SPRINGER BRIEFS IN FOOD, HEALTH,

AND NUTRITION

Anthony Keith Thompson

Suriyan Supapvanich
Jiraporn Sirison

Banana
Ripening
Science and
Technology

123
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Anthony Keith Thompson • Suriyan Supapvanich
Jiraporn Sirison

Banana Ripening
Science and Technology
Anthony Keith Thompson Suriyan Supapvanich
King Mongkut’s Institute King Mongkut’s Institute
of Technology Ladkrabang of Technology Ladkrabang
Bangkok, Thailand Bangkok, Thailand

Jiraporn Sirison
King Mongkut’s Institute
of Technology Ladkrabang
Bangkok, Thailand

ISSN 2197-571X     ISSN 2197-5728 (electronic)

SpringerBriefs in Food, Health, and Nutrition
ISBN 978-3-030-27738-3    ISBN 978-3-030-27739-0 (eBook)
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Preface

There have been many textbooks published dealing exclusively with Musa (bananas
and plantains) perhaps starting with the one written by my former boss and mentor
Professor C.W. Wardlaw (Banana Diseases: Including Plantains and Abaca 24 edi-
tions published between 1961 and 1974) to several more excellent books. Also,
several chapters in various books have also dealt with bananas.
The objective of this book was therefore to deal exclusively with how bananas
are ripened. However, the technology used is based on, and has implications with, a
whole range of factors. These include the cultivar, growing conditions and how and
at what maturity the fruit are harvested and handled. Also, various postharvest treat-
ments can be applied to fruit and how these impact with the ripening process can
have effects on the fruit quality.
The ripening process and the changes that occur are therefore considered in detail.
There is a well-established successful technology that has evolved and is applied, par-
ticularly in more developed temperate countries where bananas are imported, but this
technology essentially applies to a single genotype. However, most bananas are mar-
keted locally in the country where they are grown, and often, different technologies are
used in the ripening process, which have technical, economic and health implication.
All these factors are discussed, and interacting implications are considered. Also, there
is often insufficient and even contradictory information on how different ripening
methods affect fruit quality including carbohydrates, proteins, phenolic, vitamins, acid-
ity and other phytochemicals as well as texture, colour and flavour.
Banana sea trade began a little more than 100 years ago, but it has helped create
great economic empires and sustained the livelihoods of millions of people in pro-
ducer countries as well as in importing countries. The international banana trade has
been very fast in employing new advancements in refrigerated transport and ripen-
ing technologies, since they can reduce transport costs and losses and increase the
distance they can be successfully transported.

 Anthony Keith Thompson

  Suriyan Supapvanich
  Jiraporn Sirison

v
Abbreviations

1-MCP 1-methylcyclopropene
AA ascorbic acid
ABA abscisic acid
ACC 1-aminocyclopropane-1-carboxylic acid
ACC oxidase an enzyme involved in biosynthesis of ethylene
ACC synthase an enzyme involved in biosynthesis of ethylene
ACC synthesis the reaction catalysed by ACC synthase
ACPd activated carbon impregnated with palladium
ATP adenosine triphosphate
AVG aminoethoxyvinylglycine
CA controlled atmosphere
CMC carboxymethyl cellulose
DFE dietary folate equivalent
DPPH 1,1-diphenyl-2-picrylhydrazyl
Ethrel ethephon (2-chloroethylphosphonic acid)
GA gibberellins
GA3 gibberellic acid
IAA indole-3-acetic acid
IC inclusion complex powder for slow ethylene release, ethylene-
alpha-cyclodextrin
Imazalil 1-[allyloxy-2,4-dichlorophenethyl]imidazole, a fungicide
LDPE low-density polyethylene film
LPE lysophosphatidylethanolamine
MA modified atmosphere
mRNA messenger ribonucleic acid
PAL phenylalanine ammonia-lyase
PE polyethylene film
PG polygalacturonase
PME pectin methylesterase
POD pyrogallol peroxidase
PPO polyphenol oxidase

vii
viii Abbreviations

pVACs provitamin A carotenoids

RAE retinol activity equivalent
RH relative humidity
RNA ribonucleic acid
SAM s-adenosyl-methionine
TA total acidity; titratable acidity
TBZ thiabendazole (2-thiazol-4-yl benzimidazole), a fungicide
ton 2240 pounds (lbs) = 1016 kg
tonne 1000 kg
TPC total phenolic compounds
TFC total flavonoid compounds
TSS total soluble solids
ULO ultralow oxygen
Contents

1 Introduction����������������������������������������������������������������������������������������������     1
Musa Taxonomy����������������������������������������������������������������������������������������     2
Musa Breeding Programmes����������������������������������������������������������������������     4
Genotypes in International Trade��������������������������������������������������������������     5
Some Common Genotypes������������������������������������������������������������������������     6
2 Preharvest Effects������������������������������������������������������������������������������������    13
Fertilizers ��������������������������������������������������������������������������������������������������    13
Organic Production������������������������������������������������������������������������������������    14
Light and Day Length��������������������������������������������������������������������������������    14
Disease ������������������������������������������������������������������������������������������������������    15
Water Stress�����������������������������������������������������������������������������������������������    18
Damage������������������������������������������������������������������������������������������������������    18
Bunch Covers��������������������������������������������������������������������������������������������    19
Harvest Maturity����������������������������������������������������������������������������������������    20
3 Fruit Ripening������������������������������������������������������������������������������������������    25
Pre-Climacteric Phase��������������������������������������������������������������������������������    26
Ripening Phase������������������������������������������������������������������������������������������    28
Internal Ethylene����������������������������������������������������������������������������������������    28
Effects of Ethylene Post Ripening Initiation����������������������������������������������    31
Genetic Effects on Ripening����������������������������������������������������������������������    32
Maturity of Fruit and Response to Ethylene����������������������������������������������    34
Factors Affecting Fruit and Response to Ethylene��������������������������������    37
Changes that Occur During Ripening��������������������������������������������������������    37
Peel Colour��������������������������������������������������������������������������������������������    37
Peel Spotting������������������������������������������������������������������������������������������    40
Peel Chemicals��������������������������������������������������������������������������������������    40
Finger Drop��������������������������������������������������������������������������������������������    41
Weight Loss�������������������������������������������������������������������������������������������    41
Moisture ������������������������������������������������������������������������������������������������    43
Texture ��������������������������������������������������������������������������������������������������    44

ix
x Contents

Flavour and Aroma��������������������������������������������������������������������������������    46

Minerals ������������������������������������������������������������������������������������������������    47
Carbohydrates����������������������������������������������������������������������������������������    48
Protein����������������������������������������������������������������������������������������������������    49
Phenolics������������������������������������������������������������������������������������������������    49
Acidity����������������������������������������������������������������������������������������������������    50
Ascorbic Acid����������������������������������������������������������������������������������������    51
Carotenoids and Vitamin A��������������������������������������������������������������������    52
Folates����������������������������������������������������������������������������������������������������    54
Other Phytochemicals����������������������������������������������������������������������������    54
4 Postharvest Treatments to Control Ripening����������������������������������������    57
Controlled Atmosphere Storage����������������������������������������������������������������    57
CA on Pre-Climacteric Bananas������������������������������������������������������������    57
CA Effects After Ripening Initiation ����������������������������������������������������    58
Hypobaric Storage ������������������������������������������������������������������������������������    59
Modified Atmosphere Packing������������������������������������������������������������������    60
Ethylene Absorbents����������������������������������������������������������������������������������    62
Chemicals��������������������������������������������������������������������������������������������������    64
Metal Ions����������������������������������������������������������������������������������������������    64
1-Methylcyclopropene ��������������������������������������������������������������������������    64
Salicylic Acid ����������������������������������������������������������������������������������������    67
Gibberellic Acid ������������������������������������������������������������������������������������    67
Diazocyciopentadiene����������������������������������������������������������������������������    68
Indole-3-Acetic Acid������������������������������������������������������������������������������    68
Abscisic Acid ����������������������������������������������������������������������������������������    68
Lysophosphatidylethanolamine��������������������������������������������������������������    69
Aminoethoxy-Vinylglycine��������������������������������������������������������������������    69
Nitrous Oxide����������������������������������������������������������������������������������������    70
Maleic Acid��������������������������������������������������������������������������������������������    70
Coatings ����������������������������������������������������������������������������������������������������    71
Irradiation��������������������������������������������������������������������������������������������������    72
Temperature ����������������������������������������������������������������������������������������������    74
Humidity����������������������������������������������������������������������������������������������������    76
5 Initiation of Ripening������������������������������������������������������������������������������    79
Ethylene ����������������������������������������������������������������������������������������������������    80
Cylinders������������������������������������������������������������������������������������������������    82
Catalytic Generators������������������������������������������������������������������������������    84
Ethrel������������������������������������������������������������������������������������������������������    84
Encapsulation����������������������������������������������������������������������������������������    87
Acetylene ��������������������������������������������������������������������������������������������������    88
Calcium Carbide������������������������������������������������������������������������������������    89
Sensory Analysis������������������������������������������������������������������������������������    91
Toxicity of Calcium Carbide������������������������������������������������������������������    92
Propylene ��������������������������������������������������������������������������������������������������    93
Contents xi

Esters����������������������������������������������������������������������������������������������������������    93
Alcohol������������������������������������������������������������������������������������������������������    93
Carbon Monoxide��������������������������������������������������������������������������������������    94
Smoking ����������������������������������������������������������������������������������������������������    95
Kerosene������������������������������������������������������������������������������������������������    95
Incense ��������������������������������������������������������������������������������������������������    96
Heat��������������������������������������������������������������������������������������������������������    96
Damage and Stress������������������������������������������������������������������������������������    97
Fruit Generation ����������������������������������������������������������������������������������������    99
Leaves��������������������������������������������������������������������������������������������������������   100
6 Ripening Technology��������������������������������������������������������������������������������   101
Ripening Rooms����������������������������������������������������������������������������������������   101
Modelling��������������������������������������������������������������������������������������������������   105
Transport����������������������������������������������������������������������������������������������������   106
Reducing Ripening Initiation in Transit����������������������������������������������������   107
Ripening in Transit������������������������������������������������������������������������������������   108
7 Conclusions����������������������������������������������������������������������������������������������   111

References ������������������������������������������������������������������������������������������������������   113

Index����������������������������������������������������������������������������������������������������������������   137
About the Authors

Jiraporn Sirison is Associate Dean and Assistant

Professor in the Faculty of Agro-Industry, King
Mongkut’s Institute of Technology, Ladkrabang,
Thailand. She studied at Maejo University and
Mahidol University both in Thailand and Kyoto
University in Japan. She is a Nutrition Chemist whose
teaching and research are mainly in the area of the
effects of processing on nutrients in food and the
application of food colloids in food nutrition. She
teaches nutrition to undergraduates and has published
many research papers on these subjects. Also, she was
brought up on her parents’ banana farm in Thailand.

Suriyan Supapvanich is Assistant Professor in

Postharvest Technology in the Faculty of Industrial
Education and Technology, King Mongkut’s Institute
of Technology, Ladkrabang, Thailand. His field of spe-
cialization is postharvest biology and biochemistry of
tropical fruit and vegetables. He studied at Kasetsart
University and King Mongkut’s University Thonburi,
both in Thailand, and the University of Nottingham,
UK, where he was awarded a PhD degree. He subse-
quently worked at the University of Natural Resources
and Life Sciences in Austria and lectured in the Faculty
Natural Resources and Agro-Industry at Kasetsart
University. He has 19 years teaching experience in
courses on postharvest technology of agricultural prod-
ucts and introduction of food science and technology to
undergraduate students and advanced postharvest tech-
nology to MSc students. He has also published more
than 40 papers in international journals and 2 book

xiii
xiv About the Authors

chapters. He is currently coordinating postharvest

research projects with KMITL Prince of Chumphon
Campus, KMUTT, Thammasat University and Anhui
Agricultural University in China.

Anthony Keith Thompson is Visiting Professor at

King Mongkut’s Institute of Technology Ladkrabang
in Thailand and was formerly Professor of Plant
Science, University of Asmara, Eritrea; Professor of
Postharvest Technology and Head of Department,
Cranfield University, UK; Team Leader on an EU proj-
ect on restructuring the banana industry in the
Windward Islands; Principal Scientific Officer,
Tropical Products Institute, London; Postharvest
Expert for the UN in Sudan, Yemen and Korea for the
Food and Agriculture Organization, Ghana and Sri
Lanka for the International Trade Centre and Gambia
for the World Bank; Advisor to the British, Jamaican
and Colombian Governments in postharvest technol-
ogy of fruit and vegetables; Research Fellow in Crop
Science, University of the West Indies, Trinidad;
Demonstrator in Biometrics, University of Leeds; as
well as Consultant for various commercial and govern-
ment organizations in many countries throughout the
world. He has published over 100 scientific papers and
many scientific textbooks.
Chapter 1
Introduction

Bananas grow everywhere in the tropics as well as in many sub-tropical countries.

FAOStat (2019) estimated that the area of production of bananas in 2017 was about
5.6 million hectares producing some 114 million tonnes. There are many genotypes,
most of which are ripened and eaten as fresh fruit. Others may be cooked before
eating either unripe or ripe. For eating fresh they are harvested before they are ripe
and then either allowed to ripen or treated in some way to initiate the ripening pro-
cess. The most obvious change in bananas during ripening is their peel colour. This
has been codified into what is called a ripening index or a colour index (Table 1.1;
Fig. 1.1), which is commonly used in many countries for communication in the
ripening and retail trades and among research and extension workers for all the
genotypes.
About 15% of the world’s banana production is for the export trade, and is based
on ‘Cavendish’ (Heslop-Harrison and Schwarzacher 2007). International trade in
bananas is crucial for the economies of many developing countries and bananas are
popular throughout the world, but can only be grown successfully in the tropics and
to a much lesser extent in the sub-tropics. Therefore, an enormous quantity of
bananas is transported around the world every day, which contribute to climate
change. Ripening centres and retail distribution represent approximately 10% of
total emissions of greenhouse gases in the banana value chain, from which 75%
correspond to energy consumption, 22% to distribution centres and 1% to ethylene
production (Calberto et al. 2015). Overseas transport and primary production of
bananas were the main contributors to the total greenhouse gas emissions including
the consumer stage that resulted in a 34% rise in carbon footprint, mainly due to
high wastage (Svanes and Aronsson 2013).

