003), although no effect of 1-MCP on
senescence processes. color was detected by Dauny and Joyce (2002).
C.B. Watkins / Biotechnology Advances 24 (2006) 389–409 393
In general, 1-MCP delays loss of titratable acidity “apple” is a fruit with a wide variety of cultivars
(TA) concentrations (Fan et al., 1999; Fan and Mattheis, with different ripening rates and susceptibility to
1999a, 2001; Watkins et al., 2000; Pre-Aymard et al., various physiological and pathological disorders, and
2003, 2005; Saftner et al., 2003; Zanella, 2003; depending on cultivar, may be stored for up to a
Defilippi et al., 2004; Bai et al., 2005; Toivonen and year from harvest (Watkins, 2003). Also, preferences
Lu, 2005). However, absence of treatment effects (Mir for fruit attributes in the marketplace vary greatly
et al., 2001) or mixed responses depending on storage with cultivar, and include those with firm and acid
type (air or CA), on TA have been reported (Watkins et characteristics as well as those that are soft and
al., 2000). aromatic. 1-MCP-treated fruit have to meet varietal
Soluble solid concentrations (SSC) in 1-MCP treated requirements.
fruit can be higher, lower or the same as those in un- The research described in Section 2.1.1 has cat-
treated fruit (Fan et al., 1999; Watkins et al., 2000; Dauny alogued a wide variety of apple fruit responses to 1-
and Joyce, 2002; DeEll et al., 2002; Saftner et al., 2003; MCP, but many of these results have been obtained
Zanella, 2003; Bai et al., 2005; Moran and McManus, under ideal laboratory conditions, often when fruit are
2005; Pre-Aymard et al., 2005). treated on the day of harvest. Under commercial
Total volatile contents are reduced by 1-MCP treat- conditions fruit are usually placed into cold storage
ment, although individual volatiles are affected differ- and may be accumulated over several days before
entially (Rupasinghe et al., 2000; Lurie et al., 2002; treatment with 1-MCP. Research that examines the
Saftner et al., 2003; Defilippi et al., 2004; Bai et al., effects of various harvest and handling procedures is
2005; Kondo et al., 2005). These effects are discussed in emerging in the literature, and is resulting in
detail in Section 3.5. development of protocols to ensure that maximum
Apples are susceptible to a wide variety of physio- benefits of the technology are realized by the industry.
logical and pathological disorders, and the impact of The relationship between 1-MCP concentration and
1-MCP on susceptibility of fruit to these disorders is storage period is affected by cultivar and storage type
discussed in Sections 3.7 and 3.8. (Rupasinghe et al., 2000; Watkins et al., 2000; Pre-
Little formal sensory research is available on the Aymard et al., 2003). Longer exposure periods are
effects of 1-MCP on consumer acceptance of apples, required as the treatment temperature decreases (DeEll
partly because recent registration has precluded con- et al., 2002), although little difference in responses of
sumer testing. The rapidly ripening summer apple fruit treated with 1-MCP for 24h on the day of harvest
‘Anna’ treated with 1-MCP that had less fruity, ripe at 20 °C or after cooling overnight were detected
and overall aromas, and were firmer, crisper, juicier and (Dauny and Joyce, 2002; Watkins and Nock, 2005). In
less mealy, were more preferred in sensory analyses than addition, 1-MCP may lead to modification of CA
untreated fruit (Lurie et al., 2002; Pre-Aymard et al., recommendations, for example, it may reduce the
2005). requirement for CO2 during storage to maintain
firmness (DeEll et al., 2005).
2.1.2. Commercial application of 1-MCP Ethylene production by apple fruit occurs both on
The apple fruit provides an important example of a and off the tree when the climacteric is initiated, and
horticultural product to illustrate the opportunities and therefore the effectiveness of 1-MCP is affected both
limitations of 1-MCP-based technologies. The apple by the maturity/ripening stage at harvest and by the
was the first crop that received registration for 1-MCP period of time that the fruit are kept in cold storage
use, and 1-MCP use has been incorporated rapidly by before treatment. These two factors are inter-related as
industries around the world. The feature of 1-MCP more mature fruit at harvest produce autocatalytic
technology that has made it widely accepted by apple ethylene sooner than earlier harvested fruit. Cultivar
industries is that treated apple fruit maintain texture effects are also important; Mir et al. (2001) found
after removal from storage; in contrast, while CA relatively small effects of harvest maturity for ‘Deli-
storage can also maintain quality of apples during cious’, but large effects were shown for ‘Empire’
storage, these benefits are often lost during the (Watkins and Nock, 2005). Higher 1-MCP concentra-
subsequent shipping and marketing periods resulting tions result in better responses in fruit of advanced
in overly ripe and soft apple at the retail level. maturity (Watkins, unpublished data). However, these
Nevertheless, the rapid commercial adoption of 1- may exceed the maximum treatment concentrations of
MCP has not surprisingly resulted in a number of 1-MCP, which are established by regulation for each
challenges to growers and storage operators. The country, for example, 1μl l− 1 for apples in the US.
Watkins and Nock (2005) found that the effect of and with skin color changes appropriate to the cultivar.
delays between harvest and application of 1-MCP is Fruit responses to 1-MCP are ‘concentration × exposure
affected by cultivar, storage type and storage length. The time’-dependent in most (Feng et al., 2000; Jeong et al.,
suppliers of the commercial product, SmartFresh™, 2002), but not all (Woolf et al., 2005), studies. Both the
provide recommendations for maximum delays between timing of the peak of ethylene production and the
harvest and treatment, but these times are variable de- respiratory climacteric are delayed (Feng et al., 2000,
pending on fruit physiological condition, and a rule of 2004; Jeong et al., 2002, 2003; Hershkovitz et al., 2005).
thumb is that the delay times must be as short as possible Maximum rates of ethylene and carbon dioxide were
if longer storage periods are desired by the storage usually lower, but the magnitude can be higher in some
operator. It may be possible to store treated fruit at cultivars (Jeong et al., 2003; Hershkovitz et al., 2005).
warmer than normal cold storage temperatures if decay Treated fruit are firmer, slower to soften, and slower
and loss of other quality attributes such as TA is not to change skin color (Feng et al., 2000, 2004; Hofman et
unacceptably high (Mir et al., 2001; Jayanty et al., al., 2001; Jeong et al., 2002; Adkins et al., 2005;
2004). Hershkovitz et al., 2005; Woolf et al., 2005). 1-MCP
CA can prolong the impact of 1-MCP on both treated fruit have lower weight loss than untreated fruit
physical and sensory responses of apple fruit (Rupa- (Jeong et al., 2002). 1-MCP treatment combined with
singhe et al., 2000; Watkins et al., 2000; Mattheis et al., waxing does not extend the shelf life more than 1-MCP
2005). It is possible that 1-MCP can be an alternative to alone, but results in reduced weight loss and better
CA storage, but the two technologies generally are maintenance of green color (Jeong et al., 2003).
more effective when used in combination (Rupasinghe
et al., 2000; Watkins et al., 2000). Importantly, 2.2.2. Factors affecting commercial application of 1-MCP
variation in response of different fruit lots to 1-MCP Successful commercial utilization of 1-MCP for
can occur under commercial conditions, and CA use as avocado is dependent on an appropriate balance
a supplement can become more critical. Fruit from between excessively delayed ripening that can increase
orchard lots within a room that do not respond to 1- decay development, especially after removal from
MCP continue to deteriorate as storage times increase storage, and desirable increases in storage potential
resulting in unacceptable commercial losses (Watkins, with reduced internal disorders (Woolf et al., 2005).
