The Basics of Respiration for Horticultural Products

Oluwafemi J Caleb, Werner B Herppich, and Pramod V Mahajan, Leibniz Institute for Agricultural Engineering (ATB), Potsdam,
Germany
Ó 2016 Elsevier Inc. All rights reserved.

Introduction 1
Growth and Maintenance Respiration 2
Factors Affecting Respiration Rate 3
Maturity or Developmental Stage 4
Temperature 4
Chilling Stress 4
Atmospheric Composition 4
Other Postharvest Handling Stress 5
Measurement of Respiration Rate 5
Acknowledgments 6
References 7

Introduction

Horticultural crops after harvest are alive and continue all the characteristic functions of living organisms. One of the most impor-
tant of these processes is respiratory metabolism. Respiration denotes a complex catena of highly regulated and interactive meta-
bolic processes distributed among different cellular compartments. Respiration not only produces ATP by oxidative
decarboxylation but also provides organic and amino acids for various cellular synthetic reactions. In green tissue, these compounds
are essential in sustaining photosynthetic carbon assimilation (Krömer, 1995; Nunes-Nesi et al., 2011). Consequently, during the
day, mitochondrial respiration is still highly active retaining up to 50% of its nocturnal activity in these plants (Herppich and
Peckmann, 2000).
If fat or lipids are not used as substrates, the various steps of respiration normally include the primarily conversion of sugars into
glucose and start with the glycolytic breakdown into pyruvate within the cytoplasm. The latter enzyme-catalyzed pathway conserves
the high internal energy of sugars in the biochemical energy-mediating compound ATP (adenosine triphosphate) and in the elec-
tron-mediating compound NADH2 (reduced nicotinamide adenine dinucleotide). The mitochondria, another cell compartment,
house two other important processes involved in respiration, namely the tricarboxylic acid (TCA) cycle (also known as Krebs or
citric acid cycle) and the respiratory electron transfer system. The TCA cycle is the site of citric acid synthesis and oxidation and
is involved in the formation and breakdown of malic acid. The TCA cycle is, in general, an important switch of various organic acids
but also releases the electron-mediating metabolites NADH2 and FADH2. Also very importantly, in several steps of the TCA cycle,
CO2 is released.
The mitochondrial respiratory electron transfer system or ‘cytochrome pathway’ uses the power of electrons provided by
NADH2 and FADH2 to generate a transmembrane proton gradient, which itself drives the conversion of ADP to the biochemical
energy–mediating compound, ATP. The final enzyme of the complex respiratory electron chain, the cyanide-sensitive cytochrome
oxidase, reduces in particular atmospheric oxygen (O2) to water. Rates of production of ATP and, hence, biochemically usable
energy, can be fine-tuned to the exact metabolic needs by several shortcuts of the entire electron transport chain, the cyanide-
insensitive (cyanide-resistant) ‘alternative pathway’ and the rotenone-insensitive (rotenone-resistant) pathway (Buchanan et al.,
2000; Peckmann et al., 2012).
Generally speaking, the entire respiration processes involve the production of carbon dioxide (CO2) and water at the ‘expense’ of
atmospheric oxygen. Respiration reactions also liberate quite an amount of heat, so-called respiratory heat or ‘vital heat.’ In partic-
ular in harvested, stored produces, this heat creates a microenvironment with higher temperatures around the commodity if it is not
effectively managed by refrigeration or cool ventilation. This scenario highlights the critical importance of maintaining an optimum
cold chain system for horticultural commodities as the heat produced during respiration contributes to the refrigeration load and
must be considered in designing cold chain systems.
Similarly, the CO2 produced as a by-product of respiration can accumulate around the produce or inside the packaging. This may
partially have positive effects on quality maintenance because high CO2 concentrations reduce generally metabolic activity in a kind
of feedback inhibition. In addition it has been shown to reduce germination, growth, and spreading of microbes (Caleb et al.,
2013). On the other hand, if CO2 gets too high, it may also lead to serious tissue damages. Furthermore, at continuous produce
respiration, O2 may deplete in a storage facility. This finally can result in anaerobiosis below the critical O2 limit. Under such condi-
tions, metabolism switches from oxygen-related processes to fermentative metabolism to keep NADH2 and pyruvate concentrations
low and glycolysis running (Ke et al., 1995). In most plants, and hence plant parts, this switch results in undesired production and

Reference Module in Food Sciences http://dx.doi.org/10.1016/B978-0-08-100596-5.21003-2 1