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2019 1

A. K. Thompson et al., Banana Ripening, SpringerBriefs in Food, Health,
and Nutrition, https://doi.org/10.1007/978-3-030-27739-0_1
2 1 Introduction

Table 1.1 Changes in peel colour and starch and sugar content of pulp of ‘Gros Michel’ during
ripening, based on data from United Fruit Company 1942. Source: Wardlaw (1961) with
modifications
Ripening index Peel colour Total sugars (%) Starch (%)
1 Green 0.1–2.0 19.5–21.5
2 Green – trace of yellow 2.0–5.0 16.5–19.5
3 More green than yellow 3.5–7.0 14.5–18.0
4 More yellow than green 6.0–12.0 9.0–15.0
5 Green tips 10.0–18.0 2.5–10.5
6 All yellow 16.5–19.5 1.0–4.0
7 Yellow flecked with brown 17.5–19.0 1.0–2.5
8 Yellow with large brown spots 18.5–19.0 1.0–1.5

Fig. 1.1 Time lapse photographs of a single ‘Valery’ banana after 24 h exposure to 1000 μL L−1
ethylene and then kept at 20 °C and 90% RH

Musa Taxonomy

Bananas belong to the family Musaceae, genus Musa. Christelová et al. (2011)
reported that Musa was divided into the following sections: Ingentimusa with a
chromosome number 2n = 14, Australimusa has been merged with Callimusa and
has a chromosome number 2n = 20 (Musa beccarii, which is part of Callimusa sec-
tion, has 18 chromosomes) and Eumusa and Rhodochlamys with chromosome num-
bers of 2n = 22. Fe’i belong to Callimusa and are easily recognized by their erect
bunches.
Carl Linnaeus in 1783 gave the name M. sapientium L. to bananas that are eaten
fresh and M. paradisiaca L. for those that are cooked before eating. Many authors
still use the Linnaean classification and many other names have been used, but
Cheesman (1947–1949) defined M. balbisiana Colla and M. acuminata Colla as the
basis for almost all cultivated bananas and plantains. Simmonds and Shepherd
Musa Taxonomy 3

(1955) and Stover and Simmonds (1987) confirmed this and reported that many
desert varieties are derived from M. acuminata, some being diploid and a few being
tetraploid but most being triploid (2n = 3x = 33). M. balbisiana has also contributed
to the origin of desert bananas and plantains by hybridisation with M. acuminata.
For example, ‘Pelipita’ (Musa ABB) has 8 M. acuminata chromosomes and 25 M.
balbisiana chromosomes. Simmonds and Shepherd (1955) recommended that in
place of the species name, an A genome for M. acuminate and a B genome M. bal-
bisiana should be used showing the origin and contribution of the two species. So,
a triploid genotype whose origin is M. acuminate, e.g. ‘Giant Cavendish’ and ‘Gros
Michel’ would be referred to as Musa AAA. A triploid that has one third M. bulbi-
siana and two thirds M. acuminate would be referred to as Musa AAB. They also
recommended a subgroup within a species for example Musa AAA (Cavendish sub-
group) ‘Robusta’, Musa AA ‘Pisang Mas’ and Musa ABBB ‘Kluai Teparod’. The
centre of origin is thought to be South-East Asia, where they occur from India to
Polynesia (Simmonds 1962). The centre of diversity has been placed in Malaysia or
Indonesia (Daniells et al. 2001), although considerable diversity is known through-
out the range. The centre of origin of M. acuminata is thought to have been in
Malaysia or Indonesia (Simmonds 1962) and M. balbisiana in India, Myanmar,
Thailand and Philippines (Daniells et al. 2001). The origin of Fe’i is thought to be
the Pacific Islands probably derive mainly from Musa maclayi F.J.H. von Mueller
ex N.N. Miklouho-Maclay or Musa × troglodytarum L. They can be classified, for
example, as Musa (Fe’i Group) ‘Utafun’.
A taxonomic scoring method is used to classify the edible bananas and to pro-
vide evidence on their evolution. Edible diploid forms of M. muminata are thought
to be the primary source of the whole group to which another species, M. balbisi-
ana, has contributed by hybridization. Thus, there exist diploid and triploid edible
forms of M. muminata and diploid, triploid and tetraploid hybrid types of genetic
constitutions that vary according to their history. There is a slight possibility that a
third wild species has contributed to the origins of a small group of triploid hybrid
types. Triploidy was probably established under human selection for vigour and
fruit size; tetraploidy is rare. Indo-Malaysia and Malaysia are probably the primary
centre of origin of the group. The species M. paradisiaca and M. sapientum, named
by Linnaeus to identifiable edible varieties, are both shown to be of hybrid origin
(Simmonds and Shepherd 1955; Deepthi 2016).
Almost all genotypes used commercially in international trade have arisen from
mutations of Musa AAA (‘Cavendish’ subgroup) selected in the field. These include
‘Grand Naine’, ‘Lacatan’, ‘Poyo’, ‘Robusta’, ‘Valery’ and ‘Williams’, which are
largely distinguished from each other by the height of the pseudostem. In addition,
‘Dwarf Cavendish’ is tolerant to a wide range of climates including cool conditions.
‘Grand Naine’ responds well to optimum growing conditions, but does not grow
or yield well in sub-optimum conditions including soils with low fertility or insuf-
ficient rainfall or irrigation. The ‘Sucrier’, ‘Pisang Mas’, ‘Honey’ genotypes are
very sweet and have small fruit, thin skins, yellowish flesh and small bunches (up to
about 13 kg). ‘Gros Michel’ was the fruit that dominated the international trade
from its beginnings in the mid-nineteenth century until the late 1940s. It has long
4 1 Introduction

been grown in South-East Asia and Sri Lanka. Jean Francois Pouyat, a French bota-
nist and chemist who settled in Jamaica in 1820 probably introduced it to the
Caribbean region. He brought the fruit from Martinique to his coffee estate in
Jamaica where it was originally called ‘Banana Pouyat’ then later this became the
‘Martinique Banana’ and finally ‘Gros Michel’ and ‘Big Mike’ in USA and ‘Hom
Thong’ in Thailand. The Agricultural Society in Jamaica awarded Pouyat a dou-
bloon for introducing such a valuable genotype (Von Loesecke 1949; Davies 1990).
It is a tall heavy bearing genotype, but is susceptible to Panama disease (caused by
the fungus Fusarium oxysporum f. sp. cubense). It was reported as early as 1928 in
Trinidad that there was concern about the susceptibility of ‘Gros Michel’ to Panama
disease (Wardlaw 1961). Three races of Panama disease have been described that
can affect bananas. Race 1 caused the epidemics on ‘Gros Michel’ and also affects
some other varieties and tetraploids. Race 2 affects cooking bananas e.g. ‘Bluggoe’
and some tetraploids while Race 4 affects race 1 and race 2 susceptible varieties as
well as the ‘Cavendish’ varieties. Race 4 has been shown to consist of a sub-tropical
and a tropical race. The tropical race 4 can attack unstressed plants while sub-­
tropical race 4 usually only attacks when plants are in a stressed condition. ‘Namwa’
(AAB) was reported to be very resistant to Fusarium oxysporum f. sp. cubense,
tropical race 4. Infection incidence were less than 1% while susceptible varieties
like ‘Lakatan’ and ‘Cavendish’ sustained more than 90% in the first crop. ‘Namwa’
was also observed resistant to Banana Bunchy Top Virus (BBTV) and Black
Sigatoka (Molina 2019).

Musa Breeding Programmes

Breeding programmes have developed tetraploid bananas and plantains, starting in

1922 at the Imperial College of Tropical Agriculture in Trinidad. This programme
was subsequently transferred to establish the banana breeding programme at Bodles
in Jamaica (producing, among others, ‘Bodles Altafort’ which is Musa AAAA).
Other breeding programmes include the Centro Nacional de Pesquisa de Mandioca
e Fruticultura Tropical and Empresa Brasileira de Pesquisa Agropecuária
(EMBRAPA) that were established in Brazil and the Fundacion Hondureña de
Investigacion Agricola (FHIA) that was originally established at a research station
of the United Fruit Company in Honduras. These programmes have largely concen-
trated in breeding tetraploids. ‘FHIA-21’ (Musa AAAB), with cooking banana with
plantain in its pedigree, has been reported to be resistant to Black Sigatoka disease.
‘FHIA-01’, also called ‘Goldfinger’ (Musa AAAB), is a desert banana that has been
bred to be high yielding and resistant to Black Sigatoka, however this resistance has
been questioned. There is also concern about the tetraploids containing M. balbisi-
ana because the Banana Streak Virus (BSV) is integrated in the B genome and may
be activated, especially if they are exposed to stress or are propagated by tissue
culture (Jones 2009). The International Institute of Tropical Agriculture (IITA) in
Nigeria and Kenya have programmes involving crossing Musa AA with Musa AAB
Genotypes in International Trade 5

to generate triploid and tetraploids (Musa AAAB) with improved disease resistance
and agronomic characters. Several cultivars have been released by ITA that are
either triploid or tetraploid and are given numbers following the initials, PITA for
plantains, for example PITA 14, and BITA for cooking bananas, for example BITA
3. Centre Africain de Recherches sur Bananiers et Plantains (CARBAP) in West
Africa and Centre de cooperation Internationale en Recherche Agronomique pour le
Développement (CIRAD) in the Guadeloupe also have banana breeding projects.

Genotypes in International Trade

Currently almost all varieties used commercially in international trade have arisen
from mutations of Musa AAA ‘Cavendish’ subgroup (Fig. 1.2) selected in the field
including ‘Grand Naine’, ‘Lacatan’, ‘Poyo’, ‘Robusta’, ‘Valery’ and ‘Williams’.
From their origin in South-East Asia some authorities report that bananas were
introduced into Africa in the early middle ages, and in 1402 Portuguese colonists
took banana plants from Guinea to the Canary Islands. They were thought to have
been taken to the Americas by Portuguese explorers in the late fifteenth century. In
1826 Charles Telfair obtained ‘Cavendish’ plants from southern China and took
them to Mauritius and in 1829 some plants were sent from there to England. In
1834, the 6th Duke of Devonshire wrote to the Chaplain at Alton Towers: “My Dear
Sir, A thousand thanks for the banana, it arrived quite safe and I am delighted to
have an opportunity of seeing that most beautiful and curious fruit. It is the

Fig. 1.2 ‘Giant

Cavendish’ bunch ready
for harvesting in St Lucia
6 1 Introduction

admiration of everybody and has been feasted upon at dinner today according to the
directions”. The Duke eventually obtained plants at Chatsworth House, which is the
seat of the Dukes of Devonshire. Their family name is ‘Cavendish’ and the Duke
named it Musa cavendishii. Joseph Paxton was their head gardener who cultivated
the bananas and got them to produce fruit in a conservatory at Chatsworth. A few
years later the Duke supplied two cases of plants to a missionary named John
Williams, destined for Samoa. Only one M. cavendishii plant survived the journey
and that single specimen was the forebear of bananas that flourish in Samoa and
other South Sea Islands. Probably in 1853 some plants were received from
Chatsworth and although they were in poor condition some survived and were dis-
tributed among the islands. Also, the first banana plant from Chatsworth may also
have been the origin of stock introduced to the Canaries in 1855 (Anonymous 2012).
Apparently, suckers were also sent to the West Indies about the same time.

Some Common Genotypes

As indicate above the botanical names for different bananas are based on the publi-
cations of Cheesman (1947–1949) and subsequently those of Simmonds and
Shepherd (1955) and Stover and Simmonds (1987). A cultivar is a plant that has
been produced in cultivation and a variety is a naturally occurring form of plant,
both are within a species. Therefore, most of the names used for bananas could be
called varieties since they have occurred naturally through selections of somatic
mutations from a cultivated population. However, a few, particularly the tetraploids,
are cultivars since they have been specifically bred for certain desirable
characteristics.
There are many local names used for different types of banana therefore deciding
which names are the same and which names are different is difficult and at best
arbitrary. Some botanical names are different depending on the references, for
example ‘Bluggoe’ is given as Musa AAB (Christelová et al. 2017) and Musa ABB
(Porcher 1998), but for simplicity the one given by INIBAP (Daniells et al. 2001)
will be used. Also, transliterations between languages can cause confusion. In some
cases, different botanical names are given for fruit with the same common name.
Therefore, the following list of banana genotypes, mostly referred to in this book, is
by no means definitive, just an indication and an attempt to reduce confusion. With
the susceptibility of bananas and plantains to somatic mutation, many small-scale
growers will have their own often exclusive selection that may be unique. Here they
are listed under common names to provide a reference. For a much more detailed
and comprehensive description, refer to Stover and Simmonds (1987), Daniells
et al. (2001) and the Musa Germplasm Information System.
‘Apem’ Musa AAB (plantain subgroup).
‘Apple banana’ Musa AA ‘Manzano’ has a sub-acid flavour and should be allowed
to ripen fully before eating.
Some Common Genotypes 7

‘Amritsagar’ Musa AAA is a good table variety in Assam with medium sized plant,
good size fruit and medium thick rind, and the ripe banana develops a bright yel-
low colour (Goswami and Handique 2013).
‘Bananito’ Musa AA ‘Kluai Namwa’ ‘Lady Finger’ (Fig. 1.3).

‘Bluggoe’ Musa ABB ‘Kluai Som’, ‘Horse Plantain’ tends to be highly susceptible
to Fusarium oxysporum f. sp. cubense race 1, 2 and 4 (Jones 2000).
‘Bodles Altafort’ Musa AAAA developed from the banana breeding programme at
Bodles in Jamaica by Ken Shepherd in the 1960s and was a cross between ‘Gros
Michel’ (AAA) and ‘Pisang Lilin’ (AA).
‘Cavendish’ Musa AAA. has been the main group of varieties in international trade
since the late 1940s and selections include ‘Grand Naine’, ‘Lacatan’, ‘Poyo’,
‘Robusta’, ‘Valery’ and ‘Williams’, which are largely distinguished from each
other by the height of the pseudostem. The tallest is ‘Lacatan’ (up to 5.5 m) fol-
lowed by ‘Robusta’ and ‘Giant Cavendish’ (up to 5 m) and the smallest is the
‘Dwarf Cavendish’ (1.2–2.1 m). ‘Grand Naine’ responds well to optimum grow-
ing conditions, but does not grow or yield well in sub-optimum conditions
including soils with low fertility or insufficient rainfall or irrigation.
‘Cheni Champa’ Musa AAB is one of the hardiest medium tall bananas in Assam,
resistant to Fusarium wilt and fairly resistant to bunchy top disease with small
size fruits, thin peel, creamy pulp and sub-acid taste (Goswami and Handique
2013).
‘Giant Cavendish’ Musa AAA is synonymous with ‘Cavendish’.
‘Dominico’ Musa AAB (plantain subgroup).
‘Dominico Harton’ Musa AAB (plantain subgroup).
‘Dwarf Brazilian’ Musa AAB.
‘Dwarf Cavendish’ Musa AAA (Cavendish subgroup). ‘Dwarf Cavendish’ is toler-
ant to a wide range of climates including cool conditions.
‘FHIA-01’ Musa AAAB ‘Goldfinger’, is a desert banana that can also be used in
cooking when green. It was bred to be high yielding and resistant to Black

Fig. 1.3 Musa AA on sale

in UK supermarkets in
2018 as ‘Bananitos’
8 1 Introduction

­ igatoka. Mean bunch weight 34.1 lbs, taste 3.2 and acceptability 3.7 (Nelson
S
and Javier 2007).
‘FHIA-02’ Musa AAAA ‘Mona Lisa’ is sweet and similar to Cavendish. Mean
bunch weight 23.7 lbs, taste 3.0 and acceptability 3.6 (Nelson and Javier 2007).
‘FHIA-03’ Musa AABB ‘Sweetheart’ a cooking banana that can also be used for
dessert. Mean bunch weight 36.4 lbs, taste 3.0 and acceptability 3.6 (Nelson and
Javier 2007).
‘FHIA-17’ Musa AAAA a ‘Gros Michel’ type dessert banana that can also be
cooked. Mean bunch weight 27.4 lbs, taste 3.3 and acceptability 4.2 (Nelson and
Javier 2007).
‘FHIA-18’ Musa AAAB a sweet acid apple flavoured dessert banana. Mean bunch
weight 27.6 lbs, taste 2.9 and acceptability 3.8 (Nelson and Javier 2007).
‘FHIA-21’ Musa AAAB is a French plantain type.
‘FHIA-23’ Musa AAAA dessert banana. Mean bunch weight 26.7 lbs, taste 3.0 and
acceptability 3.7 (Nelson and Javier 2007).
‘Grand Naine’ Musa AAA (Cavendish subgroup). It responds well to optimum
growing conditions, but does not grow or yield well in sub-optimum conditions
including soils with low fertility or insufficient rainfall or irrigation. Mean bunch
weight 18.3 lbs, taste 3.2 and acceptability 3.1 (Nelson and Javier 2007).
‘Gros Michel’ Musa AAA dominated the international trade from its beginnings in
the mid-nineteenth century until the late 1940s. It is a tall heavy bearing geno-
type, but is susceptible to Panama disease (caused by the fungus Fusarium oxys-
porum f. sp. cubense). It is still grown commercially, for example in parts of
Colombia and Thailand where it is mainly called ‘Kluai Hom Thong’.
‘Guineo’ Musa AAA.
‘Guineo Negro’ Musa AAA.
‘Hari chhal’ (Musa acuminata) grown commercially in Sindh Province in Pakistan
and in India.
‘Harton’ Musa AAB (plantain subgroup).
‘Highgate’ Musa AAA semi-dwarf mutant of ‘Gros Michel’.
‘Honey’ Musa AA.
‘Horn’ Musa AAB (plantain subgroup).
‘Khai’ Musa AA (Fig. 1.4).

