Scientia Horticulturae 250 (2019) 113–120

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Scientia Horticulturae
journal homepage: www.elsevier.com/locate/scihorti

Excess water loss induced by simulated transport vibration in postharvest T

kiwifruit
⁎
Xiaopeng Weia, Dandan Xiea, Linchun Maoa, , Changjie Xub, Zisheng Luoa, Ming Xiaa,
Xiaoxiao Zhaoa, Xueyuan Hana, Wenjing Luc
a
College of Biosystems Engineering and Food Science, Zhejiang Key Laboratory of Agro-Food Processing, Key Laboratory of Agro-Products Postharvest Handling of Ministry
of Agriculture and Rural Affairs, Zhejiang University, Hangzhou, 310058, China
b
College of Agriculture and Biotechnology, Zhejiang Provincial Key Laboratory of Horticultural Plant Integrative Biology, Zijingang Campus, Zhejiang University,
Hangzhou, 310058, China
c
Institute of Food Science, Zhejiang Academy of Agricultural Sciences, Hangzhou, 310021, China

A R T I C LE I N FO A B S T R A C T

Keywords: Fruit usually suffers from water loss after transportation and storage, which largely blemishes the quality
Water loss characteristics including appearance, saleable weight and texture. This study was conducted to interpret the
Cell damage potential relations between fruit water loss and transport vibration. Kiwifruit were subjected to vibration using
Kiwifruit an electrodynamic shaker followed by storing at 25 °C and 75% RH (relative humidity) in dark for 12 d. Fruit
Shrivel
weight loss and water content in the epidermis (EP), outer pericarp (OP), inner pericarp (IP) and core tissues
Simulated transport vibration
were determined. Shrinkage of EP cells and water movement were evaluated using laser scanning confocal
microscopy and magnetic resonance imaging, respectively. Cell damage was investigated by observing the ul-
trastructure and electrolyte leakage. The results showed that fruit water loss was accompanied with the spatial
movement of water from interior IP, OP and external EP. Vibration damaged the fruit cells with plasmolysis,
membrane contracting and increased electrolyte leakage, which accelerated movement of water from the tissues
to outward resulting in severe water loss of fruit. However, the electrolyte leakage and water content in the core
were little changed in both control and vibrated fruit. Shrinkage of EP cells coupled with fruit shrivel appeared
by 4 d of storage after vibration. However, the symptom in control fruit was not observed until 8 d of storage. It
could be concluded that the simulated transport vibration caused intracellular damage in fruit tissues, which
accelerated the water loss and shriveling process.

1. Introduction
Water is the unique universal solvent accounting for a major pro-
portion of fresh fruit (De Ita and Flores, 2017), and is involved in
material transport (Rancic et al., 2015), physiological and biochemical
reactions (Bondada and Shutthanandan, 2012; Morinaga and Sykes,
2015), and maintains the appearance (Burdon et al., 2014) and firmness
of fruit (Tsuchida et al., 2008). Fruit quality and postharvest life are
highly determined by water loss which causes fruit shrivel and
browning (Lin et al., 2010; Burdon et al., 2014), fresh weight loss and
deterioration (Li et al., 2011). Water loss from harvested fruit con-
tinuously occurs through transpiration (Leide et al., 2007) that strictly
depends on the epidermis features and environmental conditions
(Morandi et al., 2007). The fruit epidermis acts as a critical barrier to
resist the movement of water both as a liquid and vapors from fruit
tissue to the outer surface, and allows the fruit to maintain a high water
content (Veraverbeke et al., 2003; DíazPérez et al., 2007). Mechanical
damage including surface and internal damage that occur in different
fruit harvest and postharvest stages can greatly accelerate the rate of
water loss from fruit (Li and Thomas, 2014). Surface damage allows
greater flux of water as a liquid and vapors through the damaged epi-
dermis (Workneh et al., 2012). However, the relations between in-
tracellular damage and fruit water loss have not been reported.
Transportation is necessary to distribute fruit after harvest, but vi-
bration generated by vehicles during road transportation is one of the
major causes for critical surface and internal damage to fruit
(Zeebroeck et al., 2007; Fadiji et al., 2016). The physical and chemical
property changes of fruit caused by vibration have been tested by Li
et al. (2000) and Zhou et al. (2007). Much attention has been paid to
assess mechanical damage to different species of fruit during transport,

⁎
Corresponding author.
E-mail address: linchun@zju.edu.cnl (L. Mao).