2 The Basics of Respiration for Horticultural Products

Table 1 Classification of horticultural commodities according to their respiration rates

Classification RCO2 (mg kg
1 h
1) at 5  C Commodities

Excessively high >60 Asparagus, mushroom, pea, spinach, sweet corn

Very high 40 to 60 Artichoke, broccoli, snap bean, bean sprouts, Brussels sprouts, cut flowers, endive, green onions,
okra, kale, and watercress
High 20 to 40 Strawberry, blackberry, raspberry, cauliflower, lima bean, avocado, carrot and radish (with tops),
leeks, and lettuce (leaves)
Moderate 10 to 20 Apricot, banana, blueberry, gooseberry, cherry, celeriac, peach, nectarine, pear, plum, fig, olive,
topped radish, cabbage, topped carrot, lettuce (head), cucumber, pepper, tomato, summer
squash, and immature potato
Low 5 to 10 Apple, citrus, grape, beet, celery, cranberry, honeydew melon, watermelon, papaya, kiwifruit,
garlic, onion, sweet potato, matured potato
Very low <5 Nuts, dates, and dried fruits

Source: Kader, A.A., 2002. Postharvest Technology of Horticultural Crops, third ed. University of California Agriculture and Natural Resources Publication 3311; Watkins, C.B.,
Nock, J.F., 2012. Production Guide for Storage of Organic Fruits and Vegetables. NYS IPM Publication No. 10.

accumulation of acetaldehyde and ethanol (in a few cases, lactic acid may also be the resulting fermentative metabolite). Prolonged
anaerobic respiration will render the produce unacceptable for normal consumption (Ke et al., 1995).
The storability and postharvest qualities of horticultural products are significantly influenced by the rates and the conditions of
respiration. Sugars and organic acids (and other value-adding organic substrates) of the produce are consumed during ongoing
respiration. This makes the effort of preserving postharvest quality and extending storage life of fresh produce from farm to
consumers a critical race against time, as the intrinsic factors influencing respiration after harvest are largely a function of the fresh
produce. As a general rule, the rate of postharvest degradation of various fruits and vegetables is attributed to their respective respi-
ration rates as shown in Table 1 (Kader, 2002; Watkins and Nock, 2012). Thus, this article presents a comprehensive overview on
the functional role of respiration in horticultural crops pre- and postharvest. The various factors affecting respiration of horticultural
commodities postharvest and the role of respiratory quotient (RQ) in designing efficient packaging and storage systems are
discussed.

Growth and Maintenance Respiration

The pathway of energy utilization for horticultural commodities can be described by two respiratory components: growth respira-
tion and maintenance respiration as shown in Figure 1 (Thornley, 2011). Growth respiration is defined as the amount of – mostly –
carbohydrates respired and biochemical energy and resulting metabolites used in metabolic processes that lead to a net gain in plant
biomass. This involves the energy required to convert assimilates into new cellular structural components, reductant for biosynthetic
processes, cellular transport, and uptake of nutrient (Chiariello et al., 1989; Iersel and Seymour, 2000). The rate of growth

Figure 1 Description of growth and maintenance respiration for horticultural crops from tree plant to maturity. Respiration rate was described as
a function of relative growth rate (RGR), and maintenance respiration is taken as the rate of respiration when RGR ¼ 0. Source: Kriedemann, P.E.,
Virgona, J.M., Atkin, O.K., 2016. Growth analysis: a quantitative approach. In: Price, C., Munns, R. (Eds.), Plants in Action: A Resource for Teachers
and Students of Plant Science, second ed. Australian Society of Plant Scientists.
 The Basics of Respiration for Horticultural Products 3