‘Karpooravalli’ ‘Karpuravalli’ Musa ABB.

‘Khuai Hom Khiew Korm’ synonymous with ‘Dwarf Cavendish’ Musa AAA
(Fig. 1.5).

‘Kluai Hom Khiew’ Musa AAA (Cavendish subgroup) synonymous with ‘Pisang
Masak Hijau’.
‘Kluai Hom Taiwan’ Musa AAA synonymous with ‘Gros Michel’.
‘Kluai Hom Thong’ Musa AAA synonymous with ‘Gros Michel’.
‘Kluai Hom’ Musa AAA (Cavendish subgroup).
‘Kluai Hug Mook’ Musa AA synonymous with ‘Silver Bluggoe’, ‘Kluai Som’.
Some Common Genotypes 9

Fig. 1.4 ‘Khai’ fruit at harvest and after storage at 13 or 25 °C

Fig. 1.5 ‘Hom Khiew’ on

sale in Thailand in 2019

‘Kluai Khai’ Musa AA synonymous with: ‘Kluai kha’ ‘Pisang Mas’, ‘Sunny
Bunch’, ‘Golden’, ‘Sucrier’.
‘Kluai Leb Mua Nang’ Musa AA (Fig. 1.6).

‘Kluai Nam Wah’ Musa AA synonymous with: ‘Pisang Awak’, ‘Kluai Hin’ and
‘Saba’.
‘Kluai Nark’ Musa AAA (Cavendish subgroup) synonymous with ‘Red banana’
‘Lacatan’.
‘Kluai Teparod’ Musa ABBB.
‘Lady Finger’ Musa AA (Fig. 1.7).

‘Latundan’ Musa ABB.

‘Malbhog’ Musa AAB synonymous with: ‘Amrithapani’, ‘Digjowa’, ‘Dudhsagar’,
‘Honda’, ‘Kozhikodu’, ‘Latundan’, ‘Madhuranga’, ‘Manzana’, ‘Nanjankode
Rasabale’, ‘Pisang Rasthali’ ‘Silk’, ‘Silk Fig’ ‘Saapkal’, ‘Sabri’, ‘Suvandal’ and
‘Thozhuvan’. It is a popular table banana in Assam with a sweet aroma, taste and
long postharvest life (Goswami and Handique 2013).
10 1 Introduction

Fig. 1.6 ‘Kluai Leb Mua Nang’ in Thailand in 2019 at harvest and after ripening in ambient con-
ditions (about 30 °C)

Fig. 1.7 Golden, Lady

Finger on sale in Thailand
in 2019

‘Namwa’ Musa AAB synonymous with: ‘Pisang Awak’, which is a popular banana
in Thailand and Malaysia. Flavours were similar to ‘Cavendish’ except that they
were stronger (Renoo Yenket, nd).
‘Nanica’ Musa AAA (Cavendish subgroup).
‘Ney Poovan’ Musa AB ‘Safet Velchi’, ‘Chini Champa’, ‘Ranel’, ‘Kisubi’, ‘Apple’,
‘Farine France’, ‘Lady’s Finger’ has a sweet, sub-acid flavoured fruit.
‘Orishele’ Musa AAB.
‘Pacovan’ Musa AAB, subgroup Prata. is mostly cultivated in Northeast Brazilian
but is also grown in India, Australia and the Western Pacific islands, where it is
known as ‘Pachanadan’ or ‘Pacha Naadan’, ‘Improved Lady’s Finger’ and
‘Lady’s finger’ respectively (Ploetz et al. 2007).
‘Pei Chiao’ Musa AAA (Cavendish subgroup) introduced to Taiwan from southern
China in the eighteenth century.
‘Petite Naine’ Musa AAA (Cavendish subgroup) synonyms include: ‘Extra-dwarf’,
‘Kiri Tia Mwin’ and ‘Dwarf Cavendish’.
Some Common Genotypes 11

‘Pisang Ambon Lumut’ Musa AAA.

‘Pisang Awak’ Musa AAB.
‘Pisang Mas’ Musa AA synonyms include: ‘Amas’, ‘Sucrier’, ‘Kluai Khai’, ‘Figue
Sucrée.
‘Pisang Lilin’ Musa AA synonyms include: Kluai Thong Ki Maew, Kaveri banana.
Pisang ekor kuda, Pisang empat puluh, Pisang lemak manis, Pisang lemak manis
terenganu, Pisang lidi, Pisang mas sagura, Pisang muli, Kluai Leb Mu Nang,
Kluai Thong Kab Dam, Kluai tong kee maew Kluai thong khi meew.
‘Pisang Ustrali’ Musa AAAA synonyms with Papua New Guinea banana.
‘PITA 3’ Musa AAAB.
‘PITA 314’ Musa AAAB.
‘PITA 24’ plantain Musa AAB.
‘Pouyat’ Musa AAA ‘Gros Michel’ is named after Jean Francois Pouyat, a French
botanist and chemist who settled in Martinique.
‘Poyo’ Musa AAA.
‘Prata‘Musa AAB. Approximately 60% of the harvested area of bananas in Brazil
are ‘Prata’, ‘Prata Anã’ or ‘Pacovan’ (Embrapa 2014).
‘Prata Anã’ Musa AAB synonyms with: ‘Lady’s finger’ and ‘Santa Catarina Prata’
in Hawai’i.
‘Rasabale’ Musa AAB (Silk subgroup).
‘Red Decca’ Musa AAA (Cavendish subgroup).
‘Red Banana’ Musa AAA (Cavendish subgroup).
‘Robusta’ Musa AAA (Cavendish subgroup).
‘Saba’ Musa BBB.
‘Santa Catarina Prata’ Musa AAA.
‘Sucrier’ Musa AA. ‘Pisang Mas’ and ‘Honey’ are very sweet and have small fruit,
thin skins, yellowish flesh and small bunches (up to about 13 kg). Plants are up
to 3.5 m high, prefer light shade and are not well adapted to cooler
temperatures.
‘Sugar’ Musa AAB.
‘Valery’ Musa AAA (Cavendish subgroup).
‘Williams’ Musa AAA (Cavendish subgroup).
Where bunch flavour and acceptability are given above, these are based on the scale
of Nelson and Javier (2007) where: 5 = excellent, 4 = very good, 3 = good, 2 = fair,
1 = poor.
Chapter 2
Preharvest Effects

The conditions and environment in which a crop is grown can affect its postharvest
life including ripening of fruit. Chillet and de Lapeyre de Bellaire (2002) found a
weak correlation between the manganese concentration and wound ethylene pro-
duction in lowland bananas grown in the West Indies. They also showed that in the
wet season lowland fruit were more fragile and produced more wound ethylene than
highland fruit. In Thailand ‘Williams’ and ‘Grand Naine’ bananas grown under low
chemical production systems tended to have higher levels of sugars and acids but
were softer and there was some indication that they contained less starch than those
from a conventional production system (Ambuko et al. 2006).

Fertilizers

No direct information could be found on the effects of fertilizer application to the

growing plant on the subsequent behaviour during ripening of bananas. However,
Ramesh Kumar and Kumar (2007) assessed potash foliar sprays on ‘Neypoovan’
bananas and found their postharvest green life was 4.5, 4.8, 5.2 and 5.3 days as a
result of foliar sprays with 0, 0.5, 1.0 and 1.5% P, respectively. Also, TSS, total
sugars and sugar acid ratio increased and weight loss and % acidity decreased as K
foliar sprays were increased. Patel et al. (2017) found marked improvement in fruit
quality of ‘Grand Naine’ when RDK (a concentrated N, P, S and Mg fertilizer) was
applied to the plants. The quality factors they were assessing were: pulp to peel
ratio, TSS, TA, ascorbic acid, reducing sugars, total sugars, sugar acid ratio and
shelf-life.
In a long-term study of the effects of N, P and K fertilizer levels on postharvest
respiration of apple, Letham (1969) found that fruit that had had the highest P con-
centration had the lowest respiration rate while those with the lowest P c­ oncentration

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2019 13

A. K. Thompson et al., Banana Ripening, SpringerBriefs in Food, Health,
and Nutrition, https://doi.org/10.1007/978-3-030-27739-0_2
14 2 Preharvest Effects

had the highest respiration rate. Apples grown with low P initiated the climacteric
sooner than those grown with high P, indicating that lower levels of tissue P were
sufficient to alter fruit developmental physiology. Lin and Ehret (1991) showed that
the shelf-life of greenhouse grown cucumbers could be improved by increasing the
concentrations of N, P, K, Ca, S and Mg in the nutrient solution. Cucumbers were
also grown in a greenhouse under low and high P fertilizer regimes by Knowles
et al. (2001). The respiration rate of low P fruits was 21% higher than that of high P
fruits and they began the climacteric rise about 40 h after harvest, reached a maxi-
mum at 72 h and declined to pre-climacteric levels by 90 h. The difference in respi-
ration rate between low and high P fruits was up to 57% during the climacteric. The
climacteric was different to the low P fruits and was not associated with an increase
in fruit ethylene concentration or ripening.

Organic Production

In a case study of the postharvest qualities of conventionally versus organically

grown bananas in the Dominican Republic, Caussiol and Joyce (2004) found that
significant differences (p < 0.05) in measured quality attributes between conven-
tionally and organically grown fruit were “few and marginal”. Moreover, any differ-
ences were inconsistent across harvest-times and during shelf-life. Thus, organically
and conventionally grown bananas had almost identical qualities. Sensory compari-
son confirmed that there was no flavour difference. Nyanjage et al. (2000) found
that organically grown ‘Robusta’ bananas ripened faster at 22–25 °C than non-­
organically grown bananas as measured by peel colour change, but ripe fruit had
similar TSS levels from both production systems. The peel of non-organic fruits had
higher N and lower P contents than organic fruits. In a survey of fruit quality of
Philippine bananas from non-chemical production, the problems highlighted all
related to management practices and none to the effects of organic production on
postharvest aspects (Alvindia et al. 2000).

Light and Day Length

Banana is a day neutral crop and no information could be found on the effects of
light intensity or day length and banana ripening, although direct exposure to the
sun can damage the fruit (Fig. 2.1). Work has been carried out on other fruit. For
example, Woolf et al. (2000) showed that during ripening of avocados at 20 °C, fruit
that had been exposed to direct sunlight showed a delay of 2–5 days in their ethyl-
ene peak compared with fruits that had been grown in the shade. Aborisade and
Ayibiowu (2010) studied the effects of day length on ripening of ‘Roma’ and
‘Beske’ tomatoes. They were harvested at the mature-green, breaker, turning or
pink stages and then stored at 28 °C either in alternating light, 12 h light and 12 h in
Disease 15

Fig. 2.1 Sun damage on

bananas in Eritrea

complete darkness. Ripening progressed in complete darkness more quickly than in

the alternating light and dark in fruit at the breaker stage. However, the difference
was more pronounced in ‘Roma’ than in ‘Beske’. Fruit initially at the turning stage
did not show any significant effect of photoperiod in ‘Roma’ but did by day 6 in
‘Beske’. Generally, photoperiod had more effect at the mature-green and breaker
stages than at the turning and pink stages of ripening.

Disease

Generally, if a crop has suffered an infection during development its storage or mar-
ketable life may be adversely affected. Bananas may ripen prematurely or abnor-
mally after harvest because of leaf infections by fungi during growth, which caused
stress and therefore shortened their storage life. This can be observed on the crop
before harvesting or it may only be observed as a physiological disorder posthar-
vest. Fungicide applications in the field to control both leaf spot diseases,
Mycosphaerella musicola and M. fijiensis, were shown to reduce premature ripen-
ing (Thompson and Burden 1995). In addition, yield was reduced in areas where
plants were infected with leaf spot, as it can become necessary to harvest fruit at a
lower grade (i.e. younger) in order to lessen the chances of premature ripening of
the fruit in transit (Stover 1972; Stover and Simmonds 1987). Black Sigatoka is
caused by infection with M. fijiensis, which is endemic to most banana exporting
countries. It does not infect the fruit, but infection can cause damage to leaves or
even kill them. This reduces the photosynthetic area of the plant that can lead to
reduction in yield. This leaf damage in turn causes stress to the plants and a reduc-
tion in the pre-climacteric life of fruit harvested from infected plants. Bananas har-
vested from Black Sigatoka infected plants may behave as though they were
physiologically 1–2 weeks older than those of non-infected plants of the same age.
In trials reported by Turner (1997), reducing the number of leaves from 12 to 7
16 2 Preharvest Effects

during fruit growth did not affect bunch weight, but reduced pre-climacteric life of
fruit by 6 days. With even fewer leaves there was no further reduction in pre-­
climacteric life, but bunch weight was reduced by 8%. Similar results were reported
by Ramsey et al. (1990) who found that plants with less than five of their leaves not
greatly affected by Black Sigatoka at harvest produced smaller bunches. All bunches
from plants with fewer than 4 leaves were “field-ripe”, that is suffering from Pulpa
Crema. Pulpa Crema is where bananas are ripe but the peel remains green. This is
because the bananas are initiated to ripen on the plant, but because the ambient
temperature is above 25 °C the pulp ripens but the chlorophyll in the skin is not fully
broken down. This effect occurs in ‘Cavendish’ but not in plantains (Fig. 2.2)
(Seymour et al. 1987) and many other Musa genotypes including ‘Kluai Hom
Thong’.
Since the introduction of the shipment of bananas as hands packed in fibreboard
boxes the principal postharvest disease problem has been Crown Rot. When the
export trade began bananas were transported as bunches. It was not until bunches
were cut into hands or clusters for packing into fibreboard boxes that Crown Rot
became a problem that needed treating. For local marketing in many producing
countries, a portion of the stem is retained still attached to the crown (Figs. 1.5 and
1.6), which appears to prevent Crown Rot. Crown Rot develops from the infection
of the cut surface of the crown, where it has been cut from the fruit stalk, by fungi.
If not treated these infections develop during transport and ripening in the importing
country and cause decay of the crown tissue, which may spread into the fruit pedicel
or even the fruit itself during ripening and marketing. Several different fungi have
been found to be associated with Crown Rot (Griffee and Pinegar 1974; Griffee and
Burden 1976), the most common being the banana anthracnose fungus Colletotrichum

Fig. 2.2 Changes in the pigment level of ‘Cavendish’ bananas during ripening at either 20 °C
(.....) or 35 °C (___) for 6 days. The fruit were exposed to 1000 μL L−1 before being ripened in air.
(Source: Seymour 1985)
Disease 17

musae, which often occurs in mixed infections with Fusarium semitectum and other
fungi (Knight et al. 1977). C. musae also infects other injuries on bananas, particu-
larly those on the ridges of angular fruit, causing decay which spreads into the pulp
during ripening. Anthracnose (C. musae) gets its name from the ability of the fungi
to cause latent infections of banana fruit. These infections occur at any time during
the development of the fruit in the field, and usually only develop as the fruit ripens,
causing round lesion on the peel surface that can cause rotting in the pulp during
ripening. This was a problem when bananas were shipped as bunches with pro-
longed shipping times, or when ripened at temperatures above 18 °C. It is rarely
seen in hands packed in boxes. For international trade bananas are treated posthar-
vest with a chemical fungicide (usually Thiabendazole or Imazalil) to control Crown
Rot and other fungal diseases (Fig. 2.3) although other non-fungicidal methods have
been tested including covering the cut areas of the crown with cling film (Fig. 2.4).
Other fungi associated with postharvest diseases of banana fruit include:
Botryodiplodia theobromae Stalk and Fruit Rot,
Ceratocystis paradoxa Stem End Rot,
Magnapothe grisea, Pyricularia grisea Pitting Disease,
Verticillium dahliae Cigar End Rot.
Most of these diseases are only occasionally serious where infection levels are
high and favourable conditions occur; none are as widespread as Crown Rot. Pitting
Disease, however, is a serious field problem in some production areas. It can also be
serious problem on fruit after harvest due to the development of latent infections
that are not controlled by postharvest treatments (Meredith 1963; Stover 1972). All
postharvest disease organisms are widespread in the field, growing and sporulating
on decaying banana flowers, bracts and leaves. The spores are blown by wind or

Fig. 2.3 Cascade application of a fungicide to freshly harvested ‘Valery’ in Ecuador, before being
packed for export in 1996
18 2 Preharvest Effects

Fig. 2.4 Plastic covering

for control of crown rot
disease on organic
‘Cavendish’ bananas,
applied directly after
harvest. Photograph
taken in 2013 after
importation in UK and
ripening

splashed by rain onto the fruit and can also be carried on bunches to contaminate the
packing station environment, including the washing water. Daundasekera et al.
(2008) showed that C. musae can produce ethylene in vitro on bananas. They found
that C. musae isolate CM100 was capable of producing ethylene in vitro on
methionine-­supplemented basal medium and on banana peel extracts that contained
methionine.