https://doi.org/10.1016/j.scienta.2019.02.009
Received 18 October 2018; Received in revised form 14 January 2019; Accepted 4 February 2019
Available online 20 February 2019
0304-4238/ © 2019 Elsevier B.V. All rights reserved.
X. Wei, et al. Scientia Horticulturae 250 (2019) 113–120

including apples (Vursavus and Ozguven, 2004), pears (Berardinelli

et al., 2005), tomatoes (Wu and Wang, 2014) and kiwifruit
(Tabatabaekoloor et al., 2013). Unlike surface damage, the internal
damage is difficult to be detected and assessed quantitatively mainly
due to the invisible wound, but could cause rapid deterioration to
subjected fruit in a short period of time (Li et al., 2013). This damage is
a critical problem affecting the fruit firmness and ripening (Li et al.,
2000), sugar and acid contents (Montero et al., 2009; Alfatni et al.,
2013), internal browning (Gonzalez et al., 2001) and core breakdown
(Milczarek et al., 2009).
Kiwifruit is a popular and commercial fruit partly due to its deli-
cious flavor and succulence, but the fruit commonly suffers from the
surface damage (Lallu et al., 1999; Tabatabaekoloor et al., 2013) and
internal damage during the transport (Li et al., 2000). In addition, ki-
wifruit had a propensity to shrivel when suffering from water loss
(Burdon and Clark, 2001). Our previous study noticed that shrivel of Fig. 2. Transverse image of kiwifruit with tissue annotation.
kiwifruit became particularly obvious after simulated transport vibra-
tion, although there was no visible damage on the fruit surface. In this
study, the contributions of the internal damage and real-time water loss
on vibration-mediated shrivel in kiwifruit were investigated by de-
terminations of water movement and cell ultrastructure including cell
wall and membrane integrity.

2. Materials and methods

2.1. Fruit and vibration

Kiwifruit (Actinidia deliciosa cv. Xuxiang) with similar size and

shape, free from mechanical damage or infections, were harvested at
commercial maturity (firmness 115.5 ± 5.0 N, soluble solids content
12.0 ± 0.3%) in Hangzhou, Zhejiang Province, China. The electro-
dynamic vibration system (D–200–3, Suzhou Sushi Testing Instrument
Co., China) consisting of a RC–2000 digital vibration controller, a SA–3
power amplifier and a D–200–3 vibration shaker with a platform
(500 × 500 × 30 mm), was used to simulate transport vibration. An
accelerometer fixed at the bottom of the platform, and the computer Fig. 3. Weight loss of kiwifruit over 12 d of storage. 0 d, immediate determi-
installed with LMS Test Lab software were used for data acquisition and nation after vibration. Data are the mean ± SD of triplicate measurement and
analysis (Fig. 1). Vibration sweep tests were done to determine the compared by one-way ANOVA. Different letters above the points indicate sig-
frequencies with the response on the platform, by sweeping over the nificantly different values (P < 0.05) by LSD test.

frequency ranges normally encountered road transportation. The 85

kiwifruit were fixed in foam-rubber cushion, and the fruit were placed
about 2–3 cm apart to avoid friction for each other. Kiwifruit on the
platform were subjected to vibration at 20 Hz at an amplitude of 0.9 g at
specified duration of 5 h (Fadiji et al., 2016), followed by storing in an
incubator (HWS, Ningbo Southeast Instrument Co., China) in dark at
25 °C and 75% RH for 12 d. Three replicates were conducted for the
vibration test and fruit without vibration were set as control. The ki-
wifruit after vibration had no visible damage on the surface.

2.2. Laser scanning confocal microscopy (LSCM)

Cellular morphology of the epidermis (EP) was observed using

Fig. 1. Schematic layout of vibration system. Test parameters were given to the
computer installed with LMS Test Lab software. RC-200 vibration controller
directed the SA-3 power amplifier to amplify the signal from the computer and
to drive the D-200-3 shaker for simulating transport vibration. And then the
vibration parameters were detected and monitored by the accelerometer that
was controlled through RC-200 vibration controller.
LSCM according to Wei et al. (2018). Cubes (2 × 2 × 2 mm) of EP
tissue were excised with a razor blade and embedded in freezing
medium (Leica Biosystems, USA) and cooled to −20 °C. Frozen cubes
were cut into sections of 10 μm in thickness using a microtome (Leica
CM 1520, Leica Microsystems Co., Germany). The sections were ob-
served under a Leica TCS-SP8 laser scanning confocal microscope (Leica
Microsystems Co., Germany) with a HCX PL APO CS10 ×/0.40 DRY
objective using a fluorescence excitation filter at 470–495 nm and
emission filter at 510–550 nm. Fluorescence image processing was ac-
complished using LAS-AF 1.6.3 software (Leica Microsystems Co.,
Germany).