unpublished data). Nevertheless, 1-MCP may be an Adkins et al. (2005) concluded that 1-MCP could extend
excellent replacement for CA storage for short term air marketing periods, but that treated fruit must have low
storage, especially for maintaining quality of summer disease inoculum load or be more resistant to infection
apples (Toivonen and Lu, 2005) and of other cultivars to avoid increased decay potential.
that deteriorate after only 2–3 months of storage Diffuse flesh discoloration, a symptom of internal
(Watkins et al., 2000; Bai et al., 2005), especially for chilling injury, was reduced in 1-MCP-treated fruit,
smaller local market operations that do not have CA although external chilling injury (skin blackening) in
facilities. ‘Hass’ was not affected by treatment (Pesis et al., 2002;
The availability of 1-MCP for the apple is allowing Woolf et al., 2005). Therefore, 1-MCP could allow
the rapid identification of commercial opportunities and storage of fruit at lower temperatures.
limitations of the technology. The success of 1-MCP for 1-MCP delayed ripening after ethylene treatment if
apple fruit appears to be based greatly on the main- fruit softening was not initiated (Jeong and Huber,
tenance of texture, especially because apple, unlike 2004). Ethylene treatment could overcome inhibition of
many other fruit types, do not soften markedly after ripening by 1-MCP at lower treatment concentrations,
harvest, and to date, at least the minimum demands for but ethylene recovery of 1-MCP ripening inhibition was
flavor have been met for most treated cultivars. An only partial and differed by ripening factor (Jeong and
extensive literature describing many aspects of the res- Huber, 2004).
ponses of apple fruit can be expected in the next several
years. 2.3. Banana (Musa sp., AAA group, Cavendish subgroup)
2.2. Avocado 2.3.1. Ripening physiology and quality

The banana fruit are typically harvested at the mature
2.2.1. Ripening physiology and quality green stage of maturity, transported, and then ripened
Ripening of avocado fruit is not initiated until they are artificially with ethylene before being sent to market.
harvested, and the fruit then softens to an edible texture Banana had increased ‘green life’ when treated with
1-MCP, responses being ‘concentration × exposure time’ 2.4. Pear (Pyrus communis L.)
dependent (Jiang et al., 1999b; Harris et al., 2000;
Bagnato et al., 2003). Ethylene production and respira- 2.4.1. Ripening physiology and quality
tion rates were lower in 1-MCP-treated fruit than in The effects of 1-MCP have been investigated using
untreated fruit (Golding et al., 1998, 1999; Pathak et al., summer, autumn and winter pears. High quality pear
2003; Pelayo et al., 2003; Lohani et al., 2004). Softening fruit have a buttery texture, with color change appropri-
of fruit was inhibited by 1-MCP treatment (Jiang et al., ate to the cultivar, and development of characteristic taste
1999a,b; Macnish et al., 2000; Botrel et al., 2002; and aroma associated with sugar and acid contents and
Pelayo et al., 2003; Lohani et al., 2004). Color changes volatile production (Kappel et al., 1995; Ma et al., 2000).
were also delayed in 1-MCP treated fruit (Botrel et al., Pears require exposure to chilling temperatures to ripen
2002), and may have been associated with disrupted or properly, with winter pears requiring as long as 8 weeks.
incomplete and uneven yellowing, even in the presence Ethylene production was inhibited by 1-MCP treatment
of propylene (Golding et al., 1998; Harris et al., 2000; (Argenta et al., 2003; Hiwasa et al., 2003; Kubo et al.,
Macnish et al., 2000). Total volatile production of fruit 2003; Ekman et al., 2004; Larrigaudiere et al., 2004;
was inhibited by 1-MCP treatment, and quantitatively, Trinchero et al., 2004; Mwaniki et al., 2005). Respiration
ester concentrations were lower, while those of alcohols rates were also lower in 1-MCP-treated fruit (Argenta et
were higher, in treated fruit (Golding et al., 1998). Little al., 2003; Kubo et al., 2003; Ekman et al., 2004).
is known about the effect of 1-MCP on other 1-MCP delayed or prevented fruit softening, the
compositional changes of banana fruit, although sugar degree of response depending on the cultivar and
content was not affected by treatment (Golding et al., 1-MCP concentration applied (Baritelle et al., 2001;
1998). Argenta et al., 2003; Hiwasa et al., 2003; Kubo et al.,
2003; Calvo and Sozzi, 2004; Ekman et al., 2004;
2.3.2. Factors affecting commercial application of 1-MCP Trinchero et al., 2004). Loss of greenness or yellowing
The limitation for commercial application of 1-MCP but not SSC, was inhibited by 1-MCP (Calvo and Sozzi,
for bananas may be the partial disruption of ripening 2004; Larrigaudiere et al., 2004; Trinchero et al., 2004).
events in treated fruit (Golding et al., 1998) that Treated fruit had higher TA in one study (Argenta et al.,
highlight the integrative role of ethylene in ripening 2003) but no effect of 1-MCP was found by others
(Golding et al., 1999). Harris et al. (2000) concluded (Calvo and Sozzi, 2004; Larrigaudiere et al., 2004;
that 1-MCP had limited commercial potential because Trinchero et al., 2004).
of the uneven color development, and that this problem
was exacerbated because of the range of maturities 2.4.2. Factors affecting commercial application of 1-MCP
present in a commercial consignment. In contrast, ‘1-MCP concentration × time’ relationships for delays
however, Bagnato et al. (2003) found that while of ripening have been shown (Argenta et al., 2003), and
ripening was inhibited and decay increased at high 1- the 1-MCP concentrations that delay, but do not
MCP concentrations, a lower concentration of 300nl ultimately prevent, normal ripening are variable. Appli-
l− 1 resulted in delayed ripening but firmness, color, cation of 0.2 μl l− 1 1-MCP resulted in normal ripening
SSC and aroma profiles were similar with those of with reduction in over-ripening problems (Calvo and
untreated fruit when compared at the same ripening Sozzi, 2004), while concentrations as high as 10 and
stages. 100 μl l− 1 to fruit in which ripening was initiated by
One strategy for use of 1-MCP on fruit such as chilling temperatures resulted in maintenance of optimal
banana for which ripening is artificially initiated with eating firmness for extended periods and prevention of
ethylene treatment could be to apply 1-MCP after this senescent breakdown development after storage (Kubo
treatment. Pelayo et al. (2003) concluded, however, et al., 2003).
that variability in responses of partially ripe bananas Repeated 1-MCP applications can further slow down
was too variable for commercial application. Also, if ripening (Ekman et al., 2004), but sensitivity to second
1-MCP was applied to fruit 24 h after propylene applications are affected if ethylene production by fruit
treatment to induce ripening, then the onset of has been initiated during storage (Trinchero et al., 2004).
ethylene and respiration rates was not affected but The efficacy of post-storage ethylene treatments to ini-
color and volatile production was inhibited (Golding tiate ripening of 1-MCP-treated fruit after storage
et al., 1998). Treatment of pre-climacteric fruit, or depends on 1-MCP concentration and storage duration
earlier 1-MCP treatment, resulted in inhibition of all (Argenta et al., 2003; Calvo and Sozzi, 2004; Ekman et
processes. al., 2004).