respiration is proportional to plant growth, and its constant is referred to as the growth respiration coefficient. It can be used to
calculate the conversion efficiency (Y), based on the amount of plant dry matter (g) that is produced per mass (g) of glucose
(Yg, g g
1) (Adu-Bredu et al., 1997; Iersel and Seymour, 2000).
On the other hand, maintenance respiration is defined as the respiration needed to provide the energy for all plant metabolic
processes that do not result in a net gain in plant dry matter. These include processes such as maintenance of ion gradients across
cellular membranes and reproducing degraded organic compounds (Chiariello et al., 1989). Maintenance respiration represents the
energy required to maintain mature tissue biomass (status quo) when the growth rate is zero (Lavigne and Ryan, 1997; Thornley,
2011). Thus, maintenance respiration can be used to determine the maintenance coefficient, based on the amount of glucose assim-
ilated for maintenance per unit dry mass per day (g g
1 day
1).
Growth respiration can be distinguished from maintenance respiration by relating variation in respiration rates to variation in
relative growth rate (RGR) over short time intervals (Figure 1). This approach assumes a model for respiration where:
Total respiration ¼ Maintenance respiration þ ((Growth respiration, costs of growth)  RGR)
In this case, it is generally assumed that growth and maintenance respiration is proportional to the growth and dry mass of the
plant, respectively. The latter assumption is often questioned, since maintenance respiration depends on both mass and organic
chemical composition. Both the conversion efficiency and maintenance respiration coefficient are important parameters in under-
standing respiration of a horticultural commodity from farm to after harvest. However, after harvest the maintenance respiration
coefficient is the most responsive of the functional components of respiration to environmental change (Ryan, 1991).
Whether or not maintenance respiration plays the major or the only role in this context strongly depends on the type of
commodity. For fully mature fruit, any further development may lead to senescence and decay. Consequently, there is no need
for growth and, hence, growth respiration any longer. In contrast, in metabolically highly active vegetables such as broccoli inflo-
rescences or asparagus spears, i.e., developing and growing shoots, the situation may be quite different. Indeed, for stored (up to
6 days) fresh white asparagus spears, it was shown that the growth respiration component may contribute up to 60% of the entire
respiration activity. This effect is only to a small part depending on storage temperature (Herppich and Huyskens-Keil, 2008). In
this experiment, potential respiration activity declined by about 50% during early storage, then remained more or less constant
virtually irrespective of temperature. The authors found large differences between the actual respiratory demand indicated by
measured rates of respiration and the actual supply, that is, the changes in soluble sugars. The latter was less than 30% of the
cumulated average respiration rate at 20  C. Analyses of the relative changes in soluble sugars and total content of the cell wall
showed that 60% (at 20  C) to 70% (at 5  C) of the soluble sugars that were ‘lost’ may have been used for cell wall enlargement,
i.e., growth.
High CO2 concentrations during storage affected spears’ sugar consumption only at ambient temperature but not at low temper-
ature (10  C). At 20  C, controlled atmosphere conditions suppressed growth respiration partially inhibiting energy supply for
spear toughening, cell wall synthesis, and prevented decline in storage carbohydrates to some extent. In this context, comparing
variations in sugar content and in cell wall components highlighted a close correlation. In harvested asparagus spears, cell wall
synthesis, i.e., growth and not maintenance respiration is the major sink for free soluble carbohydrates. This example clearly points
out that it may be sometimes necessary to take the actual biological function of the respective produce into account.
In growing fruit, both respiration components are active, and their relative contribution highly depends on the respective stage of
fruit development. These effects have been extensively studied in particular in cucumbers (Marcelis and Hofman-Eijer, 1995) and in
tomatoes (Grange and Andrew, 1995). As expected, the maintenance component is low in young fruit (approximately 20%) and
nonlinearly increases with maturation to more than 75% close to harvest. Although maintenance respiration is generally highly and
linearly dependent on temperature, growth respiration is nonlinearly related to temperature and indirectly controlled by the
temperature effect on cucumber fruit growth. Indeed, the growth component was found to be directly proportional to volume
expansion rates of tomato fruit, as manipulated by temperature or by adjusting the plants’ source–sink relationship.
RQ refers to the ratio between the concentration of CO2 produced to O2 consumed (utilized) during the respiration process of
horticultural products. Generally, the organic and inorganic content of a horticultural commodity influences the substrates used
during respiration, and this reflects on calculated RQ value. Furthermore, depending on the utilized substrate during respiration,
RQ values range from 0.7 to 1.3 for aerobic respiration (Caleb et al., 2013). For example, when carbohydrates are being utilized
under aerobic respiration, the RQ value is approximately 1. This is because the stoichiometric coefficients for CO2 and O2 are equal.
On the other hand, the RQ value for the oxidation of lipid is <1 and for organic acids is >1. A very high RQ value often indicates
a rapid shift from aerobic to anaerobic respiration (Fonseca et al., 2002a,b). Thus, understanding the dynamics of RQ for horticul-
tural commodities is essential in successful design of controlled atmosphere storage facilities and modified atmosphere packaging
systems.