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X. Wei, et al. Scientia Horticulturae 250 (2019) 113–120

Fig. 4. Fruit appearance and autofluorescence

micrographs of epidermis cells in control and
vibrated kiwifruit over 12 d of storage. AeD,
kiwifruit without vibration; EeH, kiwifruit
subjected to transport vibration; a–d, auto-
fluorescence micrographs of epidermis cells in
kiwifruit without vibration; e–h, auto-
fluorescence micrographs of epidermis cells in
vibrated kiwifruit.

Fig. 5. Transmission electron micrographs of epidermis cells in control and vibrated kiwifruit. CW, cell wall; lv, lytic vacuole; M, cell membrane. Bar scale 1 μm.

115
X. Wei, et al.
Fig. 6. MR proton density images of the intact kiwifruit (A–H) and the corresponding water contents in the EP, OP, IP and core tissues of control (J) and vibrated
fruits (K) over 12 d of storage. EP, epidermis; OP, outer pericarp; IP, inner pericarp.
2.3. Transmission electron microscopy (TEM)
TEM observation was performed according to Bar-Dror et al. (2011).
At 0 and 12 d after vibration, control and vibrated fruit were sampled.
Fresh cut cubes (2 × 2 × 2 mm) from the EP tissues were excised with a
razor blade, paralleling to the fruit longitudinal axis. Cubes were fixed
in 3% (W/V) glutaraldehyde in 0.1 mol L−1 phosphate buffer (pH 7.2)
for 2 h, rinsed with the phosphate buffer, and subsequently post-fixed in
1% (W/V) osmium tetroxide in 0.1 mol L−1 phosphate buffer (pH 7.2)
for 2 h. All steps were performed at 4 °C. The samples were dehydrated
in an ethanol series and acetone once, and then infiltrated and em-
bedded with epoxy. Sections (60 nm in thickness) were obtained by
ultramicrotome (EM UC7, Leica Microsystems Co., Germany), and
stained with uranyl acetate and lead citrate. The stained sections were
observed under a JEOL JEM 1010 transmission electron microscope
(H–7650, Hitachi High-technologies Co., Japan).
2.4. Magnetic resonance imaging (MRI)
Proton magnetic resonance imaging (1H-MRI) of kiwifruit were
obtained in a 3 T MR scanner (MAGNETOM Prisma, siemens
Healthcare, Erlangen, Germany) (Mazhar et al., 2015). Kiwifruit were
placed into a 32–channel head coil and positioned to make the fruit axis
perpendicular to the main magnetic field. T2 weighted turbo-spin echo
(TSE) images were acquired with the following parameters: recycle
time (TR) = 6000 ms, echo time (TE) = 100 ms, slice thickness = 2.0
mm, field of view = 240 × 240 mm, matrix size = 320 × 320 pixels, in
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Scientia Horticulturae 250 (2019) 113–120
plane resolution = 0.8 × 0.8 mm. Obtained 1H-MRI images were ana-
lyzed with OsiriX (Pixmeo SARL, Bernex, Switzerland) DICOM viewer
software.
2.5. Weight loss and water content determinations
The 25 kiwifruit for each treatment were weighed at indicated time
points to determine weight loss, and three replicates were performed.
Weight loss was calculated as the percentage of the pre-vibration
weight that had been lost. Water content of excised pieces of EP, OP, IP
and core tissues (Fig. 2) was determined by drying at 70 °C for 24 h
(Burdon et al., 2014).
2.6. Electrolyte leakage determination
Electrolyte leakage was measured using a conductivity meter
(Mettler Toledo, Switzerland) according to Zhou et al. (2007). Disks of
EP were obtained with a cork borer, diameter 11 mm and weighing 3 g,
and the disks (11 mm in diameter, 4 mm in thickness) were also ob-
tained from OP, IP and core tissues. Ten disks were placed in 40 mL of
distilled deionized water. The first electrical conductivity was assayed
after 10 min vacuum filtration (0.05 MPa). The disks subsequently
boiled for 15 min and the electrical conductivity was re-assayed after
the disks reached room temperature. The percent value of relative
electrical conductivity was calculated as electrolyte leakage by the first
result (electrical conductivity after vacuum filtration) divided by the
second result (electrical conductivity after boiling).
X. Wei, et al.
Fig. 7. Water content of the EP (A), OP (B), IP (C) and core (D) tissues over 12 d of storage. Data are the mean ± SD of triplicate measurement and compared by one-
way ANOVA. Different letters above the points indicate significantly different values (P < 0.05) by LSD test.
2.7. Statistical analysis
Experiments were performed in a completely randomized design
and repeated in triplicate at each time point. Data were analyzed by a
one-way analysis of variance (ANOVA), using SPSS Statistics Version 24
(USA), and presented as mean plus/minus standard deviation. The least
squares difference (LSD) test was employed to determine the statistical
