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Effect of O2 and CO2 partial pressure on selected phenomena affecting fruit and
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DOI: 10.1016/S0925-5214(98)00092-1

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 Postharvest Biology and Technology 15 (1999) 293 – 303

Effect of O2 and CO2 partial pressure on selected phenomena

affecting fruit and vegetable quality

R.M. Beaudry *
Department of Horticulture, Michigan State Uni!ersity, East Lansing, MI, USA

Received 30 June 1998; received in revised form 6 November 1998; accepted 11 November 1998

Abstract

It is likely that from the time of the Roman Empire and perhaps before, people involved in the storage of plant
material as food recognized that atmospheric modification can provide some benefit in improving storability.
However, active, commercial modification of the atmosphere for the preservation of fresh fruit and vegetables dates
to the early part of this century. Early successes with apple fruit has lead to the attempt to apply modified
atmospheres to a wide range of commodities. Responses to atmospheric modification are found to vary dramatically
among plant species, organ type and developmental stage and include both unwanted and beneficial physiological
responses. Desirable responses include a reduction in respiration, a reduction in oxidative tissue damage or
discoloration, a reduction in the rate of chlorophyll degradation and a reduction in ethylene sensitivity with the
concomitant reduction in the rate of ripening and other ethylene-mediated phenomena. Undesirable responses have
included the induction of fermentation, the development of disagreeable flavors, a reduction in aroma biosynthesis,
the induction of tissue injury and an alteration in the makeup of microbial fauna. The physiological bases for some
of these responses to elevated CO2 and reduced O2 are discussed. © 1999 Elsevier Science B.V. All rights reserved.

Keywords: Oxygen; Carbon dioxide; Ethylene; Respiration; Controlled-atmosphere; Modified-atmosphere

O2 and CO2 are biologically-active molecules of
importance in primary and secondary metabolic
processes in plants. Their global influence on
metabolism, has lead to their use for modification
of plant behavior for the purpose of extending
storage duration and shelf-life. The first recorded
* Tel.: + 1-517-353-3769; fax: +1-517-353-0890; e-mail:
beaudry@pilot.msu.edu.
instance of the occurrence of modified atmosphere
use in agriculture was in the Roman era (Varro,
1800) although the technique likely dates back
several thousand years previously in the Middle
East (Kays, 1991). Underground pits or silos used
for the storage of grains developed ‘foul air’ such
that entry into the pits could be dangerous. Evi-
dently, the respiratory activity of the plant mate-
rial reduced the O2 and elevated CO2 sufficiently

0925-5214/99/$ - see front matter © 1999 Elsevier Science B.V. All rights reserved.
PII: S 0 9 2 5 - 5 2 1 4 ( 9 8 ) 0 0 0 9 2 - 1
294 R.M. Beaudry / Posthar!est Biology and Technology 15 (1999) 293–303