Factors Affecting Respiration Rate

Respiration is a metabolic activity that provides the energy needed for other plant biochemical reactions. It involves the oxidative
breakdown of complex organic compounds such as carbohydrates, lipids, and organic acids into simple organic molecules with the
release of CO2, water, and energy. This physiological process plays a significant role in the postharvest quality of horticultural
commodities. Table 2 presents a summary of a wide range of factors that influence respiration rate of horticultural commodities
(Saltveit, 1996; Caleb et al., 2013; Mahajan et al., 2014).
4 The Basics of Respiration for Horticultural Products

Table 2 Factors influencing respiration rate

Preharvest Postharvest storage Minimal processing

Cultivar Storage temperature Size and type of cut

Growing season Concentration of O2 Sharpness of cutting blade
Farming system Concentration of CO2 Hot air treatment
Growing region Storage duration Hot water dipping
Produce maturity level Ethylene exposure Antibrowning treatment
Pretreatment Edible coatings
Radiation Irradiation and plasma treatment

Source: Saltveit, M.E. 1996. Physical and physiological changes in minimally processed fruits and vegetables. In: Tomás-
Barberán, F.A. (Ed.), Phytochemistry of Fruit and Vegetables. Oxford University Press, New York, pp. 205–220; Caleb,
O.J., Mahajan, P.V., Al-Said, F.A., Opara, U.L., 2013. Modified atmosphere packaging technology of fresh and fresh-cut
produce and the microbial consequences – a review. Food Bioprocess Technol. 6, 303–329; Mahajan, P.V., Caleb, O.J.,
Zora, S., Watkins, C., Geyer, M., 2014. Postharvest treatments of fresh produce. Phil. Trans. A Math. Phys. Eng. Sci.
372, 1–19.

Maturity or Developmental Stage

Respiration rate of horticultural commodities is significantly influenced by the growth stage wherein they are harvested. Commod-
ities harvested at full maturity such as nuts and tubers have lower respiration rates compared to those with floral or vegetative meri-
stems such as asparagus, cauliflower, and broccoli (Watada et al., 1984; Seefeldt et al., 2012; Watkins and Nock, 2012). This implies
that the rate of deterioration is higher in commodities harvested during active growth phase due to high respiration rates, while
mature fruits have relatively low rates.

Temperature
Storage or postharvest handling temperature is the most important factor affecting postharvest life of fruit and vegetables. It has
a profound impact on the rates of various biochemical and biological reactions (Kader, 2002). According to the Van’t Hoff Rule,
“the velocity of a biological reaction increases two- to threefold for every 10  C rise in temperature.” This phenomenon and the
detrimental effects of high temperature have been extensively reported in postharvest literature for a whole range of fruit and vege-
tables (Kader, 2002; Saltveit, 1996; Iqbal et al., 2009). Therefore, it is crucial to lower temperature around horticultural commod-
ities as quickly as possible after harvest in order to slow down metabolic reactions; except when curing is incorporated into the
postharvest management of certain commodities such as onions and garlic.
Low storage and postharvest handling temperatures help to reduce respiration rate, water loss, and sensitivity to ethylene and to
retard the growth of spoilage microorganism (Caleb et al., 2013). Thus, it is of significant importance to establish an optimum
temperature range for every individual commodity. It is generally assumed that the lower the storage temperature, the lower the
respiration rate, and the longer the postharvest storage life of a given horticultural commodity, as long as this condition is above
the freezing temperature of the product.

Chilling Stress
The benefit of low temperature storage condition is not universal for all commodities. Many products are sensitive to low temper-
atures and develop chilling injuries such as pit skin surface, irregular ripening, internal browning, and increased susceptibility to
decay (Lurie and Crisosto, 2005). Ideally respiration is reduced at lower temperature, but chilling-sensitive commodities (mainly
those originating from subtropical and tropical regions (Table 3)) exhibit abnormal respiration patterns when the storage temper-
ature is below 10–12  C (Saltveit, 1996; Watkins and Nock, 2012). In some cases, respiration may increase dramatically at the chill-
ing temperature conditions or thereafter when the commodity is returned to higher temperatures. This physiological response could
be attributed to the cells’ effort to remove intermediate metabolites accumulated during chilling and repair damaged cellular and
subcellular structures.