Both positive and negative aspects of 1-MCP that 2.5.2. Factors affecting commercial application of 1-MCP
may be commercially important for pear fruit have been Responses of fruit to 1-MCP are affected by
identified. 1-MCP reduces susceptibility of fruit to skin concentration and exposure period, but not treatment
browning, and vibration and impact bruising, and there- temperature (Liguori et al., 2004). Optimal 1-MCP
fore its use may permit greater flexibility during grading, concentrations vary greatly from as low as 0.4μl l− 1
packaging and transport operations (Calvo and Sozzi, (Liu et al., 2005) to 5 μl l− 1 (Liguori et al., 2004), the
2004; Ekman et al., 2004). However, increased storage latter concentration being higher than that registered for
periods of 1-MCP-treated fruit may result in greater use. Inhibition of fruit ripening is transitory in all
weight loss (Calvo and Sozzi, 2004). Ethylene appears to published studies, but repeated 1-MCP applications
be required for both initiation and progression of rip- helps maintain suppression of ripening (Liu et al., 2005).
ening of pears (Hiwasa et al., 2003). Ekman et al. (2004) The transitory effect of 1-MCP is not related to diffusion
found that skin color and firmness was dissociated and limitations within the flesh (Hayama et al., 2005). 1-
suggested that this could problematic at the retail level. MCP treatment at ambient temperatures could allow
Variation of individual fruit treated with 1-MCP could commercially significant extension of fruit shelf life.
result in the need for post-treatment sorting of fruit However, decreased 1-MCP effects in fruit stored at 4
o
(Trinchero et al., 2004). C (Bregoli et al., 2005), and a greater incidence of
chilling-related disorders in treated fruit (Dong et al.,
2.5. Peach and nectarine (Prunus persica L.) 2001b; Fan et al., 2002; Girardi et al., 2005) suggests
that this technology is limited for extending the storage
2.5.1. Ripening physiology and quality life of peaches and nectarines in cold storage.
Ethylene production is inhibited by 1-MCP treat-
ment of peaches and nectarines (Mathooko et al., 2.6. Plum (Prunus domestica L. and Prunus salicina L.)
2001; Fan et al., 2002; Rasori et al., 2002; Liguori et
al., 2004; Bregoli et al., 2005; Girardi et al., 2005), 2.6.1. Ripening physiology and quality
although transient increases in production were 1-MCP prevented or delayed the climacteric increase
detected immediately after treatment in one study in ethylene production of plums (Abdi et al., 1998;
(Fan et al., 2002). Ethylene production of 1-MCP Dong et al., 2001a, 2002; Martinez-Romero et al., 2003;
treated fruit was unaffected at the time of treatment, Salvador et al., 2003; Valero et al., 2003, 2004), even
but was reduced after storage (Dong et al., 2001b). when fruit were harvested close to the climacteric peak
Recovery from 1-MCP-induced ripening inhibition (Salvador et al., 2003). Respiration rates were also de-
resulted in greater production of ethylene than creased or the climacteric increase delayed (Dong et al.,
observed in untreated fruit (Rasori et al., 2002). 2002; Martinez-Romero et al., 2003; Salvador et al.,
Respiration rates of treated fruit were lower (Fan et 2003; Valero et al., 2003), but no effect of 1-MCP
al., 2002), or not affected (Dong et al., 2001b; Liguori treatment was detected by Dong et al. (2001a).
et al., 2004) by 1-MCP treatment. Softening of fruit Softening of the fruit was delayed by 1-MCP treatment
was delayed when fruit were kept at about 20 °C (Dong et al., 2001a, 2002; Skog et al., 2001; Martinez-
(Dong et al., 2001b; Mathooko et al., 2001; Fan et al., Romero et al., 2003; Salvador et al., 2003; Valero et al.,
2002; Rasori et al., 2002; Bregoli et al., 2005; Liu et 2003, 2004; Menniti et al., 2004). Skin color changes
al., 2005), and in fruit from early and late harvest were delayed by 1-MCP (Dong et al., 2002; Martinez-
(Liguori et al., 2004). However, Fan et al. (2002) Romero et al., 2003; Salvador et al., 2003; Valero et al.,
found that the effects of fruit maturity were greater 2003, 2004; Menniti et al., 2004), and weight loss
than those of 1-MCP treatment, with early harvested during and after storage decreased (Martinez-Romero et
fruit showing little response to treatment. Also, al., 2003; Valero et al., 2003).
beneficial effects of 1-MCP were lost at 4°C (Bregoli Responses of SSC and TA to 1-MCP treatment are
et al., 2005). variable. The SSC was not affected by 1-MCP treatment
1-MCP treatment either did not affect SSC of fruit (Dong et al., 2002; Salvador et al., 2003; Menniti et al.,
(Liguori et al., 2004), resulted in lower SSC (Fan et al., 2004), but its ripening-associated increase was (Valero et
2002; Bregoli et al., 2005), or the increase of SSC al., 2004). While TAwas not affected by 1-MCP treatment
during ripening was delayed (Liu et al., 2005). Loss of (Menniti et al., 2004), loss of acidity was reduced in other
TA was reduced in high acid (Fan et al., 2002; Liguori et studies (Dong et al., 2002; Salvador et al., 2003). The SSC
al., 2004; Bregoli et al., 2005; Liu et al., 2005), but not to TA ratio was lower in 1-MCP-treated than untreated
in low acid cultivars (Liguori et al., 2004). fruit (Martinez-Romero et al., 2003; Valero et al., 2003).
Although plums are categorized as climacteric fruit, include respiration rates (Wills and Ku, 2002; Colelli et
suppressed-climacteric cultivars are available (Abdi et al., 2003; Krammes et al., 2003), color change and
al., 1998; Martinez-Romero et al., 2003). The ‘sup- softening (Sisler et al., 1996; Hoeberichts et al., 2002;
pressed’ cultivars did not ripen when treated with 1- Colelli et al., 2003; Mostofi et al., 2003; Mir et al.,
MCP unless they were subsequently treated with 2004; Opiyo and Ying, 2005; Tassoni et al., in press).
propylene (Abdi et al., 1998), and the effects of 1- TA was higher in treated fruit (Moretti et al., 2002;
MCP were not dose dependent, maximum responses Wills and Ku, 2002; Krammes et al., 2003; Opiyo and
occurring at the lowest 1-MCP concentration applied Ying, 2005), but SSC were not affected by 1-MCP
(Martinez-Romero et al., 2003). In contrast, dose treatment (Moretti et al., 2002; Wills and Ku, 2002;
dependent responses were shown for a normal ethyl- Colelli et al., 2003; Krammes et al., 2003; Mir et al.,
ene-producing cultivar. 2004; Opiyo and Ying, 2005). Weight loss from fruit
was not affected by 1-MCP treatment (Wills and Ku,
2.6.2. Factors affecting commercial application of 1-MCP 2002; Colelli et al., 2003). Only small effects of
Postharvest softening and susceptibility to mechan- treatment on aroma volatiles were detected by Mir et al.
ical injury and pathogens are major factors limiting the (2004), although changes tended towards those vola-
shipping, storage and shelf life of plums. The degree tiles that were associated with harvest of fruit at earlier
of response to 1-MCP varies greatly by cultivar and ripening stages.