to create an environment that would not sustain phy in the early 1960s (Meigh et al., 1960). Burg
insect or animal life, thus preserving the grain. In and Burg (1967) ascertained that reduced partial
addition to grain, Cato recommended the storage pressures of O2 and enhanced partial pressures of
of grapes in clay pots (Varro, 1800), which may CO2 resulted in an inhibition of ethylene re-
have altered the atmospheric levels of O2 and CO2 sponses. The inhibition was interpreted to be in a
and would have certainly elevated humidity levels. manner consistent with an enzyme kinetic model
Subsequently, Brookes (1763) recounted a tech- in which a substrate (O2) must bind to a receptor
nique used to preserve mature, but not fully ripe before a dissociable activator (C2H4) can attach.
fruit by enclosing them in a glass vessel with an Their data suggested the partial pressure of oxy-
oiled paper membrane over the opening. Berard gen at which the response of ethylene is inhibited
(1821) conducted the first scientific studies, finding by 50% (Ks) is approximately 2.8 kPa. The data
that fruits utilize O2 and produce CO2 and that can also be depicted as the effect of oxygen on the
eliminating the O2 in the storage environment ethylene level (KA) required to generate half-maxi-
would prevent ripening. Over 100 years later, mal response (Fig. 1, inset). These data can be
Kidd and West (1927) demonstrated the effect of represented as a series of curves in which O2
reduced O2 and elevated CO2 in the storage envi- reduces the response to ethylene, with the effect
ronment would prolong the storage life of apple
being more pronounced as the ethylene level de-
fruit. Two years after this paper was published,
clines (Fig. 1). In comparison to the O2 sensitivity
the first controlled-atmosphere (CA) storage was
of ethylene action, oxygen uptake by the terminal
constructed and used for apple storage in England
oxidase of the electron transport chain of the
(Kays, 1991).
mitochondria is reduced by 50% only when inter-
The thesis of Kidd and West on the effects of
nal O2 levels reach approximately 0.25 kPa or 3
O2 and CO2 grew out of their early work on the
effect of atmosphere composition on reducing res- mmol ×m − 3 (Yocum and Hackett, 1957; Map-
piration and metabolic activity of white mustard son and Burton, 1962; Tucker and Laties, 1985).
seeds (Fidler, 1965). On the basis that storage life Additionally, Burg and Burg (1967) determined
was governed by metabolic activity they initiated that CO2 acts in a manner consistent with an
investigations into the use of atmosphere modifi-
cation on apple storability. The results of these
and subsequent studies was that elevating CO2
and suppressing O2 ‘‘had the expected effect of
depressing respiration, and thus, on the basis that
storage life is governed by the rate of metabolic
activity, CA conditions extend storage life’’ (Fi-
dler, 1965). Fidler (1965) further qualified this
remark, noting that it was more a description of
‘what happens’, and does not answer the question
of ‘how it happens’. Interestingly, Kidd and West
(1945) previously suggested the alternative possi-
bility that O2 depletion might also interfere with
the production and action of ethylene, although
conclusive supporting data were lacking.