Atmospheric Composition
Sufficient O2 concentration is essential to maintain an aerobic respiration process in fresh horticultural commodities. However,
decline in O2 below the critical limit shifts respiration from an aerobic to an anaerobic process, resulting in the production of unde-
sirable fermentative metabolites and development of off-odors (Kader and Saltveit, 2003). The critical O2 limit that enables aerobic
respiration varies significantly among horticultural commodities. Most products are within the range of 2–3% O2 concentration for
biological reactions and respiration (Fonseca et al., 2002a; Kader, 2002; Caleb et al., 2013). While others such as apples and pears
are capable of aerobic metabolic processes at O2 concentrations as low as 0.5–1%. At these low concentrations a significant
 The Basics of Respiration for Horticultural Products 5

Table 3 Classification of horticultural commodities according to sensitivity to

chilling injury

Chilling sensitive Nonchilling sensitive

Cassava Applesa; nectarinesa; plumsa; peachesa

Yams Garlic; carrots; cabbage; onions; radishes
Potatoes Apricots; broccoli; beets; blackberries; brussels sprouts
Pumpkins Grapes; sweet corn; cauliflower; cherries; prunes
Sweet potatoes Artichokes; strawberries; currants; endive
Eggplant Lettuce; parsley; mushrooms; celery;
Cucumbers Asparagus; raspberries; turnips; beans, lima
Cranberries
Bean, snap
a
Some cultivars within this group are chilling sensitive.
Source: Watkins, C.B., Nock, J.F., 2012. Production Guide for Storage of Organic Fruits and
Vegetables. NYS IPM Publication No. 10.

reduction in respiration rate and other metabolic reactions can be achieved, thereby, maintaining fruit quality and extending storage
life considerably.
Conversely, an increase in the CO2 concentration within permissible limits (which is product specific) helps to reduce respira-
tion and retard senescence and fungal growth. There are various suggestions on the possible roles of increase in CO2 on respiration.
These include that CO2 causes a feedback inhibition being a by-product of respiration; the consideration that elevated CO2 might
affect the enzymes and intermediate metabolites involved in the Krebs cycle; and another suggestion is that CO2 retards the produc-
tion of ethylene (Saltveit, 2003; Caleb et al., 2013). However, excessive accumulation of CO2 can alter the metabolic pathway of
fresh horticultural commodities and promote fermentation.

Other Postharvest Handling Stress

Various postharvest handling practices such as mild physical stress (due to mechanical damage and vibration), heat treatments/
shocks, minimal processing and coatings (chemical or edible) can cause considerable perturbation in respiration. For example,
bruises/wounds caused by mechanical damage can induce a broad range of physiological responses. These include increased anti-
oxidant activity, phenolic metabolism, and ethylene production in an attempt to heal the injury, which enhances respiration rate.
Similarly, extensive literature has shown the impact of minimal processing on the respiration rate of horticultural commodities. The
disruption of tissues due to shredding, cutting, or slicing results in accelerated respiration about two- to fourfold higher for pro-
cessed commodities in comparison to intact ones.

Measurement of Respiration Rate

The respiration rate of fresh produce can be measured via three possible methods. These include closed or static systems, where the
product is kept in a closed container; or in flow-through/open systems, where the gas is continuously flushed through the container;
and/or in permeable systems, in which a package with known volume and film permeability is filled with product. Changes in
concentration of O2 and/or CO2 are measured over a period of time in the static system as well as in the open system. For flow-
through systems, changes in O2 and/or CO2 concentration are measured between the inlet and the outlet. In the permeable systems,
steady-state of O2 and/or CO2 concentrations is determined, and a mass balance is used to estimate respiration rate (Fonseca et al.,
2002a,b). The initial gas composition can be either atmospheric air or modified gas composition. The measured changes in gas
composition are used to estimate or calculate respiration rates (Eqns. [1]–[6])
 
i
yf
yO O2  V


2
RO2 ¼ [1]
100  M  tf
ti

 
f i
y CO2
yCO V


2
RCO2 ¼ [2]
100  M  tf
ti

 
in
yout  F
yO 2 O2
RO2 ¼ [3]
100  M
6 The Basics of Respiration for Horticultural Products