harvest maturity (Abdi et al., 1998; Skog et al., 2001;
Martinez-Romero et al., 2003) but reports of extension 2.7.2. Factors affecting commercial application of 1-MCP
of storage periods from 1 week for untreated fruit to 4 The desirable response of fresh market tomato fruit
weeks for 1-MCP-treated fruit plus 7 days at 20°C to 1-MCP is a delay of ripening, but then ripening to
without negative effects indicate that 1-MCP may be a redness, desired softness, and flavor development.
very useful technology for this fruit. Moreover, 1- Another possible commercial benefit of 1-MCP is
MCP is effective at later stages of maturity when inhibition of abscission of cherry tomatoes from vines
better quality characteristics have developed (Salvador (Beno-Moualem et al., 2004). The extent of ripening
et al., 2003; Valero et al., 2003). Studies on inhibition of tomato fruit is affected by 1-MCP con-
climacteric and suppressed-climacteric cultivars indi- centration, exposure time and ripening stage (Sisler et
cate however, that 1-MCP concentrations will need to al., 1996; Hoeberichts et al., 2002; Moretti et al., 2002;
be calibrated for cultivars with different ethylene Wills and Ku, 2002; Mir et al., 2004; Opiyo and Ying,
production (Abdi et al., 1998; Martinez-Romero et al., 2005), and optimal treatment concentrations are also
2003). affected by cultivar (Krammes et al., 2003). Fruit
No differences were noted between treatment at 0 and recover capacity to ripen after treatment, but second
20 °C (Menniti et al., 2004), although Valero et al. (2003) applications further delay ripening (Hoeberichts et al.,
suggested that treatment at cold storage temperatures 2002; Mir et al., 2004). However, the maturity stage of
resulted in better control of ethylene. 1-MCP treatment the fruit at the time of 1-MCP treatment affects the
prior to simulated mechanical harvest decreased fruit ability of the fruit to recover; mature green and breaker
losses due to bruising (Lippert and Blanke, 2004). In one stage fruit shriveled and developed decay before
of the few studies that have investigated possible in- ripening, although fruit treated at pink and light red
teractions between 1-MCP treatment and postharvest stages ripened properly after a delay (Hurr et al.,
handling operations, Valero et al. (2004) found that 2005). The presence of ethylene was required
1-MCP treatment of fruit packed in small ventilated throughout for ripening even when 1-MCP was
cardboard boxes resulted in better ripening control than applied at advanced ripening stages (Hoeberichts et
for bulked fruit. al., 2002). Mostofi et al. (2003) also found that
treatment of mature green fruit resulted in impaired
2.7. Tomato (Solanum esculentum Mill) color development, and that the ripening temperature
was important; whereas ripening of fruit treated at the
2.7.1. Ripening physiology and quality breaker stage was coordinated at 15, 20 and 25 °C,
1-MCP treatments markedly affected ripening of development of color was out of phase with softening
tomato fruit by inhibiting ethylene production (Hoe- at 25°C. The reddening of locular tissues was later
berichts et al., 2002; Wills and Ku, 2002; Krammes et than that of pericarp tissues after treatment of mature
al., 2003; Opiyo and Ying, 2005; Tassoni et al., in green fruit, which could affect appearance of fresh cut
press). Other ripening processes that are inhibited slices (Mir et al., 2004).
Formal sensory studies on the effects of 1-MCP on Research with yeast has shown that ETR1 and ERS1,
consumer acceptability are not available. However, Wills genes encoding the ethylene binding proteins, show equal
and Ku (2002) concluded that MCP-treated fruit should sensitivity to 1-MCP (Hall et al., 2000), but little research
be of superior quality because of improved SSC / TA on expression of these genes in fruit and vegetables is yet
ratios. 1-MCP had little effect on aroma volatiles in the available. Accumulation of transcripts for the genes
study of Mir et al., (2004), but negatively and encoding ERS decreased in treated apple (Defilippi et al.,
irreversibly affected them in informal sensory analyses 2005). Rasori et al. (2002) found that 1-MCP did not
reported by Hurr et al. (2005). affect transcription of the gene PP-ETR1, but down-
regulated that of PP-ERS1. Recovery from 1-MCP
3. Physiological and biochemical responses of fruits inhibition was associated with increased accumulation
and vegetables to 1-MCP of PP-ERS1 transcripts. In tomato, recovery from
ripening inhibition was associated with increased gene
Generalizations regarding the effects of 1-MCP on expression for both ETR1 and ERS1 (Tassoni et al., in
physiological and biochemical responses of fruits and press).
vegetables are shown in Table 2. Specific examples are Ethylene production of fruit is usually inhibited by
provided here, but fuller documentation of these effects 1-MCP treatment, but the persistence of the inhibition
is described on a website bhttp://www.hort. cornell.edu/ can be variable (Fan et al., 1999; Fan and Mattheis,
mcp/N (Watkins and Miller, 2005a). 1999a; Dong et al., 2002; Jeong et al., 2002; Ergun et al.,
2005). Ethylene production is not always inhibited. The
3.1. Ethylene metabolism decline in ethylene production was slower over time in
1-MCP treated pineapples than untreated fruit (Selvar-
The action of 1-MCP is mediated through the ajah et al., 2001), and greater ethylene production has
inhibition of ethylene perception of plant tissues by been observed in fruit and vegetables depending on
interacting with the receptor and competing with cultivar, maturity, stage or ripening or 1-MCP concen-
ethylene for binding sites (Sisler et al., 1996; Sisler and tration, including treated avocado (Jeong et al., 2003;
Serek, 1997, 2003). Therefore, the effectiveness of Hershkovitz et al., 2005), banana (Golding et al., 1998),
inhibition of ripening and/or senescence of fruit and grapefruit (Mullins et al., 2000), strawberry (Tian et al.,
vegetables is a function of the 1-MCP concentration 2000), Chinese cabbage (Porter et al., 2005), coriander
applied, up to saturation of the binding sites. Depending (Jiang et al., 2002a), and parsley (Ella et al., 2003). In
on the product it can be desirable for the inhibition of some cases the increased ethylene does not appear to
ethylene-mediated responses to persist indefinitely, es- affect other senescence processes, e.g., 1-MCP citrus
pecially in the case of leafy vegetables, but for fruit, fruit remained green despite higher ethylene production
recovery from 1-MCP-induced inhibition of ripening is (Mullins et al., 2000), but increased ethylene production
often essential in order to provide a ripened product that can accelerate senescence in other tissues, e.g., parsley
is acceptable to the consumer. (Ella et al., 2003). It is likely that stimulated ethylene
The extent and longevity of 1-MCP action is affected production is due to loss of negative feedback regulation
by species, cultivar, tissue and mode of ethylene bio- of ethylene biosynthesis.