1. Ethylene responses
The comment by Kidd and West was, perhaps,
telling; following the advent of gas chromatogra-
Fig. 1. Effect of external oxygen levels on the response of pea
seedlings to ethylene. Inset: the effect of internal O2 on the
concentration of ethylene required to give a response half that
of the maximal response KA.
 R.M. Beaudry / Posthar!est Biology and Technology 15 (1999) 293–303
Fig. 2. Recommended O2 and CO2 combinations for the
storage of vegetables. The shaded area depicts atmospheres
theoretically attainable by MAP using low density
polyethylene (LDPE, lower boundary) and perforated pack-
ages (upper, dashed line) as redrawn from Mannapperuma et
al. (1989). The darkened area represents atmospheres observed
in commercial MA packages of mixed lettuce-based salads
(Cameron et al., 1995).
inhibitor of ethylene binding, reducing ethylene
action 50% at 1.55 kPa. They concluded that, in
the case of climacteric fruits, O2 and CO2 likely
act to delay ripening, not solely through their
influence on respiration per se, but largely
through their inhibitory effects on ethylene action.
These findings have been borne out by recent
work on the influence of the ethylene binding
inhibitor 1-methylcyclopropene (Serek et al.,
1995) and the development of ethylene binding
site-deficient mutants (Bleeker et al., 1988; Lana-
han et al., 1994) where the absence of ethylene
perception and action prevents ripening and most
of the attendant changes in physiology.
2. O2 and CO2 tolerances
The focus of postharvest technology on the use
of atmospheric modification to improve shelf-life
has lead to the investigation of the effects of low
295
O2 and elevated CO2 partial pressures on a wide
range of fruits and vegetables. Inasmuch as the
biological factors limiting the shelf life of these
varied commodities differ (i.e. not all are related
to ethylene responsiveness), the success of atmo-
sphere modification has been varied and the range
of recommended atmospheres varies widely as
well (Gorny, 1997; Kader, 1997a; Reid, 1997;
Saltveit, 1997). For example, atmosphere recom-
mendations for vegetables range from ambient O2
levels and 10–15 kPa CO2 for asparagus to 1.5–
2.5 kPa O2 and 0 kPa CO2 for lettuce (Fig. 2;
Mannapperuma et al., 1989).
Technologies used to apply modified atmo-
spheres include the aforementioned CA as well as
the use of specialized packaging. Films that are
highly permeable to O2 and CO2 are used in a
technique known as modified-atmosphere packag-
ing (MAP) to establish modified atmospheres
within packages containing horticultural products
as the product’s respiration depletes O2 and con-
tributes to the headspace CO2. In that atmosphere
modification is largely passive in MAP, there are
constraints on the possible combinations of O2
and CO2 due to the nature of gas permeation
through the package (Cameron et al., 1994, 1995);
an optimal atmosphere, therefore, may not be
attainable (Figs. 2 and 3). In packages composed
of low density polyethylene (LDPE), for instance,
the permeability to O2 is approximately 25% of
that to CO2. If respiration is aerobic and the
respiratory quotient (RQ) is one, then the steady-
state partial pressure gradient of O2 will be about
four times that of CO2. The combinations of
atmospheres attainable in packages composed of
permeable films having no perforations are along
a line between 21 kPa O2, 0 kPa CO2 and 0 kPa
O2, ! 3–5 kPa CO2 as long as the rate of O2
uptake and CO2 production are equal. If the level
of O2 is sufficiently low, however, fermentation
will occur, resulting in an elevated RQ and signifi-
cantly higher CO2 levels in the package than those
predicted (see data for commercial lettuce pack-
ages, Fig. 2). In packages that solely rely on
perforations for atmosphere modification, O2 en-
ters the package at the nearly the same rate that
CO2 leaves the package. The combinations of
296 R.M. Beaudry / Posthar!est Biology and Technology 15 (1999) 293–303
Fig. 3. Recommended O2 and CO2 combinations for the
storage of fruit. The shaded area depicts atmospheres theoret-
ically attainable by MAP by film permeation alone (low
density polyethylene, LDPE, lower boundary) and via perfora-
tions alone (upper, dashed line) or their combination (shaded
area). Data are adapted from Kader (1997a,b).
atmospheres attainable in perforated packages are
along a line between 21 kPa O2, 0 kPa CO2 and 0
kPa O2, 21 kPa CO2 as long as the rate of O2
uptake and CO2 production are equal. Packages
that rely on both perforations and permeation can
potentially achieve combinations of O2 and CO2
in the area between the two lines described. The
effect of temperature on package atmospheres is
worthy of special mention in that packages that
generate low O2 levels near the fermentation
threshold at low temperatures can develop O2
partial pressures that will induce fermentation as
temperature increases (Beaudry et al., 1992;
Cameron et al., 1994). This effect is greater on
packages relying on the diffusion of gases through
perforations for atmosphere modification than for
those dependent upon permeation of gases
through the film barrier (Cameron et al., 1994).
Additionally, variation in temperature, package
thickness and product respiration can contribute
to variability that can result in the generation of
harmful atmospheres in a portion of the packages
in any given population, with the proportion that
are ‘at risk’ increasing as the target atmospheres
are nearer to the tolerance limits of the commod-
ity (Cameron et al., 1993).
It is important to recognize that extremely low
O2 levels or excessively high CO2 levels that in-
duce fermentation (Thomas, 1925) can result in
the generation of off-flavors (Cohen et al., 1990;
Dostal-Lange and Beaudry, 1991; Richardson and
Kosittrakun, 1995) or visible tissue damage
(Lidster et al., 1990). There is an interaction be-
tween O2 and CO2 in that elevated CO2 levels
make plant material more sensitive to low levels
of O2 such that the fermentation threshold occurs
at higher O2 partial pressures as the partial pres-
sure of CO2 increases (Thomas, 1925; Beaudry,
1993).
There are occasions, however, when the ‘opti-
mal’ atmosphere for a commodity, based on its
physiological tolerance to O2 and CO2, is less
desirable than a stressful atmosphere (i.e. an at-
mosphere sufficiently low in O2 or high in CO2 to
have the potential to induce fermentation, off-
flavor or tissue damage) that confers a benefit that
outweighs the risk of a loss in quality or value
incurred by the application of the extreme atmo-
sphere. Within the apple storage industry, for
instance, storage operators have recently been
successful in preventing the storage disorder su-
perficial scald on ‘Delicious’ apples in CA storage
using O2 partial pressures as low as 0.7 kPa (Lau,
1997), which is very near the fermentation
threshold for that cultivar (Gran and Beaudry,
1993). A high partial pressure of CO2 (10–20 kPa)
is able to retard fungal growth and spore germi-
nation (Brown, 1922) and is thereby able to en-
hance storage life of a number of commodities
(Thornton, 1930; Brooks et al., 1932). This effect
of CO2 forms the basis for the relatively common
practice of supplementing the CO2 in the atmo-
sphere surrounding blackberry, raspberry, cherry
and strawberry fruit during shipment in the USA.
High levels of CO2 are also useful for the preven-
tion of chlorophyll degradation in a number of
tissues. Extreme levels of O2 ( " 0.5 kPa) and CO2
( #40 kPa) also have potential for use in quaran-
tine treatments for insect control and is being
assessed as a replacement for the fumigant methyl
bromide (Aharoni et al., 1979; Mitcham et al.,
1997). One interesting application of extreme at-
 R.M. Beaudry / Posthar!est Biology and Technology 15 (1999) 293–303 297