 
out
yin
yCO 2 CO2  F
RCO2 ¼ [4]
100  M

PO2  A  
e
RO2 ¼  yO
yO2 [5]
100  L  M 2

PCO2  A  
e
RCO2 ¼  yCO2
yCO [6]
100  L  M 2

All symbols, superscripts and subscripts presented for Eqns. [1]–[6] are as defined by Fonseca et al. (2002a). Generally, respiration
(consumption (RO2)/production (RCO2)) rate can be expressed as mol kg
1 h
1, mL kg
1 h
1 or mg kg
1 h
1, but, the SI unit
(mol kg
1 s
1) is ideal (Banks et al., 1995).
In most of the studies published before 2001, measurement of the O2 and CO2 content was based on using a gas chromatograph.
This approach was time consuming, laborious, and expensive (Fonseca et al., 2002b). However, since year 2002, there has been
a growing interest to employ simple and easy-to-use O2 and CO2 gas analyzers for measurement of the gas content (Caleb et al.,
2013). These analyzers are based on electrochemical (O2) and infrared (CO2) sensors. The analyzers are commercially available
in different formats or brands such as CheckMate (Dansensor, Ringsted, DK), PacCheck (MOCON, Minneapolis, MN), Dualtrak
(Quantek Instruments Inc., Grafton, MA), and GasSpace Advance instrument (Systech Instruments Ltd, Oxfordshire, the United
Kingdom). The gas analyzers, although simple and easy to use, all produce discrete data based on headspace sampling. A gas sample
of 3–15 ml is taken from the headspace for every measurement. With numerous measurements, this can create vacuum pressure in
the static system (Fonseca et al., 2002a). Also the intervals for measurements should be representative of the changing respiration
rates and must be chosen with great care. Since respiration, like many other biological processes, is a dynamic and continuous
process, the O2 and CO2 contents change continuously (Fonseca et al., 2002a).
Noninvasive O2 sensors emerged on the market for continuous O2 measurements (OpTech, Mocon, Minneapolis, MN; PreSens,
PreSens – Precision Sensing GmbH, Regensburg, Germany). These sensors are based on monitoring an oxygen-dependent fluores-
cence signal on a spot stick to the inside of a transparent container; however, these sensors are expensive and only available for O2
content of the container. Løkke et al. (2011) and Seefeldt et al. (2012) applied a wireless sensor network for online, noninvasive O2
measurement in a respirometer containing fresh produce without gas exchange. However, the sensor could measure the O2 content
in the range from 6.3 to 20.9 kPa, and RQ could not be determined as the CO2 content was not measured. PerfoTec (PerfoTec B.V.,
Mijdrecht, Netherlands) also developed the fast respirometer for commercial use by fresh produce industries. It measures the respi-
ration rate of fresh produce within 4 h. The limitation of the device is that it is large and cannot be easily moved into controlled
environment conditions. Thus, measurements in cold storage facilities could be challenging. Various studies have reported the
use of continuous measurements of O2 as well as CO2 contents in flow-through systems filled with fresh produce. However, infor-
mation from the flow-through systems is not representative for the gas conditions inside MAP with fresh produce where the gas
exchange is restricted during storage.
It is well known that storage temperature is the most critical parameter for fluctuations in respiration rates. It has been demon-
strated that the respiration rates increase at elevated temperatures (Caleb et al., 2012; Kader and Saltveit, 2003). In experimental
storage conditions it is assumed that the temperature is strictly controlled during gas sampling. However, Seefeldt et al. (2012) re-
ported that the transfer of a respirometer from a climate chamber to a laboratory bench for discrete gas measurements with a Check-
Mate instrument has significant impact on the accuracy of respiration rates as shown with data obtained via wireless O2 sensor. Such
data are rare in literature because it is difficult to measure minute changes in the respiration rates due to short-time temperature
fluctuations by the use of traditional gas analysis. Even measurements with online fluorescence-based equipment would not be
possible as these systems require constant illumination of the inside spot to give continuous readings of the O2 content.
Recently, Mahajan et al. (2016) developed small and flexible respirometers for the determination of real-time respiration rates.
The minisphere configuration consists of O2, CO2, and RH sensors with high sensitivity and accuracy, which can be directly inside
the glass jar either for closed or flow-through systems. Major advantages of this device included the possibility to determine the gas
content continuously and noninvasively; to calculate the produce respiration rate in real time; to optimize the gas sampling effi-
ciency with little manual handling; and being miniature offers flexibility. The respirometer also offers the capability to analyze
the effect of extrinsic factors such as temperature, gas composition, and storage time on the respiration rate to calculate RQ and
determine the low O2 limit in real time. Collection of such detailed information provides better understanding of the respiration
process and enables a more knowledge-intensive design of packaging and storage systems for fresh horticultural produce. Due to the
complexity of the respiration process, the success of various packaging and storage solutions/designs for horticultural products
greatly depends on the accuracy of the measurement methods. Therefore, more research is needed in this area.