synthesis induction. A ‘concentration × time’ effect is The genes encoding two key enzymes of the ethylene
apparent with longer exposure periods required for lower biosynthetic pathway, 1-aminocyclopropane carboxylic
1-MCP concentrations to obtain the same physiological acid oxidase (ACO) and 1-aminocyclopropane carboxylic
effects (Sisler and Serek, 1997). Some products such as acid synthase (ACS), and their respective enzyme acti-
pea require higher concentrations (40 nl l − 1 ) than vities have been studied in several crops. In apple, banana,
carnations (0.5 nl l− 1) and banana (0.7 nl l− 1), suggesting melon and pear fruit, inhibition of ethylene production by
that new receptors are produced in growing tissues or 1-MCP was accompanied by lower expression of these
that a low affinity form of the receptors is present (Sisler genes (Lelievre et al., 1997; de Wild et al., 1999; Defilippi
and Serek, 2003). Synthesis of new binding sites may be et al., 2005), and lower activities of ACS and ACO (Dong
affected by temperature; in banana, temperatures bet- et al., 2001b; Pathak et al., 2003; Defilippi et al., 2005). In
ween 30 and 40°C results in faster recovery of ripening, peach, inhibited ethylene production was associated with
while application of 1-MCP at 2.5 °C is less effective reduced activity of ACO and a reduction in PP-ACO1 and
than at 15 and 20 °C suggesting that binding of 1-MCP at PP-ACO2 transcript accumulation, but 1-aminocyclopro-
low temperatures was incomplete (Jiang et al., 2002b, pane carboxylic acid (ACC) accumulated in treated fruit,
2004c). and PP-ACS1 expression and ACS activity was not
Table 2
Generalizations regarding the effects of 1-MCP on metabolism of fruit and vegetables
Attribute or process affected Enzyme activity or associated gene Increased (↑), decreased (↓), or
expression unchanged (↔)
Ethylene metabolism
Ethylene perception ETR1, ERS1 ↓↔
Ethylene production ↓↑
ACC synthase (ACS) expression and ↓
activity
ACC oxidase (ACO) expression and ↓
activity
Respiratory metabolism
Respiration rate ↓ ↑↔
SSC ↓ ↑↔
TA ↓ ↑↔
Pigments Chlorophyll degradation ↓
Lycopene accumulation ↓
Anthocyanin accumulation ↓
Chlorophyllase activity ↓
Phenolic metabolism
Total phenolic content ↓
Phenylalanine ammonia lyase (PAL) ↓
activity
Polyphenol oxidase (PPO) activity ↓
Cell wall metabolism
Soluble polyuronide content ↓
Polygalacturonase (PG) activity ↓
Pectin methylesterase (PME) ↓
Endo-β-1,4-glucanase (EGase) ↓
Glycosidases ↓↔
Volatile compound metabolism
Esters ↓
Aldehydes ↔
Terpenoid biosynthesis ↔
Acetaldehyde and ethanol ↓↑
accumulation
Alcohol acyl transferase activity ↓
Alcohol dehydrogenase activity ↔
Nutritional Vitamin C loss ↓
Anthocyanin contents ↓
Phenolic contents ↓↔
Antioxidant activity ↓
loss
Physiological disorders
Senescent disorders ↓
Chilling injury ↓↑
Superficial scald (apples and ↓
pears)
Ethylene-induced disorders ↓
Controlled atmosphere- ↑
induced
Abscission ↓
Pathological disorders
Susceptibility to pathogens ↑↓↔
Fungal growth ↓↔
affected by treatment (Mathooko et al., 2001). 1-MCP Application of 1-MCP to avocados at the pre-climacteric
treated nectarines had lower ACS, ACO1 and ACO2 stage and at the onset of the climacteric inhibited ACS and
transcript accumulations than untreated fruit at ambient ACO activities, and the transcription of PA-ACS1, and
but not cold storage temperatures (Bregoli et al., 2005). suppressed PA-ACO and PA-ERS1 mRNAs to trace
levels (Owino et al., 2002). Discontinuation of 1-MCP 1999; Watkins et al., 2000; Benassi et al., 2003). The
action resulted in super-induction of these genes in the sugar content of banana was not affected by 1-MCP
fruit. In citrus, 1-MCP-induced increases of ethylene treatment (Golding et al., 1998), but lower SSCs in 1-
production were associated with greater ACC accumula- MCP treated fruit were associated with lower sucrose
tion and higher ACS transcript accumulation and enzyme concentrations (Defilippi et al., 2004).
activity (Mullins et al., 2000).
In tomato, transcript accumulations of ACO1, phy- 3.3. Pigment metabolism
toene synthase 1 (PSY1) and expansin 1 (EXP1), used as
indicators of treatment effects on ethylene biosynthesis, Loss of greenness, or yellowing, in most products is
color and softening, respectively, were decreased by inhibited by 1-MCP. For many products, especially
1-MCP (Hoeberichts et al., 2002). Increases in transcript leafy vegetables and certain fruit such as apple, main-
abundance of LE-ACS2, LE-ACS4 and LE-ACO1 tenance of green color is desirable in the marketplace as
mRNAs in ripening fruit were inhibited by 1-MCP, but yellowness is regarded as a sign of senescence.
ethylene production, ACC content and ACS and ACO However, for many fruit loss of chlorophyll and
activities were not inhibited to the expected levels sug- development, or unmasking, of colored pigments is an
gesting involvement of negatively regulated genes in essential aspect of ripening (Kays, 1997). Therefore,
ethylene biosynthesis (Nakatsuka et al., 1997). Reduced successful 1-MCP use requires a delay, but not
ethylene production of fruit that were treated at the turning irreversible inhibition, of the processes involved in
and pink stages was accompanied by inhibited ACS2, pigment metabolism. 1-MCP inhibited anthocyanin
ACS4, ACO1 and ACO4 transcript accumulation, while increases in strawberry fruit (Jiang et al., 2001), but
several other ACS genes were not affected (Nakatsuka et loss of chlorophylls and development of colors
al., 1998). 1-MCP abolished expression of E4 mRNA in (anthocyanins, lycopene) eventually reached desirable
wounded tomato fruit, but not expression profiles of LE- levels of those of untreated fruit. However, yellowing of
ACS2, LE-ACS6 and LE-ACO1 (Yokotani et al., 2004). banana fruit could be disrupted or incomplete and
Itai et al. (2003) showed that α-L-arabinofuranosidase uneven, even in the presence of propylene (Golding et
(LeARF1) expression was negatively regulated by ethy- al., 1998; Harris et al., 2000; Macnish et al., 2000), and
lene, while two B-D-xylosidase genes were independent there was a separation of color changes from other
of ethylene action. NR transcripts were immediately ripening attributes in pears (Ekman et al., 2004).
suppressed by 1-MCP treatment but as fruit ripened, they Ethylene-induced degreening of 1-MCP-treated cucum-
recovered from 1-MCP in a similar pattern to that of the bers was totally inhibited or occurred with development
ACS and ACO genes (Tassoni et al., in press). of uneven yellowing (Nilsson, 2005).