Water Stress

Water stress can induce increased ethylene production (Kubo et al. 1990) in bananas,
which may explain accelerated ripening in water-stressed bananas (Burdon et al.
1994). Karikari et al. (1979) reported that ‘Apem’ responded to water stress by
marked reduction in their pre-climacteric period, which is consistent with previous
reports by Sanchez-Nieva et al. (1970) and Thompson et al. (1974).

Damage

In experiments in Ghana, Ferris et al. (1993, 1995) showed that mechanical damage
promoted early ripening in 3 plantain genotypes: ‘Ubok Iba’ (True Horn), ‘Agbagba’
(False Horn) and ‘Obino I’Ewai’ (French Plantain) when harvested at either fully
mature or immature stages. The mechanical damage that caused early ripening
could be due to impact, abrasion or incision. The effect of damage on plantain ripen-
ing affected all three genotypes but the intensity of the effect varied between geno-
types. Ferris et al. (1993a) and Bugaud et al. (2014) also showed that bruise
susceptibility varied between genotypes, but as long as the impact energy was below
200 mJ, neither ‘Cavendish’ or ‘Flhorban 925’ were susceptible to bruising.
Bunch Covers 19

Bunch Covers

Banana bunch covers are plastic film tubes that are used to cover bunches shortly
after they have emerged from the top of the pseudostem (Fig. 2.5). This is to protect
the fruit from dust, bird dropping, leaf rubbing and scarring but particularly from
damage by insects particularly different types of thrips (Thrips hawaiiensis,
Chaetanaphothrips signipennis and Hercinothrips bicinctus). Sometimes bunch
covers are coated with an insecticide to improve control of thrips. Bunch covers can
also have postharvest effects. After harvest the pre-climacteric life of the covered
bunches was one or 2 days longer than the ones that had been grown with no covers.
Johns and Scott (1989) explained this effect as the fruit under the covers were less
exposed to the environment, therefore they lost less water and had a higher moisture
content, which gave the appearance of being more mature, since they would be
more rounded (see section on Harvest maturity). Choudhury et al. (1996) also
reported a longer green life for covered ‘Dwarf Cavendish’ fruits whereas Parmar
and Chundawat (1984) reported a reduction in green life using blue polyethylene
covers on ‘Basarai’ bananas. After harvest they found that the fruits grown under a
cover lost more weight after being harvested than those that had not been covered.
They attributed this to be probably as a consequence of going from a higher humid-
ity environment during growing to a lower humidity environment after harvest.
Parmar and Chundawat (1984), on the contrary, recorded lower postharvest weight
losses in fruits that had been grown under bunch covers. They also found, with
regard to quality, that the fruit that had been covered, except for those under non-­
transparent covers, had a higher TSS content than those not covered. They

Fig. 2.5 The protective

plastic sleeve is tied in a
knot at the bottom but a
gap is left to allow water to
escape and a coloured
ribbon (in this case red) is
tied on to identify the age
of the bunch
20 2 Preharvest Effects

conjectured that this was probably because the higher temperature under the cover
favoured the conversion of starch into sugars. The reduction in the content of TSS
in fruits grown under non-transparent covers might also be due to the higher mois-
ture content of these fruits. The sugar to acid ratio of the covered fruits was also
higher than those not covered fruit.

Harvest Maturity

The maturity at which bananas are harvested clearly affects their quality and ripen-
ing (see Chap. 3 section: Maturity of fruit and response to ethylene). Various ways
are used by growers to assess harvest maturity, but the most commonly used in
international trade is “ribbon tagging”. The time between flowering and fruit being
ready for harvesting may be quite constant, so in order to identify exactly when
anthesis occurred a coloured plastic ribbon is attached to the bunch directly after
anthesis, usually together with a plastic bunch cover (Fig. 2.5). The same colour is
used for 1 week and changed to another colour the following week and so on. This
means that at the harvest time the age of is bunch is precisely known. Wilson
Wijeratnum et al. (1993) showed that the ‘Ambul’ bananas grown in Sri Lanka
reached physiological maturity 8–9 weeks after the flowers had opened. Fruit
growth and development continued until the thirteenth week but changes in other
physical and chemical parameters were minimal after 11 weeks. For ‘Cavendsh’ in
Ecuador the maximum time from anthesis to harvest is usually 12 weeks and in the
Windward Islands it is 13 weeks. Harvest maturity has been shown to affect eating
quality of other fruit. For example, Blissett et al. (2019) compared ‘East Indian’
mangoes that had been allowed to ripen before and after harvesting. That is fully
ripe golden yellow and yellow-orange skin harvested 120 days after anthesis and
mature-green of a comparable size 112 days after anthesis and both harvest maturi-
ties were ripened at 27 °C. At eating ripeness, they found no significant difference
in TSS but tree ripened mangoes had lower total reducing sugar, titratable acidity,
higher total phenolics and higher DPPH (1,1-diphenyl-2-picrylhydrazyl) free radi-
cal scavenging activity than postharvest ripened of the mangoes.
The way that ribbon tagging works in practice is that the grower has options
when he needs to harvest a bunch and chooses which bunches to harvest, not only
on their age, but also their angularity and calliper grade. In a hypothetical example
(Fig. 2.6) the grower can harvest only bunches with white, green, yellow of blue
ribbons. Bunches with a white ribbon were 10 weeks old, bunches with a green rib-
bon 11 weeks old, bunches with a yellow ribbon 12 weeks old and bunches with a
blue ribbon 13 weeks old. The grower must harvest all bunches with blue ribbons (if
their calliper grade is still below 37 mm or over 46 mm they must not be packed for
export), but the grower has the option of harvesting yellow, green and white depend-
ing on their angularity and calliper grade.
The thickness of individual fingers increases as the fruit matures and this thick-
ness can be used to determine harvest maturity. It is called calliper grade and
Harvest Maturity 21

Week
Number of bunches harvested
beginning

7 January white green yellow blue

14 January brown white green yellow

21 January red brown white green

28 January purple red brown white

Fig. 2.6 Simple chart to keep record of the number of bunches harvested and their age or
maturity

Fig. 2.7 Plastic template for measuring maximum and minimum calliper grade for harvesting
bananas for export in the Windward Islands in the 1970s

­ easures the width on an individual finger on a bunch. Callipers are used to mea-
m
sure the maximum acceptable width for fruit and the minimum acceptable widths
for harvesting and various devices are supplied to growers (Fig. 2.7) to assist
harvesters.
Because there is variation in maturity between hands on the same bunch, so in
order to standardize the selection, calliper grade is measured only on a middle finger
on the second hand from the top (Fig. 2.8).
Recommendations vary but commonly fruit with a calliper grade (diameter) of
over 46 mm should not be packed for export and fruit less than 37 mm should not
be harvested before 13 weeks after anthesis. Any bunch that still has not achieved a
calliper grade of 37 mm after 13 weeks should be harvested and discarded or mar-
keted locally, as the fruit may begin to ripen during transport.
The shape of each finger in cross section changes as the fruit matures. This
change in shape is called angularity and is used, often in combination with ribbon
tagging and other methods, to determine when to harvest a bunch. Various terms are
used in producer countries to classify the angularity of banana fingers (Fig. 2.9). In
some countries the marketing agents for export companies require farmers to har-
vest only a particular grade. In the Caribbean Islands grades were classified, for
many years, from ‘light three-quarters’ to ‘round full’, which have long been used
22 2 Preharvest Effects

Fig. 2.8 Diagram showing

the second hand form the
top of the bunch that
should be selected for
measuring calliper grade

Fig. 2.9 Cross section of ‘Cavendish’ banana fingers showing the changes in shape of fruit as they
grow and relating the shape to their width in mm

as the basis for judging when to harvest bunches for export (von Loesecke 1949;
Stover and Simmonds 1987). These are based on a subjective combination of angu-
larity and thickness of individual finger.
The maturity of hands in the bunch varies slightly; those at the proximal end of
the bunch being more mature than those at the distal end, so the estimate of matu-
rity is based on the fullness of fruit of the middle hand. In Latin America, where
large-­scale production is centred on plantation units of around 400 ha, in which
uniform production conditions are achieved, maturity is assessed by the harvester
using a specially designed spring calliper to measure the thickness of a standard
finger in the bunch. This is usually taken as the middle finger of the outer whorl of
the second hand from the distal end of the bunch. In some countries, farmers har-
vest less mature bananas, even for local markets, because they have cash-flow
problems and they harvest as soon as the ripening room operator will accept the
fruit (Thompson 1981).
Harvest Maturity 23

Fig. 2.10 Showing a

banana finger that has split
postharvest, perhaps
because of a combination
of over maturity at
harvest and irrigation or
rainfall just prior to
harvesting

Ahmad et al. (2001) found very strong evidence for ‘Robusta’ that the fruit had
much better organoleptic properties the more mature they were when harvested.
However, if bananas are allowed to mature fully before harvest and harvesting is
shortly after rainfall or irrigation, the fruit can easily split during handling opera-
tions (Fig. 2.10), allowing microorganism infection and postharvest rotting
(Thompson and Burden 1995).
Chapter 3
Fruit Ripening

Generally, all fruits have been classified into either non-climacteric or climacteric
on the basis of their respiratory pattern and ethylene evolution during ripening
(Biale 1964). The ripening process is recognised as a complex irreversible series of
processes, which is initiated by a range of phytohormones primarily ethylene but
also others such as abscisic acid (ABA) (Iqbal et al. 2017). It was previously thought
that only climacteric fruit produced ethylene, but with the advent of precise analyti-
cal techniques it has been shown that all fruit and vegetables that were tested can
both produce ethylene and react to exogenous ethylene. For example, Solomos and
Biale (1975) showed that exposure to exogenous ethylene stimulated the respiration
rate of many different types of fruit and vegetables (Table 3.1).
The rapid increase in endogenous synthesis of ethylene in climacteric fruit occurs
at completion of fruit development and involves many physiological changes as
well as increased respiration rate. In non-climacteric fruit, the upsurge of respiration
rate and ethylene biosynthesis were not observed or are transitory even after exog-
enous ethylene application (Tucker 1993; Duan, et al. 2007; Kays and Paull 2004).
It was contended by Obando et al. (2007) that the classification of fruit into cli-
macteric and non-climacteric categories is an over-simplification. The simple dis-
tinction between climacteric and non-climacteric fruits has been queried because
many fruits such as guavas, melons, Japanese plums, Asian pears and peppers show
climacteric as well as non-climacteric behaviours depending on the cultivar. Studies
on genetic and inheritance patterns indicate that ethylene-dependent and ethylene-­
independent pathways coexist in both climacteric and non-climacteric fruit (Paul
et al. 2012). Zhang et al. (2009) contended that many non-climacteric fruit show a
climacteric pattern of ripening. Also, Barry and Giovannoni (2007) contended that
there was evidence that climacteric and non-climacteric fruits share some similar
pathways of ripening. For example, in plums there are non-climacteric, suppressed-­
climacteric and climacteric cultivars. Some cultivars of plums ripen very slowly
since ethylene production is suppressed and are referred to as suppressed-­climacteric

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2019 25

A. K. Thompson et al., Banana Ripening, SpringerBriefs in Food, Health,
and Nutrition, https://doi.org/10.1007/978-3-030-27739-0_3
26 3 Fruit Ripening

Table 3.1 The effects of Ethylene addition

exposure to exogenous
Crop 0 +
ethylene on respiration rate
(μL O2 g−1 h−1) during storage Apple 6 16
of various fruit and Avocado 35 150
vegetables compared to those Beet 11 22
not exposed to exogenous Carrot 12 20
ethylene. Modified from Cherimoya 35 160
Solomos and Biale (1975)
Grapefruit 11 30
Lemon 7 16
Potato 3 14
Rutabaga 9 18
Sweet potato 18 22

cultivars. They may remain under-ripe if harvested too early. Other cultivars e.g.
‘Early Golden’ ripen very rapidly after harvest. Genetic analysis showed identical
DNA profiles for the suppressed-climacteric cultivars ‘Santa Rosa’, ‘July Santa
Rosa’, ‘Late Santa Rosa’, ‘Casselman’ and ‘Roysum’ and the novel non-climacteric
‘Sweet Miriam’ (Minas et al. 2015).
ABA content of climacteric fruit is very low at immature stages, but increases
and peaks just before the onset of ripening and autocatalytic ethylene biosynthesis.
Lohani et al. (2004) suggested that ABA stimulates ripening in bananas indepen-
dently from ethylene. Exogenous ABA application was shown to enhance fruit soft-
ening, with exception to PG action. Jiang et al. (2000) suggested that ABA may act
as a coordinator during the climacteric period in bananas by enhancing their sensi-
tivity to ethylene. Ethylene and ethylene mediated responses suggest that sensitivity
towards ethylene differs in various tissues and in different developmental stages of
fruit, because of signalling interactions of ethylene with other plant hormones,
metabolites and environmental signals (Stepanova and Alonso 2005; Kendrick and
Chang 2008; Yoo, et al. 2009; Zhang et al. 2009).

Pre-Climacteric Phase

Generally, bananas are harvested at the pre-climacteric stage (colour index 1,

Table 1.1; Fig. 1.1), which is prior to the autocatalytic generation of ethylene.
During the pre-climacteric phase, the peel of the fruit is still green, the principal
component in the pulp is starch (approximately 85 % of pulp dry weight) (Cordenunsi
and Lajolo 1995) and it has a relatively low basal respiratory rate of about 10–20 mg
CO2 kg h−1 and undetectable ethylene production (Chillet et al. 2008). The low res-
piration rate of pre-climacteric bananas is concomitant with very low fructose-1,6-­
bisphosphate content (Ball et al. 1991). Sucrose content in the pre-climacteric fruit
Pre-Climacteric Phase 27

130 230

120 220
110 210

Conductance (arbitrary units)

100 200

90 190
CO2 (ml/kg/h)

80 180

70 170
60 160

50 150
40 140

30 130

20 120
0 1 2 3 4 5 6 7 8 9 10 11 12 13
Times (days)

Fig. 3.1 Changes in CO2 evolution (■) and in the conductance (+) of intact fruit during the pre-­
climacteric and climacteric of ‘Giant Cavendish’ bananas. (From Nolin 1985)

pulp increases slowly due to the very low activity of sucrose phosphate synthase
(Cordenunsi and Lajolo 1995). During the pre-climacteric period, ethylene produc-
tion of bananas was reported to be undetectable (Liu et al. 1999; Zhang et al. 2006;
Duan, et al. 2007; Larotonda et al. 2008). Liu et al. (1999) reported that in unripe
bananas, the expression of the MA-ACS1 gene is very low, which is concomitant
with the undetectable ACS activity whilst ACO activity and abundance of the
MA-ACO1 gene increase gradually during the pre-climacteric period. Thus, the very
low levels of the both ACC content and ACS activity in pre-climacteric bananas
limit the initiation of autocatalytic ethylene biosynthesis (Pathak, et al. 2003).
The duration of the pre-climacteric stage can be estimated by measuring the
grade of the fruit (see Chap. 2, Harvest maturity), but greater precision can be
obtained by measuring the minimum concentration of ethylene that can initiate the
climacteric (Nolin 1985). The end of the pre-climacteric phase can be determined
by measuring the increased softness of the pulp (New and Marriott 1974; Marriott
and New 1975; Marriott et al. 1979). Texture can be determined subjectively by
touch or objectively by the conductance of the peel and pulp and by correlating this
conductance with the rate of respiration (Nolin 1985; Marchal et al. 1988) (Fig. 3.1).
The dielectric constant and loss factor of material correlated well with moisture
content and ripeness of fruit (Soltani et al. 2011). They found that the best frequency
of sine wave, for predicting the level of ripeness in bananas was 100 kHz, which
gave a coefficient of determination (R2) of 0.94. Further factors, besides the physi-
ological state, influence the duration of the pre-climacteric, including the tempera-
ture, humidity and atmospheric composition.
28 3 Fruit Ripening

Ripening Phase

Ripening is a complex process of coordinated activation of multiple genetic and

biochemical pathways. Climacteric fruit ripening involves a transient rapid increase
in ethylene biosynthesis and respiration rate. Davies et al. (2006) suggested that
“perception of ethylene is vital not only for the initiation of ripening but also for the
continued expression of genes required for ripening”. McMurchie et al. (1972)
described two systems of ethylene biosynthesis that operate in bananas. During
maturation there is a low basal rate of ethylene biosynthesis termed System 1. This
is followed by System 2 ethylene biosynthesis, which is responsible for the auto-
catalytic climacteric rise in ethylene production. Bouzayen et al. (2010) reported
that LeACS6 and LeACS1A are expressed at the pre-climacteric stage (System 1),
while at ripening initiation, LeACS4 and LeACS1A are the most active genes and
LeACS4 continues to be express during ripening while the expression of LeACS1A
declines. The rise in ethylene production is associated with the induction of LeACS2
and the inhibition of LeACS6 and LeACS1A expression, which may be critical for
the switch from System 1 to System 2. At the onset of ripening, bananas undergo a
rapid burst in ethylene production, followed by a marked increase in respiratory rate
(Liu et al. 1999). Afterwards, the irreversibly ripening-associated mechanisms such
as the degradation of chlorophylls in fruit peel, the increase in conversion of starch
to sucrose, the reduction of peel thickness, the increase in leakage in pulp electro-
lyte and volatile production occur (Wade 1995; Cordenunsri and Lajolo; 1995;
Golding et al. 1999; Youryon and Supapvanich 2017). Unlike most climacteric fruit,
ripening in bananas is characterized by a biphasic respiratory climacteric which
proceeded by ethylene evolution (Pathak et al. 2003).