Acknowledgments

This work was supported by the Georg Forster Postdoctoral Research Fellowship (HERMES) program from the Alexander von Humboldt Foundation
(Ref. 3.4–ZAF–1160635-GFHERMES-P).
 The Basics of Respiration for Horticultural Products 7

References

Adu-Bredu, S., Yokota, T., Hagihara, A., 1997. Temperature effect on maintenance and growth respiration coefficients of young, field-grown hinoki cypress (Chamaecyparis obtusa).
Eco. Res. 12, 357–362.
Banks, N.H., Cleland, D.J., Cameron, A.C., Beaudry, R.M., Kader, A.A., 1995. Proposal for a rationalized system of units for postharvest research in gas exchange. HortSci. 30,
1129–1131.
Buchanan, B.B., Gruissem, W., Jones, R.L., 2000. Biochemistry and Molecular Biology of Plants. American Society of Plant Physiologists, Rockville, USA.
Caleb, O.J., Mahajan, P.V., Al-Said, F.A., Opara, U.L., 2013. Modified atmosphere packaging technology of fresh and fresh-cut produce and the microbial consequences – a review.
Food Bioprocess Technol. 6, 303–329.
Caleb, O.J., Mahajan, P.V., Opara, U.L., Witthuhn, C.R., 2012. Modelling the respiration rates of pomegranate fruit and arils. Postharvest Biol. Technol. 64, 49–54.
Chiariello, N.R., Mooney, H.A., Williams, K., 1989. Growth, carbon allocation and cost of plant tissues. In: Pearcy, R.W., Ehleringer, J., Mooney, H.A., Rundel, P.W. (Eds.), Plant
Physiological Ecology: Field Methods and Instruments. Chapman and Hall, New York, pp. 327–365.
Fonseca, S.C., Oliveira, F.A.R., Brecht, J.K., 2002a. Modelling respiration rate of fresh fruits and vegetables for modified atmosphere packages: a review. J. Food Eng. 52, 99–119.
Fonseca, S.C., Oliveira, F.A.R., Frias, J.M., Brecht, J.K., Chau, K.V., 2002b. Modelling respiration rate of shredded Galega kale for development of modified atmosphere packaging.
J. Food Eng. 54, 299–307.
Grange, R.I., Andrew, J., 1995. Respiration and growth of tomato fruit. Plant Cell Env. 18, 925–930.
Hanning, I., Heldt, H.W., 1993. On the function of mitochondrial metabolism during photosynthesis in spinach (Spinacia oleracea L.) leaves. Partitioning between respiration and
export of redox equivalents and precursors for nitrate assimilation products. Plant Physiol. 103, 1147–1154.
Herppich, W.B., Huyskens-Keil, S., 2008. Cell wall biochemistry and biomechanics of harvested white asparagus shoots as affected by temperature. Ann. Appl. Biol. 152, 377–388.
Herppich, W.B., Peckmann, K., 2000. Influence of drought on mitochondrial activity, photosynthesis, nocturnal acid accumulation and water relations in the CAM plants Prenia
sladeniana (ME-type) and Crassula lycopodioides (PEPCK-type). Ann. Bot. 86, 611–620.
Huyskens-Keil, S., Herppich, W.B., 2013. High CO2 effects on postharvest biochemical and textural properties of white asparagus (Asparagus officinalis L.) spears. Postharvest Biol.
Technol. 75, 45–53.
Iersel, M.W., Seymour, L., 2000. Growth respiration, maintenance respiration, and carbon fixation of vinca: a time series analysis. J. Amer. Soc. Hort. Sci. 125, 702–706.
Iqbal, T., Rodrigues, F.A.S., Mahajan, P.V., Kerry, J.P., 2009. Effect of time, temperature, and slicing on respiration rate of mushrooms. J. Food Sci. 74, E298–E303.
Iqbal, T., Rodrigues, F.A.S., Mahajan, P.V., Kerry, J.P., Gil, L., Manso, M.C., Cunha, L.M., 2008. Effect of minimal processing conditions on respiration rate of carrots. J. Food Sci.
73, 396–402.
Kader, A.A., 2002. Postharvest Technology of Horticultural Crops, third ed. University of California, Agriculture and Natural Resources Publication 3311.
Kader, A.A., Saltveit, M.E., 2003. Atmosphere modification. In: Bartz, J.A., Brecht, J.K. (Eds.), Postharvest Physiology and Pathology of Vegetables, second ed. Marcel Dekker Inc.,
New York, pp. 229–246.