Marty et al. (2005) used 1-MCP treatments to in-
3.2. Respiration rate vestigate ethylene regulation of carotenoid accumulation
and carotenogenic gene expression in orange and white
As described for several fruit in Section 2, the apricot cultivars. Both cultivars accumulated the color-
respiration rates of most treated products decreased or less phytoene and phytofluene pigments, but β-carotene
were delayed, especially in climacteric fruit where in- accumulated only in the orange fruit. While1-MCP in-
creases accompany increases of ethylene production. hibited ethylene production of both cultivars, effects on
The peak respiration rates at the climacteric were re- pigment accumulations were small.
duced by 1-MCP (Jeong et al., 2002, 2003). Enhanced Chlorophyll fluorescence changes were delayed in
respiration rates of ethylene-treated strawberry fruit 1-MCP-treated fruit kept at ambient temperatures (Mir et
were reduced by 1-MCP at earlier harvests, but not in al., 2001; Jayanty et al., 2004). Little is known about the
fruit from later harvests (Tian et al., 2000). Bower et al. effects of 1-MCP on pigment metabolism. However,
(2003) found higher respiration rates in 1-MCP-treated Gong and Mattheis (2003) and Hershkovitz et al. (2005)
strawberry fruit that may have been associated with have found that chlorophyllase activity was reduced in
earlier onset of decay. 1-MCP-treated broccoli florets and avocado fruit,
Starch degradation is sometimes delayed in 1-MCP- respectively.
treated fruit (Fan et al., 1999). SSC in treated products
might be expected to be higher than in untreated 3.4. Cell wall metabolism
products because of lower respiration rates, but can be
higher, lower or the same as in untreated fruit depending For most products, delayed rather than complete
on the product and the storage conditions (Fan et al., inhibition of softening, is desirable (Section 2). For non-
climacteric fruit such as strawberries and oranges, the 3.5. Volatile compound metabolism
effects of 1-MCP on softening were usually not de-
tectable (Porat et al., 1999; Tian et al., 2000), but soft- Flavor is a composite of taste and odor, and volatile
ening was enhanced under some treatment combinations production can be greatly affected by ethylene. There-
with ethylene (Tian et al., 2000). Studies of 1-MCP on fore, decreased and/or altered volatile production in
cell wall changes of treated fruit are limited, but a number 1-MCP compared with untreated fruits may impact
of investigations on cell wall enzymes are available. product acceptability by consumers. However, sensory
Decreased softening in 1-MCP-treated bananas is analyses are limited to few products because of the
associated with lower expression of an ethylene induced relatively recent registration of 1-MCP, and absence of
expansin (MaExp1) gene (Trivedi and Nath, 2004), and registration for many fruits and vegetables.
lower activities of pectin methylesterase (PME), poly- In apple fruit, ester production was inhibited by
galacturonase (PG), endo-β-1,4-glucanase (EGase) and 1-MCP, but results for other components were variable,
pectate lyase activities (Lohani et al., 2004). Effects of perhaps reflecting different cultivars and storage condi-
1-MCP on softening of pears were associated with de- tions. Mattheis et al. (2005) found that production of
creased β-galactosidase activity and differential effects esters, alcohols, aldehydes, acetic acid and 1-methoxy-
on expression of its genes (Mwaniki et al., 2005), lower 4-(2-propenyl)benzene was inhibited by 1-MCP, but
glycosidase activities (Trinchero et al., 2004), and their production increased when kept at room tem-
transcript accumulation of genes for PG1 and PG2, peratures after long term storage. Esters and alcohols
but not EGase (Hiwasa et al., 2003). Delayed softening were lower in 1-MCP-treated fruit than untreated fruit
of peaches was associated with delayed increases in (Defilippi et al., 2004; Kondo et al., 2005) but aldehydes
soluble pectin concentrations (Liu et al., 2005). were not affected (Defilippi et al., 2004). Esters were
Activities of exo-PG and EGase were lower in 1-MCP reduced but the proportions of alcohols and the aldehyde
treated plums than in untreated fruit, but treatment did 2-hexenal in treated fruit were higher (Lurie et al., 2002).
not affect activities of endo-PG and pectin esterase (PE). CA storage also inhibits volatile production of apple
Alterations of cell wall enzyme associated with fruit. This inhibition was enhanced in 1-MCP-treated
enhanced susceptibility of peach fruit to a chilling fruit, but the dynamics of volatile change over time were
injury expressed as woolliness is discussed in Section different in response to the two treatments (Mattheis et
3.7.2. al., 2005); the effects of CA increased, while those of
Delayed softening of 1-MCP-treated avocado fruit 1-MCP decreased, over time. The concentrations of
is reflected in similar patterns of delayed solubilization branched but not straight-chained esters recovered in
and degradation of polyuronides (Jeong et al., 2002; 1-MCP-treated fruit, though not to those of untreated
Jeong and Huber, 2004). Activities of PG and EGase fruit (Mattheis et al., 2005) confirming earlier indica-
were lower in 1-MCP-treated fruit (Feng et al., 2000). tions that sensitivity to ethylene action varies among the
Interestingly PG activity did not recover even though pathways of ester production (Fan and Mattheis, 1999b).
fruit softened to those comparable with control fruit Ethylene production clearly has some direct effects on
indicating that PG is not required for extensive volatile production of fruit, and indeed continuous ethy-
softening (Jeong et al., 2002). Changes of PME, α- lene action is required for maximum volatile production
and β-galactosidase and EGase activities were delayed (Fan et al., 1998; Fan and Mattheis, 1999b). While the
but essentially followed patterns of increase or decline mechanisms of 1-MCP action on volatiles are not
of the untreated fruit (Jeong and Huber, 2004). Lower known, Defilippi et al. (2005) found that alcohol dehy-
activities of β-galactosidase, α-arabinofuranosidase drogenase (ADH) activity was not affected by 1-MCP,
and β-xylosidase were associated with delayed but alcohol acyl transferase (AAT) transcripts and acti-
softening of 1-MCP treated kiwifruit (Boquete et al., vity were lower in treated apple fruit suggesting that
2004). modulation of this step in volatile synthesis by ethylene
1-MCP affected gene expression of cell wall- may be responsible for lower ester production. Expres-
related genes of the non-climacteric strawberry, but sion of four AAT genes were suppressed by 1-MCP
specific effects depended on fruit ripening stage treatment of melons (El-Sharkawy et al., 2005).
(Balogh et al., 2005). 1-MCP treatment resulted in In pear fruit, increases of ester and alcohol production
up-regulation of a putative EGase in green fruit, but were delayed by 1-MCP, with quantitative differences
its down-regulation in red fruit, up-regulation of a between long chain and branched esters, but volatiles were
ripening-repressed β-galactosidase, and down-regula- similar in treated and untreated pear fruit when equally
tion of pectic lyase gene. ripe (Argenta et al., 2003). Total volatile production of
bananas was inhibited by 1-MCP treatment, and quan- injury which is higher in 1-MCP-treated fruit than un-
titatively, ester concentrations were decreased, while those treated fruit (DeEll et al., 2003; Zanella, 2003; Watkins
of alcohols were increased in treated fruit (Golding et al., and Nock, 2004). The disorder is associated with early
1998). Abdi et al. (1998) found that aroma volatile harvested fruit and early exposure to carbon dioxide in
production of plums was ethylene-dependent or the storage atmosphere (Watkins et al., 1997; Fernandez-
-independent, depending on the cultivar. 1-MCP increased Trujillo et al., 2001) and the effect of 1-MCP is consistent
acetaldehyde and ethanol accumulation in citrus resulting with maintenance of the fruit in a less ripe state and thus
in off-flavor development (Porat et al., 1999), but delayed more susceptible to injury. Risk of injury is eliminated by
accumulation of these compounds in plums (Salvador et use of the antioxidant diphenylamine (DPA), registered
al., 2003). Where reduced aroma is associated with to control superficial scald, or reduced by maintaining
1-MCP treatment, the commercial implications are likely very low carbon dioxide concentrations in the storage
to vary by product type, being more critical for products atmosphere for the first few weeks of storage (Watkins
and cultivars where aroma is a quality characteristic and Nock, 2004).