significance of the differences between the means (P < 0.05).
3. Results
3.1. Fruit weight loss and shrivel incidence
Weight loss of control and vibrated fruit gradually increased with
the storage time, however, vibrated fruit suffered from more significant
weight loss (Fig. 3). By 4 d of storage weight loss of vibrated fruit
reached 2.3% and fruit shrivel appeared (Fig. 4F). The appearance was
more severe with the significant increase of weight loss that was 1.7-
fold of the control by 8 d of storage and 1.5-fold of the control by 12 d
of storage. In contrast, shrivel of control fruit appeared with 2.4%
weight loss by 8 d of storage (Fig. 4C).
3.2. Morphology and ultrastructure of epidermis cell
Cellular morphology of EP was observed using laser scanning con-
focal microscopy (Fig. 4a–h). Just after the vibration (0 d), the shape
and size of EP cells had no obvious change relative to that of control. EP
cells in vibrated fruit distorted and in irregular shape with the size
contracting by 4 d of storage. After 8 d of storage, the cells became
severely shrank and tightly stacked up. In contrast, the shrinkage of EP
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Scientia Horticulturae 250 (2019) 113–120
cells in control fruit was not observed until 8 d of storage.
Cell ultrastructure in EP was observed by transmission electron
microscopy (Fig. 5). Just after vibration (0 d), cell membrane separated
from cell wall and contracted inwards, and lytic vacuoles were distorted
with uneven membranes (Fig. 5C). However, the EP cells in control fruit
had an intact membrane structure and different size of lytic vacuoles
(Fig. 5A). Further significant changes of EP cells in vibrated fruit oc-
curred by 12 d of storage. The staining intensity of cell wall material
reduced predominantly with the size contracting, and cell membrane
and lytic vacuoles were degraded (Fig. 5D). In control fruit, the cell
membrane and some lytic vacuoles were still apparent, although the
membrane separated from cell wall (Fig. 5B).
3.3. Magnetic resonance imaging (MRI) and water content
Transverse images of kiwifruit were obtained using 3 T magnetic
resonance imaging (Fig. 6A–H). A high signal intensity (bright) corre-
sponds to high water content, and the low intensity (dark) corresponds
to the low water content (Fennell and Line, 2001; Mazhar et al., 2015).
The process of water loss in fruit tissues was shown in Fig. 6. In both
control and vibrated fruit, water loss initially occurred in EP and OP
followed by IP, and vibration accelerated water loss from the tissues.
However, the water content in core area had no obvious change during
the entire period of storage.
Decrease in water content mainly occurred in EP, OP and IP, where
the initial water contents were higher than that of core (Fig. 7). In vi-
brated fruit, water content in EP and OP deceased rapidly during the
entire period of storage, and their water contents were 1.0-fold of the
control by 12 d of storage. However, changes in IP water content were
not significant within the early 4 d of storage in both control and
X. Wei, et al.
Fig. 8. Electrolyte leakage of the EP (A), OP (B), IP (C) and core (D) tissues over 12 d of storage. Data are the mean ± SD of triplicate measurement and compared by
one-way ANOVA. Different letters above the points indicate significantly different values (P < 0.05) by LSD test.
vibrated fruit, but followed by a more rapid decrease than EP and OP
during the late period. In contrast, there were little changes in water
content of core tissue, which was also little influenced by the vibration.
3.4. Electrolyte leakage
Electrolyte leakage is an important indicator of cell damage (Deng
et al., 2005; Khare et al., 2010). In both control and vibrated fruit,
electrolyte leakage of EP, OP and IP tissues increased with the storage
time (Fig. 8). Vibration dramatically increased electrolyte leakage of
EP, which was 1.3-fold of the control by 12 d of storage. Similarly,
vibration significantly increased the electrolyte leakage of OP and IP
which were 1.1-fold and 1.2-fold higher than that of control by 12 d of
storage. In contrast, electrolyte leakage of core was almost unchanged
over 12 d of storage, which retained 15.6–17.7% in both control and
vibrated fruit.
4. Discussion
Vibration is the principal contributor for fruit damage during
transportation. Mechanical damage can lead to excess water loss of fruit
(Zhou et al., 2007; Pan et al., 2015), further affecting the fruit ap-
pearance and shelf life (Burdon et al., 2015; Bouzo and Gariglio, 2016).
In this study, we firstly found that simulated transport vibration ac-