Internal Ethylene

Unlike most other climacteric fruits, a huge production of ethylene in bananas starts
at the onset of climacteric phase and then rapid falls during the rise of the respiration
rate (Karikari et al. 1979; Duan et al. 2007). Peak ethylene production in some cli-
macteric fruit was given by Belitz et al. (2009) as: avocados 500 μg L−1, bananas
40 μg L−1, pears 40 μg L−1, tomatoes 35 μg L−1 and mangoes 3 μg L−1. The change
in physiology of climacteric fruit from maturation to ripening is initiated when cel-
lular quantities of ethylene reach a threshold level (Yang and Hoffman 1984). The
metabolic pathway within the cells of higher plant that result in ethylene production
involves a series of steps culminating in the conversion of methionine to SAM
(S-adenosyl-methionine) which is converted to ACC (1-aminocyclopropane-1-­
carboxylic acid) by the action of ACC synthase. ACC is the immediate precursor of
ethylene biosynthesis in plants. ACC oxidase is required to convert ACC to ethylene
(Hamilton et al. 1990). In the pre-climacteric phase of banana development, ethyl-
ene production is very low and starts to increase at the onset of climacteric phase
(Duan et al. 2007). Previous work in bananas has shown that the induction of
Internal Ethylene 29

ethylene biosynthesis is primarily regulated by the expression of ACS and ACO

genes (Liu, et al. 1999; Zhang et al. 2006). They found that among MA-ACS genes,
MA-ACS1 was the key ripening gene in bananas playing a crucial role in ethylene
biosynthesis regulation during ripening, which is the only gene expressed during
ripening. Whereas ACO activity increased gradually during the pre-climacteric
phase and then there was an abrupt increase in ACO activity in parallel with the
burst of ethylene biosynthesis (Liu et al. 1999). Karmawan et al. (2009) identified
nine ACS gene family members in Musa AAA, called MA-ACS1–9, but only
MA-ACS1 correlated with fruit ripening. Lòpez-Gòmez et al. (1997) found the large
increase in ethylene biosynthesis in bananas during ripening was also associated
with the increased expression of MA-ACO1 gene and ACO activity. After the cli-
macteric peak, ethylene production decreased during the increase in respiration rate.
The decrease in ethylene biosynthesis, associated with a rapid reduction of ACO
activity, was limited through the decline of its cofactors (ascorbate and iron) or
other unknown factors (Liu et al. 1999). Xuejun Liu et al. (1999) ripened ‘Grand
Nain’ bananas at 22 °C naturally or after treatment with 100 μL L−1 ethylene for
18 hours. They also found that ethylene synthesis was regulated by transcription of
MA-ACS1, but only until the climacteric rise in respiration and may subsequently be
limited by accessibility of cofactors in situ for example ascorbate and iron. They
showed that the typical pattern of ethylene biosynthesis when bananas are allowed
to ripen without the application of exogenous ethylene (Fig. 3.2) was changed when
ripening was initiated with exogenous ethylene. (Fig. 3.3). Cyanide is produced
along with ethylene, but does not accumulate in the oxidative breakdown of ACC
that were catalysed by the ethylene forming enzymes (Blackbourn et al. 1990;

Fig. 3.2 Changes in a

ethylene biosynthesis and 0.5
1-aminocyclopropane
0.4
Ethylene(nmol g-1 h-1)

1-carboxylic acid
production in bananas
0.3
ripened naturally. Modified
from Xuejun Liu et al.
(1999) 0.2

0.1

0
b
3
ACC(nmol g-1)

0
0 2 4 6 8 10 12 14 16
Ripening days
30 3 Fruit Ripening

Fig. 3.3 Changes in

ethylene biosynthesis and
1-aminocyclopropane
1-carboxylic acid
production in bananas
initiated to ripen with
exogenous ethylene.
Modified from Xuejun Liu
et al. (1999)

1990a). Ke and Tsai (1988) found that during ripening the ACC concentration in the
pulp was higher and the ethylene-forming enzyme activity lower compared with the
peel. Ethylene production was mainly from the pulp. In the absence of the pulp, the
peel did not degreen fully unless supplied with exogenous ethylene. They concluded
that degreening of the peel relied on ethylene diffusing from the pulp to the peel.
Pathak et al. (2003) showed that bananas have a “unique biphasic respiratory
climacteric”. Working with ‘Dwarf Cavendish’ they found that ethylene evolution
on day 1, after the application of exogenous ethylene to pre-climacteric bananas,
there was a 8.3 times increment in ethylene evolution followed on day 2 by a ten
fold increase in respiration rate and on day 4 there was a 6.9 fold increase in ethyl-
ene evolution followed by a six fold increase in respiration rate on day 6. They sug-
gested that this second peak justifies the appearance of a new ethylene binding site
and the synthesis of an ethylene receptor in banana pulp. The second ethylene pro-
duction in bananas is concomitant with the breakdown of starch resulting in the
accumulation of sugars and flesh softening in the post-climacteric peak stage (Yang
and Ho 1958). The starch breakdown during banana ripening is associated with the
increase in sucrose phosphate synthase activity and the decline of sucrose synthase
activity (Cordenunsi and Lajolo 1995). The sucrose phosphate synthase activity and
sucrose synthesis in bananas is strongly stimulated by ethylene (Hubbard et al.
1990). During the post-ripening initiation phase, water content and soluble pectin of
banana pulp increased gradually (Yang and Ho 1958), which is consistent with soft-
ening being associated with cell wall hydrolases. Ethylene evolution during banana
ripening plays a key role in up-regulating PME, PG, pectate lyase and cellulase
(Lohani et al. 2004).
Effects of Ethylene Post Ripening Initiation 31

Effects of Ethylene Post Ripening Initiation

Exposure to exogenous ethylene has been shown to hasten the ripening of mangoes,
even after they had been initiated to ripen (Burg and Burg 1962), but no reports of
similar effects could be found for bananas. One supermarket chain in Thailand used
ethylene absorbing sachets in their MAP bananas in an attempt to increase their
shelf-life, both whilst being offered for sale on their shelves and for customers at
home. The temperature is relatively high in both places and even under air-­
conditioning the temperature can be about 25 °C. It was reported that this treatment
was unsuccessful in extending the shelf-life of the bananas and it was not used in
commercial practice by that company (Patcharin Chitaurjaisuk personnel communi-
cation 2019). It was suggested that this was not surprising since these bananas
would have been initiated to ripening by ethylene treatment before they were packed
and therefore absorbing any ethylene in the pack that was produced by the bananas
would have no effect on speed of ripening. This hypothesis is supported by Golding
et al. (1998), working with ‘Williams’, who found that their respiration rates, peel
colour and total volatiles production, “once engaged with autocatalytic ethylene
production, become partially independent of further ethylene action”.
However, slowing the ripening of bananas that had been initiated to ripen can be
achieved by 1-MCP treatment or changing the atmosphere around the fruit. Bananas
treated with 1-MCP, after they had been initiated to ripen, had an extended shelf-life
of 4–6 days compared to those not treated (Anonymous 2019) (See also Chap. 4
1-Methylcyclopropene). Liu (1976a) showed that bananas that had been initiated to
ripen by exposure to exogenous ethylene and then immediately stored in 1 kPa O2
at 14 oC remained firm and green for 28 days but then ripened almost immediately
when transferred to air at 21 °C. Ahmad and Thompson (2006) showed that the
marketable life of ‘Giant Cavendish’ could be extended 2.3–3.8 times, depending
on the combination of O2 + CO2 used, compared to storage in air. However, they
found that there were detrimental effects on fruit quality when 2 kPa O2 was used
and overall 4 kPa O2 was most effective in extending their storage life. CO2, in the
range tested (4–8 kPa CO2) appeared to have no positive or negative effects on mar-
ketable shelf-life or fruit quality. Klieber et al. (2002) also found that exposure of
‘Williams’ to O2 below 1 kPa at 22 °C after ripening initiation induced serious skin
injury that increased in severity with increasing exposure over the times tested
(6–24 h) and also did not extend their shelf-life. ‘Sucrier’ that had been initiated to
ripen with ethylene and then placed in polyethylene film bags (0.03 mm) at 20 °C
showed inhibited ripening and had a “fermentation flavour” (Romphophak et al.
2004). Storage of ‘Williams’, which had been initiated to ripen, in total N2 at 22 °C
had a similar shelf-life to those stored in air, but areas of brown discolouration
appeared on the peel of bananas in total N2 storage (Klieber et al. 2002). Hypobaric
storage can slow ripening of bananas after they have been initiated to ripen. Liu
(1976c) initiated ‘Dwarf Cavendish’ to ripen by exposure to 10 μL L−1 ethylene at
21 °C. They then stored them at 14 °C for 28 days under hypobaric storage of 51 or
79 mm Hg. All fruit remained green and firm for the 28 days and continued to ripen
32 3 Fruit Ripening

normally after they had been removed to atmospheric pressure at 21 °C (760 mm

Hg). See also Chap. 4 Hypobaric storage.
Controlled atmosphere storage of bananas before ripening initiation has been
shown to affect ripening. Wills et al. (1998) showed that pre-climacteric bananas
exposed to low O2 took longer to ripen when subsequently exposed to air than fruits
kept in air for the whole period. ‘Cavendish’ that had been stored for 7 days in 0.5
or 2 kPa O2 then transferring to air and initiated to ripen with ethylene, showed a
delay in the onset of the climacteric rise in respiration rate and delays in yellowing
of peel, softening and conversion of starch to sugars compared to the control that
had been stored in air throughout (Imahori et al. 2013).

Genetic Effects on Ripening

The elements of the postharvest environment that have been shown to interact with
genetic factors in fruit include temperature, ethylene, O2 and CO2. Ripening, tex-
tural and colour changes and susceptibility to diseases have been shown to be influ-
enced by gene expression (Giovannoni 2001). Banana genes associated with both
peel and pulp ripening changes have were described by Elitzur et al. (2010) and
Manrique-Trujillo et al. (2007) who showed that differential gene expression during
ripening in bananas was linked with changes in both metabolism and components.
Seymour et al. (2011) reported that in strawberries, a MADS-box SEPALLA total
acidity gene (SEP1/2) was needed for normal fruit development and ripening and in
bananas the MADS-box SEP3 gene also displayed ripening related expression
(Elitzur et al. 2010). They characterized two banana E class (SEPALLATA3) MADS-­
box genes, MaMADS1 and MaMADS2, which were homologous to the tomato RIN-­
MADS ripening gene (Seymour et al. 2013). The delay in fruit ripening in bananas
was shown to be associated with reduced biosynthesis of ethylene, and in the most
severe repressed lines, no ethylene was produced and ripening was most delayed in
those lines. However, unlike tomato rin mutants, bananas of all transgenic repres-
sion lines responded to exogenous ethylene and ripening normally (Elitzur et al.
2016).
A number of genes involved in ethylene signalling have been identified mainly
from the isolation of ethylene response mutants in Arabidopsis, defining a pathway
from ethylene perception to changes in gene expression. Johnson and Ecker (1998)
found a negative regulator of ethylene responses, CTR1, acting downstream of the
ethylene receptors. This pathway suggests that the ethylene signal is propagated
through a mitogen-activated protein kinase cascade. Yan-chao Han et al. (2016)
showed a significant induction of MaC2H2–1/2, C2H2 zinc finger proteins, tran-
scripts during the ripening of bananas with three different ripening characteristics
caused by natural, ethylene-induced and 1-MCP delayed treatments, which corre-
lated well with ethylene production. Also, MaC2H2–1/2 bound to the promoters of
the key ethylene biosynthetic genes MaACS1 and MaACO1 and repressed their
activities. They concluded that MaC2H2–1/2 are transcriptional repressors and may
Genetic Effects on Ripening 33

mediate a finely tuned regulation of ethylene production during banana ripening,

possibly via transcriptional repression of ethylene biosynthetic genes. Jian-fei
Kuang et al. (2013) reported that EIN3 binding F-box protein (EBF) is an essential
signalling component necessary for ethylene response. They isolated two EBF
genes designated MaEBF1 and MaEBF2 from bananas and found that MaEBF2
was enhanced by ethylene during fruit ripening, while MaEBF1 changed only
slightly. They concluded that MaEBF may be involved in banana ripening, at least
partly via interaction with MaEIL5. In the nucleus, the EIN3 family of DNA-binding
proteins regulates transcription in response to ethylene and an immediate target of
EIN3 is a DNA-binding protein of the AP2/EREBP family (Alonso et al. 1999).
Both elevated CO2 and reduced O2 atmospheres have beneficial effects in con-
trolling overall metabolic rate and ethylene production during the postharvest period
of fresh fruit and vegetables (Wills et al. 2007). Song et al. (2015) suggested that
high levels of CO2 in stores can compete with ethylene for binding sites in fruits.
Burg and Burg (1967) showed that in the presence of 10 kPa CO2 the biological
activity of 1 % ethylene in the surrounding atmosphere was abolished. Yang (1981)
showed that CO2 accumulation in the intercellular spaces of fruit acts as an ethylene
antagonist. Romero et al. (2008) stored grapes under 20 kPa CO2 + 20 kPa
O2 + 60 kPa N2 or air for 3 days, then air for 15 days. The grapes that had been
exposed to controlled atmosphere for 3 days activated the induction of transcription
factors belonging to different families such as ethylene response factors, in particu-
lar VviERF2-c. Rothan et al. (1997) found that CO2 induced the expression of cer-
tain stress-related genes and blocked the expression of both ethylene-dependent and
ethylene-independent ripening associated genes. Song et al. (2015) stored bananas
and plantains in high CO2 concentration (20 kPa) with O2 at 21 kPa for 6 days at
24 °C. They found that storage in high CO2 atmospheres suppressed ethylene evolu-
tion and the expression of the related biosynthesis gene, ACS, as well as up-­regulated
the expression of anaerobic respiration pathway genes, ADH and PDC, which might
be responsible for the retardation of the pulp ripening.
In climacteric fruits the final step in the biosynthesis of ethylene requires O2
(Adams and Yang 1979), therefore, a lower O2 concentration might result in an inhi-
bition of ethylene biosynthesis and slow the ripening process. Also, all fresh fruits
and vegetables synthesize O2, many in very low levels, so the effects on ethylene
biosynthesis may also be a factor. Loulakakis et al. (2006) concluded from their work
on low O2 storage of avocados that the “low oxygen, in addition to its inhibitory
effect on ethylene biosynthesis and action, exerts its effect on prolonging the storage
life of fruit by inducing a number of genes and proteins which possibly participate in
the adaptation of fruit to low oxygen and by suppressing or by un-­affecting others
without the involvement of ethylene”. Tonutti (2015) contended that members of the
ethylene responsive factor VII (transcription factors gene family) displayed differen-
tial expression suggesting their involvement in the ­modulation or controlling mecha-
nisms where there is O2 deficiency in apple cells. Imahori et al. (2013) showed that
bananas stored in an atmosphere containing 0.5 kPa O2 there was increased alcohol
dehydrogenase activity and after the bananas were transferred to regular (21 kPa O2)
air and initiated to ripen with ethylene, the onset of climacteric peak, peel yellowing
34 3 Fruit Ripening