mosphere is in the prevention of browning of cut

lettuce in prepared salad products placed in
modified-atmosphere packages (Lee et al., 1996).
In this case, while the physiologically optimal
storage O2 level is 1 kPa or greater (Mannappe-
ruma et al., 1989), packages are designed to gen-
erate less than 0.5 kPa to prevent browning
during shipping and the retail holding period,
resulting in considerable fermentation (Cameron
et al., 1995; Fig. 2). Fermentation of packaged
lettuce, however, results in the generation of min-
imal amounts of malodorous volatiles unless the
product is seriously mishandled (Smyth et al.,
1998). Thus, the product is intentionally stressed
by an extreme atmosphere, but the loss in its
aromatic quality is much less than the gain in
visual quality. Examples such as this reinforce the
importance of quantifying plant responses to ex-
tremes of atmosphere modification, especially as
new techniques and technologies are being
adopted for fruit and vegetable storage. Fig. 4. Response of asparagus to the range of O2 partial
pressures below ambient. The response to O2 is expressed as
changes in adenylate energy charge (AEC, upper graph) as
defined by Atkinson, (1968) where AEC = ([ATP] +0.5×
3. Primary metabolism responses [ADP]) ×([ATP]+[ADP]+ [AMP]) − 1, and pyruvate kinase
activity (lower graph). The response has been divided into two
In this regard, the response of primary zones: that which can sustain active metabolism, termed a
metabolism (processes of glycolysis, fermentation homeostatic zone, and that which cannot, termed a crisis zone.
and aerobic respiration) to atmosphere modifica- The homeostatic zone is divided into two parts: at O2 partial
pressures below approximately 4 – 5 kPa, is a zone of elastic
tion bears further attention. It is well established stress. The crisis zone has been divided into two parts, below
that low O2 and elevated CO2 partial pressures approximately 1 kPa O2. Data are adapted from Silva (1998).
can reduce the rate of respiration as previously
mentioned. Further, for some plant materials, the found to be causally linked to cell survival (Xia et
reduction in metabolism accompanying reduced al., 1995). In addition, Silva (1998) found that a
respiration can apparently improve storability, al- number of important glycolytic enzymes under-
though for non-climacteric tissues, this is the ex- went marked changes across this O2 range. It is
ception, rather than the rule. Recent findings by suggested that the range of O2 levels can be
Silva (1998) for asparagus spears and those of divided into two zones: that which can sustain
Al-Ani et al. (1985) for germinating seeds suggest
active metabolism for an indefinite period, which
that metabolic processes in some plant tissues are
is here termed a homeostatic zone; and that which
responsive a wide range of O2 partial pressures,
cannot, which is termed a crisis zone (Fig. 4).
well above those that limit respiration. The find-
These zones can be visualized by the influence of
ings of Silva (1998) indicate that, at least for
O2 on pyruvate kinase activity (Fig. 4, adapted
asparagus, the level and type of response differs
across the O2 partial pressure range of 0.16–16 from Silva, 1998), which is a critical glycolytic
kPa (Fig. 4). Silva (1998) found that O2 influenced control point that catalyzes the final step in gly-
the adenylate energy charge (AEC), which is colysis (Plaxton, 1996). The homeostatic zone
sometimes used as an indicator of metabolic activ- might be further divided into two parts. At high
ity (Saglio et al., 1980), although it has not been O2 partial pressures (#5 kPa O2), the plant mate-
298 R.M. Beaudry / Posthar!est Biology and Technology 15 (1999) 293–303

