Available online at www.sciencedirect.
com
Biotechnological interventions in reducing losses of
tropical fruits and vegetables
Mohammad Irfan 1, Pankaj Kumar 2,
Mohammad Feza Ahmad 3 and Mohammed Wasim Siddiqui 3,*
Tropical fruits and vegetables are predominantly cultivated in
warm climate zones, resulting in cultivar diversity in terms of
structure, features, and physiology. These constitute a variety
of bioactive ingredients such as vitamins, minerals, phenolic
acids, anthocyanins, flavonoids, fatty acids, fiber, and their
distinctive appearances attract customers across the world.
The global production of fruit and vegetables has been attained
a tremendous increase for the last few decades. However, huge
losses at pre- and postharvest levels are major constraints in
their judicious use. Traditional breeding strategies were used to
minimize these losses, but their functionality is limited due to
their time and labor intensiveness. Recent biotechnological,
computational, and multiomics approaches not only address
the losses concern but also aid in boosting crop productivity
and nutritional values. This article emphasizes molecular tools
that have been used to reduce losses of tropical fruits and
vegetables at pre- and postharvest levels.
Addresses
1
Plant Biology Section, School of Integrative Plant Science, Cornell
University, Ithaca, NY, USA
2 50% of total produce, resulting in a major economic loss
Department of Biotechnology, Dr. Y.S. Parmar University of
Horticulture and Forestry, Solan, Himachal Pradesh, India
3
Department of Food Science and Postharvest Technology, Bihar
Agricultural University, Sabour 813210, Bihar, India
Corresponding author:
Siddiqui, Mohammed Wasim (wasim_serene@yahoo.com)
*
ORCID-0000-0002-8885-3619
Current Opinion in Biotechnology 2023, 79:102850
This review comes from a themed issue on Food Biotechnology
Edited by Max I. Teplitski and Jorge M. Fonseca
For complete overview of the section, please refer to the article
collection, “Food Biotechnology (2023)”
Available online 5 December 2023
https://doi.org/10.1016/j.copbio.2022.102850
0958–1669/© 2022 Elsevier Ltd. All rights reserved.
Introduction
Fruits and vegetables, predominantly grown in tropics,
are considered as an essential component of daily human
diet all over the world. In the last few decades, the de-
mand for tropical fruits and vegetables has been in-
creased significantly, particularly in the United States
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and European Union. Among several tropical fruits and
vegetables, banana, mango, citrus, papaya, pineapple,
sweet potato, and cassava are the most prominent in
international commerce (Figure 1). Tropical fruits are
super rich in health-benefiting compounds such as an-
thocyanins, flavonoids, carotenoids, vitamins, and fibers,
serving a significant source for human nutritional re-
quirement [1].
The production of tropical fruits and vegetables is sig-
nificantly affected by various factors, including abiotic
and biotic stresses (Figure 1), for instance, in banana,
drought stress causes up to 65% yield loss [2]. Moreover,
various diseases during plant growth and development
limit fruit productivity [3]. Most of the tropical fruits are
perishable in nature, respire and transpire even after
harvest, and are sensitive to chilling. This results in
significant postharvest losses, particularly in low- and
middle-income countries, which are main contributors of
tropical produce [4]. In some cases, this loss goes up to
of such countries [5]. Because of rising global demand of
tropical fruits, their nutritional quality, and flavor, the
need for developing technologies to maintain and en-
hance the production of tropical crops and their pre- and
postharvest quality attributes is essential. In this review,
we primarily focus on biotechnological approaches to
reduce losses of tropical fruits and vegetables at pre- and
postharvest levels.
Biotechnological approaches
In contrast to temperate fruits, which includes apple,
tomato, grapes, and so on, the genetic improvement of
tropical fruits is limited possibly because of genome
complexity, long juvenile phase, and lack of infra-
structure and resources in growing countries [6]. Al-
though conventional breeding methods have been
employed to reduce the pre- and postharvest losses, but
their use is restricted due to their time-consuming and
labor-intensive procedures [7]. To augment conven-
tional breeding of tropical crops, a variety of bio-
technological approaches, such as genetic transformation
and transgenics, antisense technology, RNA inter-
ference, and genome editing, have been employed,
which efficiently address these challenges (Figure 2).