and pulp softening were delayed. Kanellis et al. (1989a; 1998b) found that storing
avocados, which had been initiated to ripen, for 6 days in an atmosphere containing
2.5 kPa O2 suppressed cellulase activity immune-reactive protein and abundance of
its mRNA and also produced an alteration in the profile of total proteins, which
involved suppression, enhancement and induction of new polypeptides (Kanellis
et al. 1989a and 1993). In addition, it has been shown that O2 levels of 2.5–5.5 kPa,
which suppressed the appearance of ripening enzymes at the protein and mRNA
levels, was similar to those O2 levels that induced the synthesis of new isoenzymes of
alcohol dehydrogenase (Kanellis et al. 1993).
Pre-storage treatments can also affect the genetic response of fruit to environ-
mental conditions. For example, Kangliang Sheng et al. (2018) found that several
key genes involved in phenylpropanoid, flavonoid and stilbenoid pathways, includ-
ing PAL, CHS, F3H, LAR, ANS, STS, showed increased expression in response to
UV-C treatment in grapes. Storage temperature, prior to ripening, can influence the
organoleptic characteristics of ripe bananas through genetic expression. Xiaoyang
Zhu et al. (2018) stored bananas at 5 °C, 13 °C or 20 °C and found that both 13 °C
and 5 °C affected volatile-related amino acid and biosynthetic precursors of fatty
acid compositions, although at 13 °C the production of only a few specific volatiles
were reduced compared to 20 °C. The expression levels of the biosynthesis-related
genes associated with volatiles, MaHPL, MaLOX, and MaAAT, were repressed at
both 13 °C and 5 °C particularly 5 °C, while MaADH and MaPDC were up-­
regulated. They concluded that storage at 5 °C reduced volatile production by regu-
lating different key enzymes and genes involved in volatiles biosynthetic pathways
and were mainly mediated via the repression of the lipoxygenase and amino acid
metabolic pathways.

Maturity of Fruit and Response to Ethylene

The three main factors affecting the response of bananas to exogenous ethylene are
harvest maturity, the period from harvest until ethylene exposure began and the dura-
tion of exposure to ethylene. Liu (1976a, b and c) found that immediately after har-
vest, bananas do not respond to exogenous ethylene, even when exposed at high
levels They reported that this effect could be due to the presence of inhibitors, which
could explain the absence of ripening of bananas before harvest. Liu (1976b) found
that bananas that are too young do not have the enzymatic systems required to syn-
thesize ethylene. Basically, very low concentrations of ethylene (10–50 μL L−1) are
enough to initiate ripening of bananas, but, in commercial practice they may be
exposed to 1000 μL L−1 of ethylene for 24 and sometimes 48 hours to ensure uniform
ripening (Thompson and Burden 1995). For the same maturity the climacteric rise in
respiration was initiated by increasing the concentration of ethylene until a threshold
value of about 0.8 μL L−1 was reached, beyond which the response remained
unchanged (Chang and Hwang 1990a, b and c). These effects depend on genotype,
temperature and on the conditions in which they were grown. Inaba and Nakamura
Maturity of Fruit and Response to Ethylene 35

(1986, 1988) have shown for ‘Cavendish’ that the minimum time of treatment with
ethylene is a function of the concentration of ethylene supplied, the temperature of
the fruit and the concentration of ACC just before treatment. Jedermann et al. (2015)
found that ripening of ‘Cavendish’ from 14 week old bunches (weeks after flower
emergence) did not differ from fruits of 15 week old bunches. They also found no
ripening differences of the younger fruits from the bottom of the bunch (hand 7 and
8) compared to the older fruit at the middle (hand 4 and 5) or the top of the bunch
(hand 1 and 2). In contrast, Ahmad et al. (2001) showed differences in ripening for
‘Cavendish’ from hands from different parts of the bunch and bunches of different
ages but, they estimated that the ripening differences of the bunch position could not
be perceived in a commercial ripening situation. They found that the time taken for
fingers to be fully ripe after harvest was significantly (p = 0.05) affected by their age
at harvest and also its position on the bunch (Table 3.2). Overall it can be interpreted
that the older the individual finger at harvest the shorter the time needed for it to
ripen. This was especially clear for fruit harvested after 10 weeks, but there was little
difference between 12 and 14 weeks after anthesis.
The colour of the fingers, as measured with a colourimeter, at colour stage 6
(Table 3.3) varied significantly (p = 0.05), with fingers having less green and more
yellow with increased harvest maturity and a similar trend from the lower to the
upper position of the finger on the bunch (Table 3.3). It can therefore be inferred that
the older the fruit at harvest the better its colour when it is fully ripe. Also, it illus-
trates that human perception of colour can be considerably different to instrumental
measurement. However, Mustaffa et al. (1998) for ‘Montel’ bananas found that
there were differences in size, weight, volume, peel colour, texture, TSS, AA, pH,
TA, starch and sugar contents between different hands and different fingers from the
same bunch. Ahmad et al. (2001) found that the texture of ‘Cavendish’ was firmer
for fruits ripened at 13 °C than at 16 °C especially for the younger fingers (Table 3.4).
This was also reflected in TSS which was lower in the fingers ripened at 13 °C than
at 16 °C. Taken with the data in Table 3.4 and Table 3.5 it can be concluded that the
fruit ripened at 13 °C were less ripe that those ripened at 16 °C in spite of the subjec-
tive assessment that they looked about the same colour. Also, TSS, firmness and
instrumental measurement of colour reflect the stage of ripeness and there were no
differences between the two temperatures. Subsequently Ahmad et al. (2007), work-
ing with ‘Cavendish’ bananas found that they ripened in 9 days at 18 °C and in

Table 3.2 Effects of harvest maturity and position of the hand in the bunch on the number of days
to ripen to colour stage 6 (Table 1.1) at 13 °C after initiation to ripen with ethylene of ‘Robusta’
and ‘Grand Naine’. Modified from Ahmad et al. (2001)
‘Robusta’ ‘Grand Naine’
Position of hand on bunch
Time after anthesis Top Middle Lower Top Middle Lower
14 weeks 18 19 21 24 24 30
12 weeks 19 20 25 24 24 32
10 weeks 28 23 34 31 32 35
36 3 Fruit Ripening

Table 3.3 Effects of harvest maturity and position of the hand on the bunch colour at colour stage
6 (Table 1.1) at 13 °C after initiation to ripen with ethylene. Modified from Ahmad et al. (2001)
Positive b* = increased
Negative a* = increased greenness yellowness
Position of hand on bunch
Time after anthesis Top Middle Lower Top Middle Lower
14 weeks −3.7 −4.5 −4.7 49 51 49
12 weeks −4.3 −4.6 −5.0 48 49 48
10 weeks −4.9 −4.9 −5.1 48 49 47

Table 3.4 Effects of harvest maturity and position of the hand on the bunch on the texture and
TSS at colour stage 6 (Table 1.1) after initiation to ripen with ethylene and storage at either
16 or 13 oC of ‘Grand Naine’. Modified from Ahmad et al. (2001)
16 °C 13 °C
Position of hand on bunch
Texture
Time after anthesis Top Middle Lower Top Middle Lower
14 weeks 3.5 3.5 3.3 5.1 5.2 5.6
12 weeks 3.4 3.4 3.2 5.6 5.6 5.8
10 weeks 3.3 3.2 3.0 5.7 5.7 8.2
TSS
Time after anthesis Top Middle Lower Top Middle Lower
14 weeks 21.4 21.0 19.1 19.1 18.7 16.1
12 weeks 19.7 19.2 18.2 18.5 17.6 16.7
10 weeks 18.8 18.2 16.8 16.7 16.6 14.5

Table 3.5 Effects of harvest maturity and position of the hand on the bunch on taste panel scores
(1 = low and 5 = high) at colour stage 6 (Table 1.1) after initiation to ripen with ethylene and
storage of ‘Grand Naine’ at 16°C. Modified from Ahmad et al. (2001)
Flavour Sweetness Acceptance
Position of hand on bunch
Time after anthesis Top Middle Lower Top Middle Lower Top Middle Lower
14 weeks 4.3 4.1 3.1 4.6 4.2 3.5 4.4 4.0 3.0
12 weeks 3.9 3.8 3.0 3.5 3.5 2.8 4.0 3.8 2.7
10 weeks 3.1 3.0 2.6 3.3 3.1 2.7 3.4 3.5 2.8

11 days at 16 °C, and there was a slight preference for flavour and acceptability for
those ripened at 18 °C.
There were no significant differences (p = 0.05) in the taste panel assessments for
ripe ‘Grand Naine’ ripened at either 13 °C (data not shown) or 16 °C (Table 3.5)
although there were clear trends that the panellists preferred the more mature fruit
in all the criteria used.
Changes that Occur During Ripening 37

Factors Affecting Fruit and Response to Ethylene

Pathak et al. (2003) commented that “ethylene induced ripening of banana is char-
acteristically different from that of other climacteric fruits and that ethylene biosyn-
thesis may have more than one mechanism operating during ripening which are
tightly controlled at various levels.” Bananas have been shown to respond to ethyl-
ene concentrations as low as 0.015–0.05 μL L−1 at 21 °C (Liu 1976a), although
concentrations greater than 0.001 μL L−1 were shown to be required for the initia-
tion of the climacteric in avocadoes (Biale and Young 1971). The production of
endogenous ethylene by different Musa genotypes placed in the same conditions
varies. For example, Karikari et al. (1979) showed that under conditions where
‘Cavendish’ (Musa AAA) produced 2.1–2.5 μL−1 kg hr.−1 ethylene, tetraploids
(Musa AAAA) produced 6.7 μL−1 kg hr.−1 ethylene and ‘Apem’ (Musa AAB) pro-
duced 34.8 μL−1 kg hr.−1 ethylene. The internal factors regulating the response to
ethylene may be associated with growth regulators (Brady 1987) and ethylene
receptors could be involved, thus explaining the inhibition of ripening by silver,
which is not overcome by exogenous ethylene as the silver is bound to the putative
ethylene receptor (Saltveit et al. 1978). Ethylene-mediated regulation of starch-­
degrading enzymes at transcriptional and translational levels is crucial for sugar
metabolism during banana ripening. Also, the crosstalk between ethylene and other
growth substances, including IAA and abscisic acid, also influences primary sugar
metabolism. The starch stored in banana pulp cells is compartmentalized within
plastids, which also contain several common starch-degrading enzymes (Peroni-­
Okita et al. 2013; Junior et al. 2006; Purgatto et al. 2001).

Changes that Occur During Ripening

A general figure for the various chemicals in banana pulp were given by USDA
(Table 3.6). Forster et al. (2003) showed that the chemical composition in banana
pulp varied with the central part being higher in nutrients, except for ascorbic acid.

Peel Colour

The pigments in the peel of bananas and plantains are chlorophylls and carotenoids
(Figs. 3.12; 3.13; 3.14). The change in colour of ripening fruits is associated with
the breakdown of chlorophylls with carotenoid levels remaining relatively constant
(Von Loesecke 1949; Seymour 1986; Montenegro 1988) (Fig. 2.2), however, this
depends on the genotype. ‘Cavendish’ banana cultivars can fail to completely
degreen when they are ripened at 24–25 °C and above (Seymour et al. 1987; Semple
and Thompson 1988). This can result in bananas, which are ripe in every other
38 3 Fruit Ripening

Table 3.6 The composition Component Content

of banana fruit for 100 g−1
Water 74.91 g
fresh weight (adapted from
USDA 2012) Energy 89 kcal
Protein 1.09 g
Total lipid (fat) 0.33 g
Carbohydrate, by difference 22.84 g
Fibre, total dietary 2.6 g
Total sugars 12.23 g
Calcium 5 mg
Iron 0.26 mg
Magnesium 27 mg
Phosphorus 22 mg
Potassium 358 mg
Sodium 1 mg
Zinc 0.15 mg
Total ascorbic acid 8.7 mg
Thiamin 0.031 mg
Riboflavin 0.073 mg
Niacin 0.665 mg
Vitamin B-6 0.367 mg
Folate, DFE 20 mcg_DFE
Vitamin B-12 0.00 μg
Vitamin A, RAE 3 mcg_RAE
Vitamin A, IU 64 IU
Vitamin E (alpha-tocopherol) 0.10 mg
Vitamin D (D2 + D3) 0.0 μg
Vitamin D 0 IU
Vitamin K (phylloquinone) 0.5 μg
Fatty acids, total saturated 0.112 g
Fatty acids, total monounsaturated 0.032 g
Fatty acids, total polyunsaturated 0.073 g

respect, remaining green. The higher the temperature the more obvious the effect. It
is one cause of the physiological disorder of ‘Cavendish’ bananas called “pulpa
crema” or “yellow pulp.” This is where bananas are initiated to ripen on the plant,
but because the ambient temperature is above 25 °C the pulp ripens but the chloro-
phyll in the skin is not fully broken down. Bowden et al. (1994) compared the peel
colour of ‘Cavendish’ and ‘Apple’ bananas by measuring their a* values in a Colour
Difference Meter (the lower the a* value the greener the peel). They found that their
pre-climacteric readings were both just above −19, but after 28 days storage at
13 °C ‘Cavendish’ had gone to +2.2 and ‘Apple’ to −6.1. When they were subse-
quently initiated to ripen with ethylene at 19 °C, the a* values were + 2.7 for
Changes that Occur During Ripening 39