Kader, A.A., Zagory, D., Kerbel, E.L., 1989. Modified atmosphere packaging of fruit and vegetables. CRC Crit. Rev. Food Sci. Nutr. 28, 1–30.
Ke, D., Yahia, E., Hess, B., Zhou, L., Kader, A.A., 1995. Regulation of fermentative metabolism in avocado fruit under oxygen and carbon dioxide stress. J. Amer. Soc. Hort. Sci.
120, 481–490.
Kriedemann, P.E., Virgona, J.M., Atkin, O.K., 2016. Growth analysis: a quantitative approach. In: Price, C., Munns, R. (Eds.), Plants in Action: A Resource for Teachers and Students
of Plant Science, second ed. Australian Society of Plant Scientists.
Krömer, S., 1995. Respiration during photosynthesis. Ann. Rev. Plant Physiol. Plant Mol. Biol. 46, 45–70.
Lavigne, M.B., Ryan, M.G., 1997. Growth and maintenance respiration rates of aspen, black spruce and jack pine stems at northern and southern BOREAS sites. Tree Physiol. 17,
543–551.
Løkke, M.M., Seefeldt, H.F., Edwards, G., Green, O., 2011. Novel wireless sensor system for monitoring oxygen, temperature and respiration rate of horticultural crops postharvest.
Sensors 11, 8456–8468.
Lurie, S., Crisosto, C.H., 2005. Chilling injury in peach and nectarine. Postharvest Biol. Technol. 37, 195–208.
Mahajan, P.V., Caleb, O.J., Zora, S., Watkins, C., Geyer, M., 2014. Postharvest treatments of fresh produce. Phil. Trans. A Math. Phys. Eng. Sci. 372, 1–19.
Mahajan, P.V., Lucas, A., Edelenbos, M., 2016. Development of a small and flexible sensor-based respirometer for real-time determination of respiration rate, respiratory quotient
and low O2 limit of fresh produce. Comp. Elect. Agric. 121, 347–353.
Marcelis, L.F.M., Baan Hofman-Eijer, L.R., 1995. Growth and maintenance respiratory costs of cucumber fruits as affected by temperature, and ontogeny and size of the fruits.
Physiol. Plant 93, 484–492.
Nunes-Nesi, A., Araujo, W.L., Fernie, A.R., 2011. Targeting mitochondrial metabolism and machinery as a means to enhance photosynthesis. Plant Physiol. 155, 101–107.
Peckmann, K., von Willert, D.J., Martin, C.E., Herppich, W.B., 2012. Mitochondrial respiration in ME-CAM, PEPCK-CAM, and C3 succulents: comparative operation of the
cytochrome, alternative, and rotenone-resistant pathways. J. Exp. Bot. 63, 2909–2919.
Ryan, M.G., 1991. Effects of climate change on plant respiration. Ecol. Appl. 1, 157–167.
Saltveit, M.E., 1996. Physical and physiological changes in minimally processed fruits and vegetables. In: Tomás-Barberán, F.A. (Ed.), Phytochemistry of Fruit and Vegetables. Oxford
University Press, New York, pp. 205–220.
Saltveit, M.E., 2003. Is it possible to find an optimal controlled atmosphere? Postharvest Biol. Technol. 27, 3–13.
Schaffer, A.A., Pharr, D.M., Madore, M.A., 1996. Cucurbits. In: Zamski, E., Schaffer, A.A. (Eds.), Photo-assimilate Distribution in Plants and Crops. Source-Sink Relationships. CRC
Press, New York, pp. 729–758.
Seefeldt, H.F., Løkke, M.M., Edelenbos, M., 2012. Effect of variety and harvest time on respiration rate of broccoli florets and wild rocket salad using a novel O2 sensor. Postharvest
Biol. Technol. 69, 7–14.
Thornley, J.H.M., 2011. Plant growth and respiration re-visited: maintenance respiration defined – it is an emergent property of, not a separate process within, the system – and
why the respiration: photosynthesis ratio is conservative. Ann. Bot. 1–16.
Tomás-Barberán, F.A., Loaiza-Velarde, J., Bonfanti, A., Saltveit, M.E., 1997. Early wound- and ethylene-induced changes in phenylpropanoid metabolism in harvested lettuce.
J. Am. Soc. Hort. Sci. 122, 399–404.
Watada, A.A., Herner, R.C., Kader, A.A., Romani, R.J., Staby, G.L., 1984. Terminology for the description of developmental stages of horticultural crops. HortSci. 19, 20–21.
Watkins, C.B., Nock, J.F., 2012. Production Guide for Storage of Organic Fruits and Vegetables. NYS IPM Publication No. 10.