rial is metabolically active and able to generate Table 1

sufficient energy, as indicated by the AEC, to Generalizations regarding the effects of O2 levels below ap-
proximately 5 kPa and CO2 levels greater than 5 kPaa
sustain most of its metabolic processes. At the
lower levels of O2 in the homeostatic zone (be- Process or attribute af- Atmospheric modificationb
tween 2 and 4 – 5 kPa O2 in Fig. 4), it is suggested fected
that an elastic stress zone is present in which O2
Reduced O2 Elevated CO2
uptake begins to decline (data not shown) and
marked metabolic readjustments must take place 1. Primary metabolism
to match carbon flux with O2 uptake. Below this Respiration rate − −, 0, or +
level of O2, it is suggested that energy levels begin Starch breakdown − −
Sugar consumption −, 0, or + −, 0, or +
to decline rapidly as indicated by the drop in the
AEC. It is possible that asparagus tissues attempt 2. Secondary
metabolism
to enhance carbon flux, despite a shortage in Ethylene metabolism
required energy, critical glycolytic control en- ACC synthase activ- − −
zymes are increased in (extractable) activity. As ity
ACC oxidase activity − − or +
the O2 level declines still further, the activity of
ACC synthase syn- − −
these critical enzymes declines, perhaps due to thesis
insufficient production of carbon skeletons as car- ACC oxidase synthe- − −
bon flux declines and reduced energy available for sis
Ethylene perception − −
the synthesis of proteins. In this zone, fermenta- Pigment metabolism
tion is induced in what might be the only mecha- Chlorophyll degrada- − −
nism left to the plant material to generate energy tion
needed for survival. The data of Silva (1998) Anthocyanin develop- − −
ment
further suggest that CO2 affects this relationship, Carotenoid biosyn- − −
but most markedly at levels of CO2 of 20 kPa and thesis
above. Even so, Silva (1998) noted also that low Phenolic metabolism
levels of CO2 (i.e. approximately 5 kPa), can Phenylalanine ammo- − +
nia lyase activity
reduce the rate at which sugars are utilized and Total phenolic con- − −
alter the extractable activities of a number of tent
enzymes critical to glycolysis. Polyphenol oxidase − −
activity
Cell wall metabolism
Polygalacturonase ac- − −
4. Secondary metabolism responses tivity
Soluble polyuronide − −
content
Apart from effects on ethylene response and the
Volatile compound
respiratory pathway, O2 and CO2 may affect metabolism
other processes associated with the quality of the Ester biosynthesis − − or 0
treated commodity (Kader, 1997b; Table 1). Terpenoid biosynthe- − −?
sis
Among these, perhaps three metabolic processes Fermentative volatile + ($1 kPa) + (%20 kPa)
deserve further attention: pigment metabolism, biosynthesis
phenolic metabolism and volatile compound 3. Decay
metabolism. Fungal growth − ("1 kPa) − (#10 kPa)
Amongst the pigments affected by atmospheric Bacterial growth − or 0 − or 0
modification, chlorophyll is most broadly associ- a
Responses are dependent upon the commodity, cultivar,
ated with fruit and vegetable quality. Chlorophyll
temperature, treatment duration and the presence of interac-
loss may be desirable for the edible product, as in tions between O2 and CO2 (Adapted from Kader, 1997b).
the case of many climacteric fruit, but it may also b
(−) decrease or inhibit; (0) no effects; (+) stimulate or
be a quality defect, as in the case of senescence-re- increase; and (?) insufficient data.
 R.M. Beaudry / Posthar!est Biology and Technology 15 (1999) 293–303
lated chlorophyll loss in green vegetables. In the
instance of climacteric fruit, chlorophyll degrada-
tion can be inhibited by low O2 and elevated CO2,
primarily by virtue of the effects of these
molecules on ethylene sensitivity. Similarly, senes-
cence-related chlorophyll degradation in non-cli-
macteric vegetative tissues can be reduced by low
O2 and elevated CO2 (Saltveit, 1997). In the case
of broccoli, the atmospheres most beneficial for
storage are similar to those used for CA storage
of climacteric fruits (Makhlouf et al., 1989a) and