Over the last few years, several studies have been pub-
lished describing particular use of transgenics, genomics,
Current Opinion in Biotechnology 2023, 79:102850
2 Food Biotechnology
Figure 1
Tropical fruits and vegetables with major consumption and commercial importance, and key factors responsible for their losses at pre- and
postharvest levels.
and genome editing in controlling pre- and postharvest
losses of tropical fruits and vegetables. Here, we briefly
review these successful approaches in the following
sections.
Transgenic (heterologous and
overexpression) approach
The modern biotechnological tools allow the manipula-
tion of genes from various sources and their insertion into
plants to impart desirable traits for crop improvement.
This approach has been used for developing several tro-
pical crops to prevent yield losses. Banana (Musa spp.) is
one of the most significant economic commodities in
tropical regions, having commercial production in over
130 countries. The banana industry is severely impacted
by sigatoka leaf spot disease caused by a fungus
Mycosphaerella fijiensisis. To address this, Vishnevetsky
et al. expressed Trichoderma harzianum endochitinase
(ThEn-42) gene together with stilbene synthase gene
from grape and a superoxide dismutase gene Cu,Zn-SOD
from tomato to develop transgenic banana plants [8].
These transgenic banana plants showed enhanced re-
sistance against sigatoka leaf spot disease in field condi-
tions without compromising yield and fruit quality.
Interestingly, the transgenic banana was also resistant to
gray mold Botrytis cinerea, a major fruit pathogen [8]. An-
other fungal disease Fusarium wilt or Panama disease
caused by Fusarium oxysporum affects many tropical fruit
crops worldwide. To improve the resistance against Fu-
sarium oxysporum f. sp. cubense tropical race 4 (TR4) in
banana, a resistance gene analog 2 from a TR4-resistant
diploid banana and an antiapoptosis gene, Ced9 from a
nematode Caenorhabditis elegans, were expressed in
Current Opinion in Biotechnology 2023, 79:102850
Current Opinion in Biotechnology
cavendish banana [9]. These two transgenic banana vari-
eties provided significant resistance improvement against
Fusarium in the field trials [9].
Papaya (Carica papaya L.) is a major subsistence food
and is an important part of general diet in the Southeast
Asia region. The potyvirus papaya ringspot virus (PRV)
poses a serious threat to papaya and reduces its economic
worth. The traditional breeding methods only control
the PRV to a certain extent. To improve the resistance
against PRV, Fitch et al. (1992) transformed immature
zygotic embryos with a PRV coat protein (cp) gene using
microprojectiles [10]. The developed papaya transgenic
lines showed significant resistance to PRV with one
transgenic line displaying total resistance. Further, the
genome of this transgenic papaya was sequenced, which
provided novel insights into its genetics and evolution
[11]. Later, this transgenic papaya was released in Hawai
and commercialized in the United States as ‘SunUp’ and
Rainbow, a hybrid of ‘SunUp’ and a nonengineered
cultivar ‘Kapoho’ [12]. Other than these examples em-
ploying transgenic technology, several other transgenic
tropical crops were also developed as enlisted in Table 1.
Antisense and RNAi technology
Antisense and RNAi technology are gene silencing ap-
proaches (Figure 2a and b) that have been successfully
applied for developing tropical crops with improved fruit
traits (Table 1). One of the recent examples is devel-
opment of banana with delayed ripening and extended
shelf life through targeting two MADS box genes, Ma-
MADS1 and MaMADS2, which are necessary for banana
ripening [13]. Recently, a genetically modified
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Figure 2
Schematic representation of mechanism of antisense (a), RNAi (b), and CRISPR–Cas9 (c) technologies. (d) Critical steps for developing genetic and
genome-engineered fruit crops using above technologies.
pineapple (Ananas comosus) with modified fruit color and
delayed ripening/senescence has been commercialized
by Del Monte as Pinkglow™ in the United States after
approval from FDA [14]. In the Pinkglow™ pineapple,
pink flesh accumulates lycopene, which is developed by
expressing tangerine (Citrus reticulata) phytoene syn-
thase gene, critical for fruit carotenoid biosynthesis, and
suppressing of endogenous lycopene-β and ε-cyclase
genes (β-LYC and ε-LYC) by RNAi. Moreover, a mer-
istem-specific 1-Aminocyclopropane-1-carboxylic acid
(ACC) synthase was also silenced by RNAi in the
Pinkglow™ pineapple to inhibit the flowering and
ethylene production eventually resulting in delayed ri-
pening [14]. Sweet potato (Ipomoea batatas (L.) Lam.) is
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Postharvest Loss reduction in fruit & vegetables Irfan et al. 3
Current Opinion in Biotechnology
an important staple food crop in Sub-Saharan Africa,
however, its production is severely affected by an insect
pest African sweet potato weevil (Cylas brunneus). As a
crop protection strategy, Christiaens et al. delivered
gene-specific dsRNA against 24 target genes of Cylas
brunneus, which resulted in effective silencing and a
rapid and high mortality of the insect [15].