expected by the consumer. For some products, certain Some interactions of 1-MCP with other technologies
aromas are associated with over-ripening and therefore have been investigated: Fan and Mattheis (2001) found
their inhibition is desirable, or aroma concentrations may that irradiation damage was aggravated in 1-MCP-trea-
be less important than texture and acid/sugar levels. ted fruit when they were kept at 20 °C. Lu and Toivonen
(2003) showed a synergistic benefit of 1-MCP and 35%
3.6. Nutritional quality CO2 on quality of ‘Gala’ apples, without injury.
Most research focus of 1-MCP and disorders of apple
The effect of 1-MCP on nutritional quality has not has been on superficial scald (syn. storage scald) because
been studied thoroughly. However, 1-MCP slows of the interaction between ethylene production and that
vitamin C loss in Chinese jujube (Jiang et al., 2004b), of α-farnesene, and early reports that 1-MCP inhibited
peaches (Liu et al., 2005), pineapples (Selvarajah et al., disorder development (Fan and Mattheis, 1999a,b;
2001), and minimally processed lettuce and pineapple Rupasinghe et al., 2000; Watkins et al., 2000). Super-
(Budu and Joyce, 2003; Tay and Perera, 2004). Although ficial scald is a physiological storage disorder of suscep-
phenolic contents of apples were not affected by 1-MCP tible apple and pear cultivars that is manifested as
treatment (Defilippi et al., 2004), 1-MCP treated ‘Deli- browning or blackening of the skin resulting from nec-
cious’ and ‘Empire’ apples maintained higher water- rosis of the hypodermal cells (Bain and Mercer, 1963).
soluble antioxidant activity than untreated fruit after cold Development of the disorder occurs during low
storage (MacLean et al., 2003). Lower phenolic and temperature storage and it is thought to be a chilling
anthocyanin contents in strawberry fruit treated with injury (Watkins et al., 1995). Injury to cells probably
high 1-MCP concentrations (Jiang et al., 2001) could results from free radical reactions associated with the
potentially reduce antioxidative activity. oxidation of α-farnesene to conjugated trienols (CTols),
predominantly 9E and 9Z isomers of 2,6,10-trimethyl-
3.7. Physiological storage disorders dodeca-2,7,9,11-tetraen-6-ol (Rowan et al., 1995; Whi-
taker et al., 1997). Accumulation of α-farnesene in the
3.7.1. Apple skin is associated with ethylene (Du and Bramlage,
The literature on physiological storage disorders is 1994; Watkins et al., 1995; Whitaker et al., 2000).
greatest for apple fruit. 1-MCP can reduce senescent Development of scald can be reduced or prevented by
breakdown (Watkins et al., 2000; DeLong et al., 2004; inhibiting α-farnesene production or its accumulation,
Moran and McManus, 2005), brown core (syn. core- e.g., by ventilation, use of oil wraps, or by inhibiting its
flush) (Fan and Mattheis, 1999a; Zanella, 2003), core oxidation to CTols by DPA (Huelin, 1968; Lurie et al.,
browning (DeLong et al., 2004), coreline browning 1989). It has been well demonstrated that inhibition of
(Moran and McManus, 2005) and soft scald (Fan and scald by 1-MCP is associated with inhibition of α-
Mattheis, 1999a), which are variously disorders associ- farnesene accumulation that restricts substrate available
ated with senescence and cold storage. Dissipation of for oxidation (Fan and Mattheis, 1999a; Rupasinghe et
watercore from fruit is reduced by 1-MCP treatment al., 2000; Watkins et al., 2000; Shaham et al., 2003;
(Watkins, unpublished data). Susceptibility of some Arquiza et al., 2005; Pechous et al., 2005).
disorders appears to be increased by 1-MCP treatment, Although it may not be considered a storage disorder
although little research on these responses has yet been sensu stricto, an undesirable feature of some apple cul-
published. An exception is external carbon dioxide tivars is development of greasiness, or slipperiness to
touch, which is disliked by consumers. Greasiness is associated with decreased responses of the fruit to
associated with changes in the wax and oil fractions in ethylene.
the skin (Morice and Shorland, 1973), and in susceptible 1-MCP treatment prevents ethylene induced devel-
cultivars, its development can occur in more mature fruit opment browning and chemical changes in lettuce and
and with longer storage periods (Leake et al., 1989a,b). carrots. 1-MCP delayed ethylene-induced russet spot-
1-MCP inhibited development of greasiness (Fan and ting of whole and minimally processed lettuce, and
Mattheis, 1999a; Watkins and Nock, 2005) suggesting browning of the cut surfaces of shredded lettuce (Wills
that it may be a useful tool to manage the problem. et al., 2002; Saltveit, 2004; Tay and Perera, 2004).
Interestingly, while these disorders are associated with
3.7.2. Other fruits and vegetables induction of phenypropanoid metabolism and greater
Development of superficial scald in pear fruit was accumulation of phenolic compounds, 1-MCP did not
also controlled by 1-MCP, but developed as fruit were interfere with wound-induced browning (Saltveit,
released from inhibition of ripening (Ekman et al., 2004). 2004). 1-MCP-treated carrots had inhibited accumula-
Thus, total control of the disorder only occurred in fruit tion of 8-hydroxy-3-methoxy-3,4-dihydro-isocoumarin,
that did not ripen. 1-MCP inhibited production of α- associated with bitter flavors and the phytoalexin, 6-
farnesene and its oxidation product, MHO, in pears methoymellin (Fan and Mattheis, 2000; Fan et al.,
(Argenta et al., 2003). Other disorders of pears that are 2000). 1-MCP prevented ethylene-induced water-soak-
inhibited by 1-MCP include senescent scald and core ing of watermelon and the associated increase in
browning (Argenta et al., 2003), watery and core phospholipid (phosphatidylcholine and phosphatidyli-
browning (Calvo and Sozzi, 2004), senescent break- nositol) degradation and activities of phospholipases C,
down (Kubo et al., 2003), internal breakdown (Ekman et phospholipases D and lipoxygenase (Mao et al., 2004).
al., 2004) and decay (Argenta et al., 2003). Postharvest pitting of citrus fruit was inhibited by 1-
Incidences of low temperature disorders of a number MCP (Dou et al., 2005).
of fruit were reduced by 1-MCP treatment, including
internal flesh browning in avocado (Pesis et al., 2002; 3.7.3. A physiological basis for disorder development in
Hershkovitz et al., 2005; Woolf et al., 2005), and pine- response to 1-MCP
apple (Selvarajah et al., 2001), and chilling injury of The effects of 1-MCP on disorders of fruit and veg-
citrus fruit (Dou et al., 2005). Reduced browning in etables can be categorized into several types:
avocados was associated with reduced polyphenol oxi-
dase (PPO) and peroxidase (POD) activities (Pesis et al., a. Disorders that are associated with senescence and
2002; Hershkovitz et al., 2005). therefore are prevented by inhibition of ethylene
In contrast, the chilling injuries, internal browning, production. These include disorders such as senes-
flesh woolliness and reddening were increased by cent breakdown of apple fruit, which are reduced by
1-MCP treatment in peaches and nectarines (Dong et 1-MCP-induced slowing of senescence processes.
al., 2001b; Fan et al., 2002; Girardi et al., 2005). Deve- b. Chilling-related disorders that are inhibited when
lopment of woolliness was associated with lower accu- ethylene production is prevented. Examples include
mulations of transcripts for ACO, and associated superficial scald of apples, and internal flesh
decreases of those encoding PG, PE, while that of browning of avocado and pineapple.