ethylene synthesis by broccoli under these atmo-
spheres is much reduced (Makhlouf et al., 1989b).
The specific inhibitor of ethylene action, 1-methyl-
cyclopropene (Serek et al., 1995), can temporarily
prevent degreening of broccoli (Mir and Beaudry,
unpublished data) and added ethylene or propy-
lene, an ethylene analogue, can enhance degreen-
ing in a dose-dependent manner (Tian et al.,
1994). It seems likely that the control of chloro-
phyll loss by low O2 and elevated CO2 in many
green tissues is in part by virtue of the effects on
these gases on ethylene perception, however, a
complete understanding of the mode of action of
CO2 in the reduction of chlorophyll loss is
lacking.
Brown pigments accumulate on the cut surfaces
of some fruits and vegetables and can present a
significant quality defect. Pigment formation in
these cases can be influenced by the composition
of the atmosphere. The brown pigments form
primarily as a result of the action of a single
enzyme, polyphenol oxidase (PPO) on plant phe-
nolics. PPO oxidizes a wide range of substrates
including monophenols, triphenols, ascorbic acid,
and o- and p-diphenols (Mayer and Harel, 1979).
The oxidation products of these reactions react
with each other to form high molecular weight
polymers and form macromolecular complexes
with amino acids and proteins, which leads to the
formation of brown pigments (Vámos-Vigyázó,
1981). The oxidation process requires the presence
of molecular oxygen and can be prevented by
reduced O2 levels. The Km for O2 is approximately
7–10 kPa (Mayer and Harel, 1979). While the
rate of oxidation can be reduced by low O2, the
reaction is sufficiently rapid that reduced O2 has
little effect if the storage duration is significant.
299
For instance, browning of an apple slice can take
place in 5–10 min. Oxygen levels at or near the
lower O2 limit of 0.3–0.5 kPa for apple slices
(Lakakul, 1994; Lakakul et al., 1999), would per-
mit the same extent of browning to occur in
approximately 4 h. Indeed, there were no differ-
ences in the extent of browning of apple slices
stored at 1 and 20 kPa O2 when held for 24 h at
0°C Lakakul (1994).
The production of volatile esters, which con-
tribute to characteristic aromas of a number of
fruit including apple, banana, pear, peaches,
strawberries and others, are affected by atmo-
sphere modification (Lidster et al., 1983; Golias,
1984; Shamaila et al., 1992; Fellman et al., 1993;
Song et al., 1998; Perez and Beaudry, unpublished
data). Aroma compounds that confer characteris-
tic odors are generally suppressed by low O2, in
part by the action of O2 on ethylene action in
climacteric fruits, but is also likely to be via action
of O2 on oxidative processes including respiration.
Song et al. (1998) found an O2 dependency of
ester production in apple that was similar to the
O2 dependency of respiration (Fig. 5). Perez and
Beaudry (unpublished data) found that ester pro-
duction in strawberry, a non-climacteric crop, was
more linearly related to O2 partial pressure. The
effect of CO2 on ester production is unclear. Streif
and Bangerth (1988) found CO2 exposure during
CA storage of apple fruit led to a significant
reduction in volatile production following storage.
However, Song et al. (1998) found no effect of
CO2 on ester biosynthesis while apple fruit were
held in the modified atmosphere. In pear, there is
some indication that low O2 may also alter ter-
penoid metabolism in that the production of !-
farnesene (a volatile sesquiterpene) is reduced by
low O2 and elevated CO2 (Chen et al., 1993).
Similar results have been found for apple (Song
and Beaudry, unpublished data).
Apart from the effects of O2 and CO2, there
exist a number of other biologically active
molecules other than ethylene that have some
promise for application in storage environments.
Among these are acids, aroma compounds and
aroma precursors that have antifungal activity
(Aharoni and Stadelbacher, 1973; Avissar et al.,
1989; Hamilton-Kemp et al., 1992; Vaughn et al.,
300 R.M. Beaudry / Posthar!est Biology and Technology 15 (1999) 293–303