Despite the successful application of transgenic, antisense,
and RNAi technology in improving tropical crop traits,
concerns have been expressed regarding high off-target
effects resulting in modified phenotypes, unintended, and
unpredictable pleiotropic effects of genetically engineered
products on human health and environment [16].
Current Opinion in Biotechnology 2023, 79:102850
 4

Table 1
Examples of genetically modified tropical fruits and vegetables with improved traits for reducing pre- and postharvest loss using biotechnological tools.

Fruit and Target gene (s) Approach Delivery method Trait (s) improved Pre- or postharvest Reference
Food Biotechnology

vegetable
Banana Hrap and Pflp10 Transgenic Agrobacterium-mediated Resistance to BXW Preharvest [45]
gene transfer
Banana MaMADS1, Antisense or RNAi Agrobacterium-mediated Delayed ripening and enhanced shelf life Postharvest [13]
MaMADS2 gene transfer
Banana BSOLV CRISPR–Cas9 Agrobacterium-mediated Resistance to banana streak virus (eBSV) Preharvest [19]
gene transfer
Banana MaACO1 CRISPR–Cas9 Agrobacterium-mediated Delayed ripening and enhanced shelf life Postharvest [21]
gene transfer

Current Opinion in Biotechnology 2023, 79:102850

Cassava IF4E, nCBP1, CRISPR-P Agrobacterium-mediated Resistance to CBSD Preharvest [23]
nCBP-2 gene transfer
Cassava AC2, AC3 CRISPR–Cas9 Agrobacterium-mediated Resistance to geminivirus Preharvest [24]
gene transfer
Papaya ACO Transgenic Biolistic gun Delayed ripening and enhanced shelf life Postharvest [52]
Papaya ACO RNAi Agrobacterium-mediated Delayed ripening and enhanced shelf life Postharvest [49]
gene transfer
Papaya CP RNAi Particle-bombardment Broader-spectrum resistance against Hainan papaya ringspot Preharvest [50]
transformation virus (PRSV)
Citrus Methionine Overexpression Agrobacterium-mediated Resistance to Xanthomonas citri Pre- and [51]
gene transfer postharvest
Citrus CsLOB1 CRISPR–Cas9 Agrobacterium-mediated Resistance against citrus canker Pre- and [26]
gene transfer postharvest
Duncan CsLOB1 CRISPR–Cas9 Agrobacterium-mediated Resistance against Xanthomonas citri subsp. citri Pre- and [27]
grapefruit gene transfer postharvest
Sweet orange MTSE1 Downregulation Agrobacterium-mediated Increased resistance to Xanthomonas citri subsp. citri and Postharvest [47]
gene transfer Penicillium digitatum
Sweet orange pthA-nls Overexpression Agrobacterium-mediated Increased resistance to Xanthomonas axonopodis pv. citri Pre- and [48]
gene transfer postharvest
Squash CP Transgenic Agrobacterium-mediated Resistance against cucumber mosaic virus (CMV), watermelon Preharvest [53]
gene transfer mosaic virus 2 (WMV 2), and zucchini yellow mosaic virus (ZZMV)
Cucumber eIF4E CRISPR–Cas9 Agrobacterium-mediated Resistance to the potyviruses Zucchini yellow mosaic virus and Preharvest [22]
gene transfer papaya ringspot mosaic virus-W
Sweet potato IbCBF3 Overexpression Agrobacterium-mediated Improved cold and drought stress tolerance Preharvest [46]
gene transfer

Hrap (hypersensitivity response-assisting protein; Pflp10 (plant ferredoxin-like protein); MaMADS1 and MaMADS2 (Musa acuminata MADS box gene); AC2 (codes for transcription activation
protein TrAP involved in gene activation, virus pathogenicity, and suppression of gene silencing); AC3 (codes for the REn protein involved in replication enhancement); RP (replicase gene); ACO
(ACO-1-aminocyclopropane-1-carboxylic acid oxidase); Methionine (modified plant thionin); CsLOB1 (citrus sinensis LATERAL ORGAN BOUNDARIES 1); MTSE1(limonene synthase that cat-
alyzes the formation of limonene); pthA-nls (pathogenesis gene encoding 3 nuclear localization signals that are critical for PthA’s function and localization to the host cell nucleus); CaERF28
(capsicum ethylene-responsive factor); IbCBF3 (Ipomoea batatas C-repeat-binding factor 3).