celerated kiwifruit water loss and shriveling due to intracellular
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Scientia Horticulturae 250 (2019) 113–120
damage.
Fruit shriveling is one of major physical changes and influencing
appearance, resulting from the shrinkage of flesh tissues, especially the
EP cells (Burdon et al., 2014). The size and shape of the cells depend on
cell turgor pressure and cell wall structure (Lewicki and Pawlak, 2005;
Joardder et al., 2015). Besides cell wall structure, turgor pressure that is
exerted by intracellular water on the cell membrane and cell wall, gives
elasticity, rigidity and strength to the cells (Lewicki and Pawlak, 2003;
Oey et al., 2007). Vibration damaged the EP cells with plasmolysis
(Fig. 5C) and increase of electrolyte leakage (Fig. 8A), which ac-
celerated the water loss from EP tissue. As the water loss and damaging
cell structure of EP tissue, the EP cells lost turgor pressure and the
mechanical properties resulting in the cells shrinkage and shrivel ap-
pearance of fruit (Fig. 4). By 12 d of storage, predominant water loss
and damages of EP cells (Fig. 5D) caused the severe shriveling in vi-
brated fruit (Fig. 4H).
Water loss of harvested fruit is driven by the water vapor pressure
between the ambient air and epidermis (DíazPérez et al., 2007), and
thereby inducing water movement in the fruit tissues (Veraverbeke
et al., 2003). In the present study, water loss in fruit tissues initially
occurred in external EP and OP followed by IP (Fig. 6), which was
consistent with the result of water content (Fig. 7). This result suggested
the spatial movement of water from interior IP, OP and outward EP
during water loss of kiwifruit. Water movement within fruit tissue is
caused by water transport in microstructures, particularly cell wall and
X. Wei, et al.
cell membrane (Nguyen et al., 2006; Van, 2007), which are crucial
microstructures affecting quality and physiology of fruit and also the
first targets of plant abiotic stresses (Roy et al., 2006). A previous study
indicated that mechanical vibration induced the electrolyte leakage and
increased the activity of lipoxygenase, which lead to over-ripe changes
in kiwifruit (Li et al., 2000). Intact cell wall and membrane are semi-
permeable and the main hindrance for water movement between cells
(Fanta et al., 2013, 2014). In the present study, vibration damaged the
cell wall and membrane (Fig. 8A–C) of fruit tissues (IP, OP, and EP) that
accelerated the unrestricted movement of water from the tissues to
outer surface. The EP plays a pivotal role in resistance to the movement
of water from interior tissue to the outer surface (Veraverbeke et al.,
2003; Celano et al., 2009; Maguire et al., 1999). Vibration damaged the
cell structure of EP with plasmolysis, membrane contracting (Fig. 5C)
and increased electrolyte leakage (Fig. 8A), which caused the loss of
natural water barrier and assisted water loss of fruit.
In addition, outward movement of water within flesh tissues was
dependent upon the spatial distance and primary water content. Rapid
water loss initially occurred in the external tissues (EP and OP), but was
delayed in the interior tissue (IP) by 4 d of storage. However, water
content in the core was almost unchanged over 12 d of storage, which
could be accounted for low water content (Fig. 7D), stable cell structure
(Fig. 8D) and remote spatial distance from the fruit surface (Fig. 2).
In conclusion, simulated transport vibration stimulated the water
loss from kiwifruit although there was no visible damage on the fruit
surface. Kiwifruit water loss was the result of spatial movement of
water from interior IP and OP to external EP. Intracellular damage
caused by vibration was the main contributor to the excess water loss,
which accelerated fruit shriveling. This study interpreted the vibration-
mediated water loss in postharvest fruit and reminded the importance
of fruit packaging and transportation to ensure the good quality to the
final consumers.
Funding
This work was financially supported by the National Key Research
and Development Program of China (2018YFD0401303) and the
National Natural Science Foundation of China (31772365).
Author contributions
LCM, DDX and XPW conceived and designed the experiments. XPW,
DDX, MX and XXZ performed the experiments. XPW and DDX analyzed
the data and wrote the manuscript. CJX, ZSL, XYH and WJL also con-
tributed to the data interpretation and writing. All authors read and
approved the final manuscript.
Conflict of interest
The authors have declared no conflict of interest.
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