Fig. 5. Effects of O2 and CO2 partial pressures on ethanol (A), ethyl acetate (B), hexyl acetate (C) and 2-methylbutyl acetate (D)
content in packages of ‘Starkrimson Delicious’ apples after 2 months storage at 0°C.

1993; Sholberg and Gaunce, 1995; Song et al.,
1996). In some cases, the applied compounds have
the potential to alter the aroma characteristics of
the commodity, but the effect is usually not per-
manent and may enhance aroma to some extent
(Song et al., 1996).
5. Summary
Future research and development needs include
the development of more detailed information on
the effects of O2 and CO2 at the biochemical and
molecular levels. In each of the metabolic pro-
cesses discussed here, there are important gaps in
information. Perhaps the one area that has the
potential to most immediately impact the quality
of fruit and vegetables pertains to the effects of
atmosphere modification on aroma. Additional
advancement in polymers to refine control over
O2 and CO2 permeability is needed to make MAP
a more widely applicable alternative to CA
storage.
In some respects, the application of modified
atmospheres to horticultural products, at least in
regard to O2 and CO2, has advanced little since
the days grain was first stored in sealed pits or
clay jars. While our understanding of the re-
sponses of plant material to modified atmospheres
has certainly improved, in the final analysis, the
economic and nutritional benefits we have ob-
tained since those early days have been almost
entirely due to trial-and-error experimentation.
We have simply taken advantage of the same
technique for food preservation in increasingly
complex and technologically advanced manners.
One might ask if there are further opportunities in
the use of atmosphere modification we may be
able to realize. In some regards, significant oppor-
tunity to alter the response of plant tissues to O2
and CO2 may exist via molecular modification. It
has been proposed, for instance, that within
plants there exists at least one O2 receptor in
addition to the terminal oxidase of the electron
transport pathway (Mapson and Burton, 1962;
Tucker and Laties, 1985; Solomos, 1997). The
potential for the effects of atmosphere modifica-
tion may expand if such a receptor/sensor can be
identified and manipulated. One might envision
inducing fruit in ambient O2 to ‘perceive’ that
they are in a low O2 partial pressure and thereby
inducing the attendant low O2 responses. While
 R.M. Beaudry / Posthar!est Biology and Technology 15 (1999) 293–303
this possibility may or may not be realizable, such
an advance would signal our conceptual entry
into a new era in food preservation.
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 Postharvest Biology and Technology 16 (1999) 199

Erratum

Erratum to ‘‘Effect of O2 and CO2 partial pressure on

selected phenomena affecting fruit and vegetable quality’’
[Postharvest Biology and Technology 15 (1999) 293–303]!
R.M. Beaudry *
Department of Horticulture, Michigan State Uni!ersity, East Lansing, MI, USA

The Publisher regrets that Fig. 3 of this article was printed incorrectly. The correct version follows.
.

Fig. 3. Recommended O2 and CO2 combinations for the

storage of fruit. The shaded area depicts atmospheres theoret-
ically attainable by MAP by film permeation alone (low
density polyethylene, LDPE, lower boundary) and via perfora-
tions alone (upper, dashed line) or their combination (shaded
area). Data are adapted from Kader (1997a,b).
* Tel.: +1-517-353-3769; fax: +1-517-353-0890.
E-mail address: beaudry@pilot.msu.edu (R.M. Beaudry)
!
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