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Genome editing
Over the last decade, genome editing approaches, par-
ticularly CRISPR–Cas9 (Figure 2c), have emerged as a
novel tool for improving the sustainable agriculture
[5,17,18]. Several tropical fruits crops with enhanced
disease resistance, fruit quality, productivity, and ex-
tended shelf life have been developed using
CRISPR–Cas9 technology (Table 1). For example, in
banana, banana streak virus strain Obino l'Ewai
(BSOLV) was targeted for resistance to banana streak
virus using CRISPR/Cas9-based targeted mutagenesis
[19]. Similarly, the resistance of banana against banana
Xanthomonas wilt (BXW) caused by Xanthomonas cam-
pestris pv. musacearum was improved by using
CRISPR–Cas9-based targeted mutagenesis [20]. Re-
cently, Hu et al. used CRISPR–Cas9 with multiple
editing strategies to create Musa acuminata-1-aminocy-
clopropane-1-carboxylic acid oxidase-1 (MaACO1)-dis-
rupted banana plants, resulting in longer fruit shelf life
via reduced ethylene production [21]. In cucumber, the
broad virus resistance against Potyviridae family viruses
(Cucumber vein yellowing virus, Zucchini yellow mosaic virus,
and Papaya ringspot mosaic virus-W) was developed by
disrupting the function of eukaryotic translation initia-
tion factor 4E (eIF4E) gene through CRISPR–Cas9
system [22]. In Cassava, brown streak disease (CBSD)
resistance was achieved by using CRISPR–Cas9 tech-
nology by targeting genes translation initiation factor
4E (IF4E), novel cap-binding protein-1 (nCBP1), and
novel cap-binding protein-2 (nCBP-2) [23]. Similarly,
viral genes AC2 and AC3 were targeted to impart gemi-
nivirus resistance [24].
In Citrus, the bacteria Xanthomonas citri subsp.
citri causes citrus canker with severe symptoms in the
stem, leaf, and fruit at both pre- and postharvest levels,
resulting in huge economic losses. To address this,
LATERAL ORGAN BOUNDARIES 1 (CsLOB1), a
susceptibility gene and key transcription factor re-
sponsible for disease expression, was knocked out
through CRISPR–Cas9 approach [25]. CsLOB1 contains
an effector-binding element (EBEPthA4) in its promoter
region, which is recognized by PthA4, the main effector
of Xanthomonas citri, to activate the expression of
CsLOB1 gene resulting in citrus canker development. To
modify the EBEPthA4 of CsLOB1 promoter, different
targeted mutations were created using CRISPR–Cas9
and several Citrus mutants showed disrupted CsLOB1
induction in response to Xanthomonas citri infection
eventually leading enhanced resistance to citrus canker
[26]. This suggests that CRISPR/Cas9-mediated
CsLOB1 promoter editing is an effective method for
developing canker-resistant citrus cultivars. Similarly,
CRISPR–Cas9-based mutants (DLOB9 and DLOB10)
with improved resistance to Xanthomonas citri were pro-
duced in duncan grapefruit (Citrus paradisi) by targeting
CsLOB1 [27].
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Postharvest Loss reduction in fruit & vegetables Irfan et al. 5
Bioinformatics and omics interventions in
tropical fruit biotechnology
In recent years, various bioinformatics and omics ap-
proaches such as genomics, transcriptomics, proteomics,
metabolomics, ionomics, and phenomics have revitalized
the plant biotechnology and played a significant role in
various crop improvement programs, including traits of
tropical fruits and vegetables. Omics offers promising
solutions to address concerns about quality attri-
bute problems and biological explanations for harvested
produce deteriorations. These methods involve selecting
slower ripening lines, modifying ethylene responses, and
reducing the softening rate to extend postharvest life
and maintain quality [28]. In the past decade, the
genome of many tropical fruits such as cassava [29], ci-
trus [30], papaya [11], banana [31], durian [32], pome-
granate [33], pineapple [34], mango [35], lychee [36],
and sweet potato [37] is sequenced, which has been
possible because of technological advancement of se-
quencing approaches in genomics. The genome se-
quence of tropical crops not only provides insights into
their molecular evolution and genetics but also offers
prospects for developing molecular tools and breeding
for improving fruit quality traits. Moreover, emerging
computational tools such as artificial intelligence and
machine learning have paved the way for predicting
gene sequence and functional relationship of plant me-
tabolic pathways and biological processes, particularly in
the rapidly accumulating but ill-annotated genomes,
including tropical crops’ genome [38].