‘Cavendish’ and + 1.7 for ‘Apple’. With plantains it was shown that complete chlo-
rophyll destruction can occur even at 35 °C (Seymour 1985, Seymour et al. 1987).
Studies on why this effect of temperature on degreening of ‘Cavendish’ bananas
occurs failed to reach a definitive conclusion (Blackbourn et al. 1990), although
there was some indication that it was related to the thylakoid ultrastructure of the
chloroplasts (Seymour 1986, Blackbourn et al. 1990). Thylakoids are an internal
system of interconnected membranes that are arranged into stacked (grana thyla-
koids) and non-stacked (stroma thylakoids) regions that are differentially enriched
in photosystem I and II complexes. Blackbourn et al. 1990) showed that during
ripening of ‘Cavendish’ at higher temperatures there was a retention of thylakoid
membranes resulting in a relatively delayed breakdown in both chlorophyll b and
chlorophyll a from the reduced dismantling of pigment-protein complexes. By con-
trast, degreening was complete within 4 days at both 20 °C and 35 °C in plantains
(Musa AAB), where the thylakoid membranes and their associated pigment-protein
complexes were lost, and there was rapid increase in chlorophyll a to b ratios at both
ripening temperatures. They suggested that the retention of thylakoid membranes is
an important factor in the failure of ‘Cavendish’ bananas to degreen when ripened
at tropical temperatures, and that the degreening problem may be related to the
comparatively high chlorophyll b content of the pre-climacteric fruit. Sultan and
Rangaraju (2014) stored ‘Grand Naine’ at 20 °C and 95 % RH with 100 ppm ethyl-
ene for 24 h. Using the Hunterlab and CIE L*a*b* systems they found that lightness
(L*) and yellowness (b*) increased to 74.25 and 51.11 respectively by the 4th day
and subsequently declined.
Colour changes during ripening of many fruits, including bananas, have been
used as a rough guide to the stage of ripeness. It is commonly used commercially in
the form of colour matching charts based on the degree of peel yellowing where
colour index 1 (Table 1.1) is allocated to fruit that are dark green and colour index
6 (Table 1.1) is for fruit which are fully yellow (Von Loesecke 1949; Lizada et al.
1990; Thompson 1996). Examples of these colour charts are supplied by commer-
cial banana companies and can also be found in Stover and Simmonds (1987),
Abdullah and Pantastico (1990) and Thompson (1996).
Medlicott et al. (1992) measured changes in peel ground colour during ripening
of bananas using peel colour scores, a colour difference meter and by extraction and
measurement of chlorophyll and carotenoid concentrations. There were close cor-
relations between the subjective colour scores and the colour meter measurements.
A similar relationship was obtained for colour scores and chlorophyll content, but
no significant correlations were found between colour scores and carotenoid c­ ontent.
Peel colour changes in relation to pulp firmness and TSS content were highly cor-
related and preceded simultaneously throughout ripening. Slaughter and Thompson
(1997.) described an optical chlorophyll sensing system that showed a high correla-
tion with spectral analysis, tristimulus colorimeter analysis and visual colour match-
ing of the peel colour of bananas. This optical chlorophyll sensing system gave a
rapid response, was simple to operate, non-destructive and low cost and was reported
to have potential application in automatic monitoring of banana ripening.
40 3 Fruit Ripening

Peel Spotting

Brown spots on the skins of bananas is a normal stage of ripening that usually
occurs when the skin has turned from green to yellow or fully yellow depending on
the genotype. Maneenuam et al. (2007) compared the effect of storage in different
O2 partial pressures at 25 °C and 90 % RH on peel spotting in ‘Sucrier’. The fruit
had first been initiated to ripen and were turning yellow. They were then transferred
to atmospheres containing either 90 kPa O2 or 18 kPa O2 in gas tight chambers. Peel
spotting and a decrease in dopamine levels were quicker in fruit in 90 kPa O2 indi-
cating that dopamine might be a substrate for the browning reaction. Dopamine
levels in ‘Cavendish’ were between 80–560 mg 100 g−1 in the peel and 2.5–10 mg
100 g−1 in the pulp, even in ripened bananas (Kanazawa and Sakakibara 2000). The
browning reaction is related to the activity of PAL that converts phenylalanine to
free phenolic substances that form the substrate that is converted to quinones by
PPO. The level of total free phenolics, both in the whole peel and in peel spots, was
lower in the high O2. They concluded that peel spotting was not correlated with in
vitro PAL and PPO activities. Decreases in dopamine levels correlated with peel
spotting, indicating that it might be used as a substrate for the browning reaction.
Kamdee et al. (2018) ripened ‘Sucrier’ to colour index 3–4 (Table 1.1) and then
placed them at 42 °C and 78 % RH for 6, 12 18 or 24 hours followed by storage at
25 °C and approximately 78 % RH. Exposure to 42 °C inhibited peel spotting but
did not impair ripening, with exposure for 24 hours being the most effective, but the
taste and odour of the ripe fruit was reduced to unacceptable levels. After the
18-hour treatment peel spotting was considerably reduced whilst the taste and odour
remained acceptable. Total phenolics and dopamine content of treated bananas were
higher and the in vitro activity of enzymes involved in the production of substrate
(PAL) and in the initial browning reactions PPO was inhibited by exposure to
42 °C. The exposure to 42 °C apparently did not delay the membrane degradation
required for the interaction between the substrate and the enzymes that catalyze the
browning reactions, suggesting that the main effect of the heat treatment was in the
enzymatic steps involved in the browning reactions.

Peel Chemicals

Changes in many chemicals that occur in banana peel during ripening have been
dealt with above. Banana peel also contains substantial amounts of phytonutrients
and minerals. Puraikalan (2018) found that powder made from banana peel con-
tained total phenols (159–235 mg 100 g−1), flavonoids (1.23–1.70 mg QE g−1), tan-
nin (127–152 mg 100 g−1), protein (9.4–11.7 %), fat (3.6–6.7 %) and
fibre(11.5–14.4 %), depending on variety, and could be incorporated into processed
foods and serve as a functional food. Anhwange et al. (2009) analysed M. sapientum
peel and gave the mineral content as potassium, calcium, sodium, iron, manganese,
bromine, rubidium, strontium, zirconium and niobium to be 78.10, 19.20, 24.30,
Changes that Occur During Ripening 41

0.61, 76.20, 0.04, 0.21, 0.03, 0.02 and 0.02 mg g−1, respectively. The concentrations
of protein, crude lipid, carbohydrate and crude fibre were: 0.9, 1.7, 59.0 and 31.7 %,
respectively. They also commented that banana peel is a good source of nutrition
and could be utilized as an animal feed supplement. Baskar et al. (2011) tested etha-
nolic extract of peel of free radical scavenging assays of peel extracts of nine local
banana genotypes. They found that peel extracts of all the nine genotypes had sig-
nificant antioxidant and phytochemical activity.

Finger Drop

Finger drop occurs during ripening and is associated with weakening of the pedicel,
which can result in reducing the market value of the fruit. Sensitivity to finger drop
was reported to vary between banana varieties (New and Marriott 1983). Esguerra
et al. (2009) stored ‘Latundan’ bananas at 23–25 °C and found that finger drop
occurred about 5–6 days after harvest when the fruit were fully yellow. A major
cause of finger drop is infection, associated with crown rot fungi (see Chap. 2
Stress), but it has also been associated with physiological effects and was shown to
be stimulated by ripening at higher temperatures (Semple and Thompson 1988).
They also found that prolonged exposure of fruit to ethylene during ripening could
increase finger drop in certain circumstances. Paull (1996) found that finger drop in
‘Santa Catarina Prata’ bananas occurred after the climacteric ethylene peak and was
associated with a second ethylene production. Holding fruit at 15 °C for up to two
weeks reduced finger drop of fruit ripened at 25 °C. Ripening fruit at 17.5 °C, or
1 day at 25 °C then at 17.5 °C reduced finger drop from up to 100 % to less than
10 %. Ethylene treatment for 1 day at 25 °C resulted in less finger drop than fruit
that did not receive exogenous ethylene. More mature hands were more prone to
finger drop than less mature hands. Finger drop was associated with enhanced eth-
ylene biosynthesis gene expression including developmental related and ripening
induced genes (MaACO1), specific ripening-induced genes (MaACS1) and wound-­
induced genes (MaACS2) (Mbéguié-A-Mbéguié 2008; Huber and Mbéguié-A-­
Mbéguié 2012). In studies of ‘Cavendish’ it was shown that a change in the
expression of major cell wall modifying genes occurred in the finger drop area
(Mbéguié-A-Mbéguié 2009). Spraying the bananas postharvest with 4% calcium
chloride delayed the onset of finger drop by 2–4 days after the attainment of full
yellow colour but treatment with GA3 or ethanol did not control finger drop
(Esguerra et al. 2009).

Weight Loss

Postharvest loss of moisture from the fruit can hasten ripening. It was found by
Finger et al. (1995) that respiration rate and the biosynthesis of ethylene was
higher when the fruits had fresh weight losses of 5 % or more. The increase in
42 3 Fruit Ripening

respiration rate was 70 % higher and ethylene production was 50 % higher for
fruits that lost 5 % when compared to the control fruits in bananas which had lost
little moisture. The results confirmed that water stress (after the harvest of fruit)
might affect the shelf-life and ripening, depending on the intensity of the water
stress.
Clearly the higher the humidity during ripening the lower will be the weight
loss (Thompson et al. 1972). Thompson and Silvis (1975) found that ‘Dwarf
Cavendish’ in the Sudan could lose as much as 27 % in weight in 0% RH compared
to about 1% in 100 % RH in ambient temperatures (24-32 °C). Also packing fruit
in plastic film or moist coir will reduce their weight loss. Generally, the more
mature the bananas are at harvest the lower their weight loss during ripening with
fully mature fruit loosing 5.2 %, less mature 6.3 % and for immature 10.4 % during
28 days at 16 °C (Ahmad et al. 2001). This trend was also true for plantains
(Thompson et al. 1972) and bananas where weight loss varied in fruit from differ-
ent positions on a single bunch (Table 3.5), but this could be confounded with
speed of ripening since the fruit from the top of the bunch generally ripened quicker
than those lower down. The temperature at which the fruit were ripened also
affected their weight loss, with those ripened at the lower temperature (13 °C) los-
ing more weight than those ripened at a higher temperature (16 °C) (Table 3.7).
However, this might also be partly due to the fruit at the lower temperature taking
longer to ripen than the ones at the higher temperature. Collin (1989) also showed
considerable differences in weight loss during storage when comparing tempera-
tures of 13.5 and 22 °C (Table 3.8).

Table 3.7 Effects of harvest maturity (time after anthesis) and position of the hand on the bunch
on weight loss at colour stage 6 (Table 1.1) at 13 or 16 °C after initiation to ripen with ethylene.
Modified from Ahmad et al. (2001)
‘Robusta’ 13 °C ‘Grand Naine’ 16 °C ‘Grand Naine’ 13 °C
Position of hand on bunch
Time after anthesis Top Middle Lower Top Middle Lower Top Middle Lower
14 weeks 2.6 3.0 3.4 1.3 1.4 1.5 3.0 3.1 3.6
12 weeks 2.9 2.9 3.9 1.4 1.5 1.7 3.1 3.1 4.0
10 weeks 3.8 3.9 4.0 1.6 1.6 2.0 3.8 4.1 4.4
LSD (p = 0.05) 0.28 0.07 0.22

Table 3.8 ‘Orishele’ plantains harvested 82 days after flowering and stored for 21 days. Modified
from Collin (1989)
Temperature Weight loss % Pulp:peel ratio Acidity meq 100 g−1 TSS % Peel colour index
13.5 °C 8.9 2.2 8.1 22.1 6
22 °C 20.0 4.2 9.0 31.2 7+
Changes that Occur During Ripening 43

Moisture

Cultural conditions and water availability during growth have a relatively limited
effect on the water content of the fruit. The pulp of ‘Cavendish’ bananas had a
higher water content than the pulp of plantains. Marchal and Mallessard (1979)
showed that the moisture content of freshly harvested fruit was higher in the peel
than the pulp ranging from 86.6 to 91.9 % for ‘Cavendish’ and 85.0–87.1 % for
plantains in the peel and 60.3–74.2 % for ‘Cavendish’ and 60.4–60.6 % for plan-
tains in the pulp. The increase in the weight ratio between pulp and peel reflects
changes in moisture content. During ripening, the stomata, which were found to
be equal in number in plantains and ‘Cavendish’ bananas (370–450 cm−2), remain
functional. A low atmospheric pressure initiates a degradation of the epidermal
and stomatal tissues, thus reducing gaseous exchange including water vapour
(Collin and Folliot 1990). During ripening water loss through transpiration from
the peel, showed a pattern of changes similar to that of respiration (Palmer 1971).
The rise in the water content of the pulp during ripening was linked to the break-
down of carbohydrates and osmotic migration of water from peel to pulp because
of the higher concentration of sugars in the pulp (Fig. 3.4). This reflected the
thickness of the pulp, which progressively became thinner during ripening
(Fig. 3.5).

14 84

12 82
Tss (%) and acidity (meq/100g)

10 80
Water content (%)

8 78

6 76

4 74

2 72

0 70
0 5 10 15 20 25 30
Time (days)

Fig. 3.4 Changes in the water content (∗), total free acidity (■), and in the total soluble solids (+)
in the pulp of ‘Giant Cavendish’ bananas during the pre-climacteric and climacteric periods
(Marchal et al. 1988)
44 3 Fruit Ripening

Fig. 3.5 Changes in the

thickness of the peel (mm)
of three plantain genotypes
during ripening in Nigerian
ambient conditions of
21-33 °C and 79–99 %
RH. Modified from Ferris
et al. (1995)

Texture

As bananas ripen both the peel and the pulp become softer. Softening of bananas
during ripening appears to be associated with two or three processes (Smith 1989):
1. The first is the breakdown of starch to sugars, which showed close correlation
with softening (Fig. 3.6), since starch granules can have a structural function in
cells.
2. The second is the breakdown of the cell walls due to the solublization of pectic
substances and even the breakdown of cellulose. Smith found evidence of
increased activity of cellulase during banana ripening. Changes in enzyme activ-
ity can be due to variations in the pH and of the fatty acid composition of the
membranes. In tomatoes cellulase activity increased during softening (Huber
1983), but the pulp pectinesterase activity declined during softening. However,
there was no significant relationship, between the activity of pectinesterase or
their starch content, or the soluble polyuronide levels (Smith and Seymour
1990). Possibly these hydrolytic enzymes are induced by the increased concen-
tration of endogenous ethylene, independently of the effect of ethylene on respi-
ration (Brady 1987). Alterations in cell membranes increased their permeability
and diminished the rupture force of both the peel and the pulp, with different
genotype effects, in about 2 days before the climacteric rise. This could be linked
Changes that Occur During Ripening 45

25 6

Mechanical resistance (kg/cm2)

20
Starch (% fresh weight)

4
15
3

10
2

5
1

0 0
0 5 10 15 20 25 30
Time (days)

Fig. 3.6 Changes in the starch content of the pulp (■) and in the mechanical resistance of the fruit
(+) during the pre-climacteric and climacteric in ‘Giant Cavendish’ bananas. (From Marchal et al.
1988.)

to a variation of water potential and of the membrane unsaturated fatty acids,

which render the membranes more fluid and mobile (Wade et al. 1980). During
softening the concentration of soluble pectic polysaccharides, uronic acid and
related enzyme activities all increase, but these increases were not necessarily
linked to the increase in soluble pectin.
3. The possible third process involved in softening is the movement of water from
the peel of the banana to its pulp during ripening. This latter process could affect
the turgidity of the skin which would be enhanced by transpirational losses. This
change in the moisture status of the fruit also contributes to the ease of which the
peel can be detached from the pulp.
Lohani et al. (2004) reported that softening during ripening is generally attrib-
uted to degradation in cell wall structure, particularly the solublization of pectin
involving increased activities of various cell wall hydrolases. Their activity is regu-
lated by ripening-related hormones and/or other signal molecules and exposure to
endogenous ethylene stimulated the activity of PME, PG, pectate lyase and cellu-
lase in ‘Dwarf Cavendish’ over a period of 7 days after ripening was initiated with
ethylene. Pre-treating the fruit with either 1-MCP or IAA suppressed the ethylene
effects with ABA stimulated activity of all hydrolases except PG and was most evi-
dent for pectate lyase. Therefore, ABA can enhance softening with or without eth-
ylene. In contrast IAA suppressed their activity. Textural changes can vary with
different banana genotypes with the peel of ‘Cavendish’ softening during storage
and ripening while the peel of ‘Apple’ actually became slightly firmer (Table 3.9).
For the pulp, softening was progressive but ‘Cavendish’ was consistently firmer
than ‘Apple’.
46 3 Fruit Ripening

Table 3.9 Comparison of two banana genotypes on their texture. Modified from Bowden et al.
(1994)
Before storage After 28 days at 13 °C After ripening at 20 °C
Peel texture N mm2
‘Cavendish’ 2.05 0.84 0.55
‘Apple’ 1.95 2.18 2.02
Pulp texture N mm2
‘Cavendish’ 0.97 0.08 0.08
‘Apple’ 0.58 0.04 0.05