EGase was increased. 1-MCP enhanced these effects c. Chilling-related disorders that are increased by
suggesting that a certain level of ethylene production by inhibition of ethylene production. Examples include
fruit is required for normal ripening after storage (Dong woolliness (mealiness) and internal breakdown in
et al., 2001b). A slight increase in internal reddening peach and nectarines, and chilling injury in citrus,
occurred in 1-MCP-treated plums (Dong et al., 2002), suggesting that these disorders are aggravated if
but no effects of 1-MCP on internal browning and flesh normal ripening processes mediated by ethylene are
gelling, have been reported. 1-MCP-treated ‘Shamouti’ prevented.
oranges (Porat et al., 1999) and banana fruit (Jiang et al., d. Ethylene-induced disorders that are inhibited by
2004a) are also more sensitive to chilling injury. In 1-MCP application. These include browning
untreated banana fruit stored at chilling-inducing tempe- responses in lettuce, increases in undesirable chem-
ratures, the development of injury was associated with ical changes such as isocoumarin accumulation in
decreased ethylene binding. 1-MCP treatment further carrots, and water-soaking of watermelon.
decreased ethylene binding, and Jiang et al. (2004a) e. Disorders associated with CA storage, e.g., CO2
suggested that enhanced chilling injury of bananas was injury of apple fruit, which is enhanced by 1-MCP.
3.8. Pathological storage disorders opment. However, sensitivity can be beneficial against
some pathogens but deleterious to resistance against
1-MCP increased disease susceptibility of avocado other pathogens. Small amounts of endogenous ethylene
(Hofman et al., 2001; Adkins et al., 2005; Woolf et al., may be necessary to maintain basic levels of resistance to
2005), custard apple, mango and papaya (Hofman et al., environmental and pathological stress because of its
2001). Treatment of strawberry fruit with higher 1-MCP involvement in regulation of plant defense genes
concentrations also increased decay (Ku et al., 1999). In (Marcos et al., 2005).
citrus fruit, 1-MCP inhibited decay at low concentra-
tions, but enhanced it at high concentrations (Dou et al., 4. Summary
2005). Also, in citrus, 1-MCP increased mold rots caused
by Penicillium digitatum and P. italicum (Porat et al., The discovery and subsequent commercialization of
1999; Marcos et al., 2005) and stem rots caused by 1-MCP has provided exciting opportunities for posthar-
Diplodia natalensis (Porat et al., 1999). In contrast, vest scientists to gain insight into the fundamental
Mullins et al. (2000) found no effect of 1-MCP on processes that are involved in ripening and senescence
progression of P. digitatum growth on innoculated of fruit and vegetables. Prospects for commercialization
grapefruit. of 1-MCP for several products appear high. For products
Susceptibility of apples to bitter rot (Colletotrichum such as most vegetables and perhaps non-climacteric fruit,
acutatum) and blue mold (P. expansum) was higher in where further senescence (e.g., yellowing) will decrease
1-MCP treated than untreated fruit (Janisiewicz et al., product value, 1-MCP applications that prevent any
2003). 1-MCP slightly increased severity of decay in change are desirable. However, for products such as many
fruit inoculated with P. expansum, especially when used climacteric fruit, success will be based on delaying rather
in conjunction with other stress treatments such as pre- than preventing ripening, in order to provide a product
storage heat (Leverentz et al., 2003). Saftner et al. (2003) that meets consumer requirements. Data obtained thus far
found that 1-MCP with or without pre-storage heat for the apple show that the issues associated with
treatment reduced decay due to wound-inoculation by P. commercialization are not trivial and provide an exciting
expansum, Botrytis cinerea, C. acutatum at the time of era for postharvest researchers as they aid horticultural
harvest and after CA storage, probably by maintaining industries to realize the full potential for 1-MCP. Under
firmness and thereby resistance to infection. commercial conditions, handling practices will need to
Decay incidence of peaches after inoculation with P. take into account many factors including commodity type,
expansum was slightly reduced by 1-MCP treatment, and cultivar, maturity or ripeness stage, time between harvest
it was suggested that resistance in these fruit was related and treatment, treatment temperature, and desired effects
to higher activities of PAL, PPO and POD (Liu et al., on quality. Each of these factors can affect the 1-MCP
2005). Reduced decay caused by brown rot, Monilinia concentrations that should be applied to each product.
laxa, was found in 1-MCP-treated plums (Menniti et al., Whether 1-MCP concentrations that will allow beneficial
2004). In strawberry, decay was more rapid in 1-MCP delays in ripening, while still allowing positive aspects of
treated fruit (Bower et al., 2003), while Ku et al. (1999) ethylene action to occur, can be optimized to meet
found inhibited decay at low 1-MCP concentrations and conditions associated with pre- and post-harvest varia-
enhanced decay with treatments of 500–1000nl l− 1. tions of some products is still uncertain.
Increased disease susceptibility in fruit treated with high Commercial utilization of this technology will also be
1-MCP concentrations may be associated with lower a function of the cost of 1-MCP application relative to its
phenylalanine ammonia lyase (PAL) activity and lower benefits for each product. The cost / benefit ratio will be
phenolic contents (Jiang et al., 2001). affected by many factors including the product response,
Relatively little about 1-MCP effects on disease especially in relation to quality as perceived by the
incidence is known, but it is likely to become an im- consumer, how successfully 1-MCP use can be incor-
portant factor in the less ideal environments that exist in porated into handling, storage and transport systems, the
the commercial world compared with laboratory-based scale of the industry involved, competition in the
experimental systems. Factors that influence the effects marketplace, and whether it provides access to markets
of 1-MCP on disease development are likely to be that are not available using current technologies.
specific to the product and its interaction with the spe- In summary, the availability of 1-MCP is likely to
cific pathogen and the environment. Delayed ripening have a dramatic impact on our understanding of the
associated with reduced ethylene production may in- involvement of ethylene in plant metabolism as well as
crease product resistance to infection and lesion devel- the storage and handling of horticultural products.
Senescence and ripening processes are complex, how- Bower JH, Blasi WV, Mitcham EJ. Effects of ethylene and 1-MCP on
ever, and even more under commercial conditions where the quality and storage life of strawberries. Postharvest Biol
Technol 2003;28:417–23.
many other factors come into play. As shown for the Bregoli AM, Ziosi V, Biondi S, Rasori A, Ciccioni M, Costa G, et al.
apple, the strengths and limitations of this compound “in Postharvest 1-methylcyclopropene application in ripening control
the real world” will only be fully appreciated when it of ‘Stark Red Gold’ nectarines: temperature-dependent effects on
becomes commercially available for other horticultural ethylene production and biosynthetic gene expression, fruit
quality, and polyamine levels. Postharvest Biol Technol
products around the world.
2005;37:111–21.
Budu AS, Joyce DC. Effect of 1-methylcyclopropene on the quality of
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Biotechnology Advances 24 (2006) 389 – 409

Research review paper

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2.5. Peach and nectarine (Prunus persica L.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 396

1. Introduction ones, as well as senescence of vegetative tissues, has

C.B. Watkins / Biotechnology Advances 24 (2006) 389–409

2.2. Avocado 2.3.1. Ripening physiology and quality

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