Roadblocks and future prospects
Although biotechnological tools and their advancement
have successfully implemented for reducing the tropical
fruit loss, however, they are still underutilized as com-
pared with temperate fruits. The biotechnological ap-
plications are mainly restricted to fewer fruits, including
banana, citrus, and sweet potato, and their usage remains
challenging in many other tropical fruits. The reasons
largely include the difficulty in genetic transformation,
recalcitrant nature, complex tissue culture systems, and
lack of genomic resources [6,7]. Therefore, developing
and optimizing robust genetic transformation and re-
generation methods are essential for broader bio-
technological applications and trait improvement of
tropical crops. Another concern for fruit bioengineering
is pleotropic effect, sometimes caused by constitutive
expression of foreign genes in plants, therefore, use of
tissue-specific, for example, fruit-specific promoters can
be useful for delivering genes in fruits or other edible
parts [39]. Recently, H2S, NO, and H2O2 have been
discovered as signaling molecules, which regulates pre-
and postharvest fruit traits [40–42]. Therefore, these
molecules and their biosynthesis and signaling genes
might be served as candidates for bioengineering of
tropical fruits and vegetables.
Current Opinion in Biotechnology 2023, 79:102850
6 Food Biotechnology
Many tropical fruit and vegetable crops are also prone to
several physiological disorders such as fruit cracking,
spongy tissue, bumpy fruits, chock throat, granulation,
pink berries, purple mark, and so on, which results in
loss in fruit quality and yield. To better understand the
molecular mechanisms behind fruit cracking, high-
throughput RNA sequencing and omics were recently
used to assess the differential expression of genes linked
to plant metabolism, such as cell wall metabolism, in
cracked fruits versus noncracked fruits. However, there
is still a need to investigate the molecular basis of the
aforementioned disorders and to investigate the gene
functions that are responsible for these phenotypic traits.
The biotechnological approaches and genetic en-
gineering could be served as sustainable tools to study
the precise mechanism behind a given cause and the
solutions to overcome it.
Among all knockdown and knockout approaches,
CRISPR–Cas9 technology offers sustainable solutions in
an efficient way for reducing losses of tropical fruits and
vegetables, however, it holds several limitations. One of
the limitations is the off-target effect, therefore, thor-
ough studies and caution are necessary for engineering
tropical crops using CRISPR–Cas9. For this, a more ef-
ficient and precise CRISPR–Cas9 technology based on
ribonucleoproteins (RNPs), which has been recently
applied to fruit crops, could be useful for many tropical
crops bioengineering [43]. This technology offers a
DNA-free editing with minimal off-target effects and
reduced cell toxicity due to rapid degradation of RNPs
by endogenous proteases and nucleases. Another con-
cern with biotechnological approaches, including the
genome editing, is the regulatory requirements, parti-
cularly in the context of tropical crops. This not only
limits investment by the fruit industry but also curtails
the efforts by the plant breeders and biotechnologists
working in this area. In the last few years, many coun-
tries have relaxed the regulatory norms particularly for
CRISPR–Cas9-mediated genome-engineered crops and
some crops have been approved for commercial use. One
of the interesting examples for this is CRISPR-edited
tomato enriched with γ-aminobutyric acid, which has
been approved by Japanese government in 2021 [44].
Conclusion
In this article, we primarily focused on the applications
of current biotechnological tools in genetic and genome
engineering of tropical crops for reducing losses of pro-
duce. An integrated approach combining breeding, bio-
technology, omics, and computational biology may
provide novel ways for sustainable cultivation of tropical
crops and improving fruit yield and postharvest stability.
It is evident that CRISPR has more advantages than
other approaches, therefore in future, developing
CRISPR-edited tropical crops offers a sustainable
Current Opinion in Biotechnology 2023, 79:102850
solution in reducing the losses, eventually benefiting
growing countries economically and for securing food for
rapidly increasing global population.
Authors contribution
MI and MWS designed the article. MI, PK, MFA
and MWS wrote and revised the article. All authors ap-
proved the final version.
Conflict of interest statement
All authors declare that they have no conflicts of interest.
Data Availability
Data will be made available on request.
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