Flavour and Aroma

More than 250 volatile compounds have been identified in bananas (Imahori et al.
2013; Brat et al. 2004). Bananas mainly produce isoamyl and isobutyl esters, includ-
ing 3-methyl butyl and 2-methyl propyl esters of acetate and butanoate. These esters
are considered to be the major contributors to the aroma of bananas (Jayanty et al.
2002), providing the fruity smell (Wendakoon et al. 2006). The alcohols pentan-­
2-­ol, 3-methyl butanol, and 2-methyl propanol also contribute to the succulent char-
acter of banana aroma and the ketone pentan-2-one, which is one of the major
volatile compounds of bananas, imparts the characteristic banana flavour (Brat et al.
2004). McCarthy et al. (1963) and Tressl and Jennings (1972) had previously
reported that amyl esters give bananas their distinctive flavour and aroma, and butyl
esters give them a fruity flavour and aroma; however, other esters and aldehydes,
alcohols, and ketones have been associated with flavour, and their production rates
can increase during ripening. Salmon et al. (1996) showed that generally esters such
as butyl acetate, isoamyl acetate, ethyl acetate, butyl butanoate and isoamyl isobu-
tanoate are responsible for the characteristic aroma of fresh banana and constitute
the major class of compounds present in banana’s volatile profile.
Volatile compounds of several banana genotypes have been widely studied includ-
ing ‘Grand Naine’ (Bugaud et al. 2009); ‘Gran Enana’, a subgroup of the ‘Cavendish’
originating from Central and South America (Vermeir et al. 2009); various varieties
grown on Madeira Island (Nogueira et al. 2003); ‘Cavendish’ from Honduras (Jordan
et al. 2001); free and glycosidically bound volatile compounds of ‘Valery’ and
‘Pequeña Enana’ (Pérez et al. 1997); ‘Cavendish’ ‘Gran Enana’ and ‘Enana’ from
the Canary Islands, ‘Enana’ from Colombia (Cano et al. 1997); ‘Cavendish’, ‘Sen-
nin’ and ‘Delicious’ (hybrid between ‘Philippine’ and ‘Taiwanese’) (Shiota 1993).
Different volatilities and concentrations of those volatiles that can vary among the
different varieties (Cano et al. 1997). They identified the compounds and the ratios of
Spanish ‘Enana’, Spanish ‘Gran Enana’ and Latin American ‘Enana’ correlating
with differences among the flavours with Spanish ‘Enana’ being the richest in flavour
compounds. McCarthy et al. (1963) assigned the various components of banana
aroma and flavour to the amyl and isoamyl esters of acetic, propionic and butyric
acid, whereas the alcohols and carbonyls gave odours that they described as green,
woody or musty. Only Spanish bananas exhibited the presence of hexyl butanoate,
Changes that Occur During Ripening 47

which was related to a banana-like flavour. Pontes et al. (2012) evaluated the volatile
profiles of ‘Dwarf Cavendish’, ‘Prata’, ‘Maçã’, ‘Ouro’, and plantains and found a
total of 68 volatile organic metabolites that were tentatively identified and used to
profile the volatile composition in different Musa genotypes. Ethyl esters were found
to comprise the largest chemical class, accounting for 80.9, 86.5, 51.2, 90.1 and
6.1 % of total peak area for ‘Dwarf Cavendish’, ‘Prata’, ‘Ouro’, ‘Maçã’, and plantain
volatile fractions, respectively. Facundo et al. (2012) determined the volatile differ-
ences in two banana genotypes during cold storage and found that cold storage
affected the volatiles more strongly in ‘Nanicão’ than in ‘Prata’. Esters such as
2-pentanol acetate, 3-methyl-1-butanol acetate, 2-methylpropyl butanoate, 3-methyl
butyl butanoate, 2-methylpropyl 3-methyl butanoate and butyl butanoate were con-
siderably reduced during cold storage of ‘Nanicão’.
In India Venkata Subbaiah et al. (2013) ripened bunches of ‘Grand Naine’ and
found that their taste and flavour progressively increased up to the 6th day and there-
after decreased during the following 10 days. Tai-Ti Liu and Tsung-Shi Yang (2002)
tested ripening of ‘Pei-Chiao’ bananas (a ‘Giant Cavendish’ clone) at 20, 25 or
30 °C over 8 days and found that development of flavour compounds was highest
for those ripened at 25 °C and ethanol developed most rapidly at 30 °C.
Sanaeifar et al. (2014) tested a metal oxide semiconductor based electronic nose
to monitoring the changes in volatile production of bananas during ripening and
found that it was possible to distinguish between different ripening stages with
98.66 % accuracy.

Minerals

A general figure for the various minerals in bananas were given by USDA (Table 3.6)
but, the mineral composition of all crops varies with cultural conditions and differ-
ences have even been observed between bananas and plantains harvested from the
same plots (Table 3.10). Bananas have a high potassium content and Lee (2008)
reported that an average sized banana had 450–467 mg K. The average K content
for bananas grown in Hawai’i (‘Dwarf Brazilian’ and ‘Williams’) was 330.6 mg
100 g−1 fresh weight and Mg content averaged 35.1 mg 100 g−1. Iron, copper, and
manganese are other minerals of nutritional importance in bananas (Wall 2006).
Hardisson et al. (2001) reported that for bananas grown in Tenerife there was
5.09 mg g−1 K, 0.59 mg g−1 P, 0.38 mg g−1 Ca and 0.38 mg g−1 Mg on a fresh weight

Table 3.10 Average mineral composition (mg 100 g−1 fresh weight) before ripening of pulp of
‘Cavendish bananas’ (Musa AAA) and plantains (Musa AAB) harvested from the same plot
(modified from Marchal and Mallessard 1979)
P K Ca Mg S
Bananas 27 460 7 36 34
Plantains 32 440 14 32 24
48 3 Fruit Ripening

basis. Wall (2006) reported that ‘Apple’ bananas had higher concentrations of P, Ca,
Mg, Mn and Zn than ‘Williams’.
In Nigeria, Adeyemi and Oladiji (2009) gave the following for different stages of
ripeness:
Unripe 73.47 % ash, 0.68 % Zn, 0.146 % Mn, 0.506 % Co.
Ripe 77.19 % ash, 0.80 % Zn, 0.271 % Mn, 0.886 % Co.
Overripe 79.22 % ash, 0.78 % Zn, 0.118 % Mn, 0.756 % Co.
Changes in mineral content during ripening must be related to the changes in
mass of the fruit through water loss and to a lesser extent through mass losses due
to respiration. The loss of water from the peel in the course of ripening can explain
its enrichment in K, Ca and Mg (expressed in relation to their fresh weight). It is,
however, possible that a proportion of these elements migrate with the water from
the peel to the pulp, the K and Ca contents of which were shown to rise equally
(Izonfuo and Omuaru 1988).

Carbohydrates

As bananas and plantains develop on the plant almost all the carbohydrate is in the
form of starch, which accumulates during maturation, and in the pre-climacteric
phase there is little change in the principal carbohydrate metabolites (Wardlaw et al.
1939). During subsequent ripening the total carbohydrate content is progressively
reduced as the starch is broken down into reducing and non-reducing sugars (Barnall
1943). During the early part of ripening, sucrose is the predominant sugar, but in the
later stages glucose and fructose predominate. Starch is broken down to sucrose by
the action of sucrose phosphate synthetase and non-reducing sugars from sucrose
by acid hydrolysis. In bananas the breakdown of starch is usually completed during
ripening, but in plantains this breakdown is not complete even when they are con-
sidered to be fully ripe (George 1981). In ‘Apple’ bananas (Musa AA) the fruit was
found to still have residual starch when they were fully yellow and they may need
to be ripened further before being eaten depending on the consumers’ taste (Wei and
Thompson 1993). To illustrate this point, the caption on the label of some Musa AA
bananas, marketed in the UK, has “wait for brown freckles on skin before eating”
(Fig. 1.3).
In bananas, ripening involves a reduction in starch content from around 15–25 %
to less than 5 % in the ripe pulp, coupled with a rise of similar magnitude in total
sugars (Barnell 1943, Desai and Deshpande 1975, Lizada et al. 1990). Wardlaw
(1961) quoted figures that showed that the starch content of ‘Gros Michel’ could go
as low as 1% and sugar content as high as 19% when they were fully ripe. Venkata
Subbaiah et al. (2013) ripened ‘Grand Naine’ and showed that the total sugars
increased from 1.22 % to 24.05 % and starch content decreased from 21.52 % to
2.53 %. Adão and Glória (2005) harvested ‘Prata’ bananas at the full three-quarter
stage and stored them in polyethylene bags at 16 ± 1 °C and 85 % RH. Starch levels
Changes that Occur During Ripening 49

decreased throughout ripening, but after 7 days sucrose was prevalent and remained
constant then decreasing after 28 days while fructose and glucose levels increased.
However, at temperatures at or below 12 °C starch was no longer converted to sugar
(Wardlaw et al. 1939). The onset of the starch-sugar conversion has been shown to
be influenced by harvest maturity, with more mature fruits responding earlier. These
changes have been demonstrated in both triploid (Musa AAA) (Madamba et al.
1977) and diploid (Musa AA) fruit (Montenegro 1988). Arora et al. (2008) reported
that ‘Karpuravalli’ (Musa ABB) was rich in carbohydrates in terms of total starch
(1786 μg g−1 dry weight in the peel and 545 μg g−1 dry weight in the pulp) and sug-
ars (53.5 μg g−1 dry weight in the peel and 39.1 μg g−1 dry weight in the pulp).

Protein

Toledo et al. (2012) found that chitinases were the most abundant types of proteins
in unripe bananas. Two isoforms in the ripe fruit were implicated in their stress/
defence response and three heat shock proteins and isoflavone reductase were also
abundant at the climacteric stage. Pectate lyase, malate dehydrogenase and starch
phosphorylase accumulated during ripening. In addition to the ethylene formation
enzyme amino cyclo carboxylic acid oxidase, the accumulation of S-adenosyl-l-­
homocysteine hydrolase was needed because of the increased ethylene synthesis
and DNA methylation that occurred in ripening bananas.
Wade et al. (1972) reported that total nitrogen extracted by 5 % (w/v) trichlorace-
tic acid did not change significantly during ripening change and they found no evi-
dence of a movement of nitrogen from the peel to the pulp during ripening. However,
Dominguez-Puigjaner et al. (1992) working with ‘Dwarf Cavendish’ bananas found
evidence of differential protein accumulation in the pulp during ripening.

Phenolics

Bananas can contain high levels of phenolic compounds especially in the peel (von
Loesecke 1949). Tannins are perhaps the most important phenolic from the point of
view of fruit utilisation because they can give fruit an astringent taste. As fruit ripens
their astringency becomes lower, which is appearently associated with a change in
the structure of the tannins, rather than a reduction in their levels, in that they form
polymers (von Loesecke 1949, Palmer 1971). Mura and Tanimura (2003) confirmed
the loss of astringency during ripening of bananas and also found that it was by the
polymerizing of polyphenol compounds with a molecular weight of 2 × 105.
However, Fatemeh et al. (2012) showed that green bananas had higher total pheno-
lic compounds and total flavonoid compounds than those of ripe fruit. Radical scav-
enging activities (inhibition of DPPH) of the extracts ranged from 26.55 to 52.66 %.
Phenolics are also responsible for the oxidative browning reaction when the pulp of
50 3 Fruit Ripening

fruit (especially immature fruit) is cut. The enzyme polyphenoloxidase is responsi-

ble for this reaction (Palmer 1971). Bennett et al. (2010) detected catechin, gallocat-
echin, epicatechin and condensed tannins, in the soluble extract of banana pulp but
no soluble anthocyanidins nor anthocyanins. Fatemeh et al. (2012) evaluated the
stage of ripeness (green and ripe) of both pulp and peel on antioxidant compounds
and antioxidant activity of ‘Cavendsh’ and ‘Berangan’. The TPC ranged from 75.01
to 685.57 mg GAE 100 g−1 and TFC ranged from 39.01 to 389.33 mg CEQ 100 g−1
of dry matter. TPC and TFC values of the peel were higher than those of the pulp.
Although in ‘Berangan’ bananas the peel extracts appeared to have low TPC and
TFC, its antioxidant activity was moderate to high, which implies that antioxidative
compounds other than phenolics and flavonoids could be responsible for inhibition
of DPPH.

Acidity

Bananas and plantains, like most other fruits, are acid with a pulp pH below 4.5 (Von
Loesecke 1949). Palmer (1971) showed that the main acids in bananas were ascor-
bic, citric, malic and oxalic and the levels of these acids normally increase during
ripening. Total acidity in the pulp was shown to increase rapidly from about 28 to
67 mL NaOH 100 g−1 fresh weight from the pre-climacteric minimum to the climac-
teric and then it slowly declined post-climacteric to about 52 mL NaOH 100 g−1 fresh
weight (Wardlaw et al. 1939). These measurements were carried out on ‘Gros
Michel’ at 29.4 °C and 85% RH over a ripening period of 14 days. Marchal et al.
(1988) showed a slight increase in total acidity in ‘Cavendish’ during ripening
(Fig. 3.4). Also, in ‘Cavendish’, the pH of immature fruit was reported to be between
5.4 and 6.0 both in the peel and in the pulp, but the free acidity of the pulp was higher
(Thomas et al. 1986). During ripening pH decreased until it reached 4.0 at the fully
ripe stage in both ‘Cavendish’ bananas and in plantains to increase gradually there-
after. In green fruits free acidity was lower in ‘Orishele’ plantain (AAB) at 2 meq
100 g−1 fresh weight (Collin and Dalnic 1991) than in ‘Giant Cavendish’ bananas at
4 meq 100 g−1 fresh weight (Marchal et al. 1988). Free acidity increased until full
ripeness, more so in ‘Orishele’ (up to 10 meq 100 g−1 fresh weight) than in the
‘Cavendish’ (up to 7 meq 100 g−1 fresh weight). Collin (1989) reported that ‘Orishele’
plantains that had been stored for 21 days in air at 13.5 °C had acidity of 8.1 meq
100 g−1 but the acidity of those that had been stored in PE film bags was 4.5 meq
100 g−1. A similar effect was shown at 22 °C where the figures were 9.0 and 2.9
respectively. Ruiling Liu et al. (2016) reported that 1-MCP maintained acidity in
apples by regulating the balance between malate biosynthesis and degradation and
also delayed the postharvest loss of malate and citrate. In mangoes Medlicott et al.
(1987), Campbell and Malo (1969) and Fuchs et al. (1975) all indicated that expos-
ing fruit to endogenous ethylene had little effect in increasing the rate of acidity loss,
although Bhova et al. (1978) showed the converse, which may be a varietal effect.
Changes that Occur During Ripening 51

Ascorbic Acid

Bananas are a good source of vitamin C with a general level of about 8.7 mg 100 g−1
(Table 3.6). Seenappa et al. (1986) reported that the vitamin C level was different in
different varieties of banana and in ‘Dwarf Cavendish’ vitamin C level fell from
7.7 mg 100 g−1 in unripe fruit to 4.4 mg 100 g−1 in ripe fruit (Table 3.11) while the
vitamin C levels for ‘Cavendish’ ranged from 2.1 to 18.7 mg 100 g−1 (Leong and
Shui 2002, USDA 2012, Vanderslice et al. 1990, Wills et al. 1984). Also, in pre-­
climacteric fruit there was considerable variation in vitamin C content in different
genotypes (Table 3.12). These results agree with Wenkam (1990), who reported
vitamin C values of 5.1 mg 100 g−1 for ‘Williams’ and 14.6 mg 100 g−1 for ‘Dwarf
Brazilian’. Wills (1990) reported that ‘Sugar’ bananas (Musa AAB) had higher vita-
min C, starch, glucose, fructose and dietary fibre than ‘Cavendish’. Wall (2006)
reported the following analysis: vitamin C 12.7 mg 100 g−1, vitamin A 12.4 mg
RAE 100 g−1, total soluble solids 17.9 % and moisture 68.5% for ‘Dwarf Brazilian’.
For ‘Williams’ she reported: vitamin C 4.5 mg 100 g−1, vitamin A 8.2 mg RAE
100 g−1, soluble solids 20.5% and moisture 73.8%. The average vitamin C content
for ‘Dwarf Brazilian’ fruit ranged from 6.3 to 17.5 mg 100 g−1, and ‘Williams’
ranged from 2.5 to 6.3 mg 100 g−1 from four locations in Hawai’i. Ascorbic acid
content can decrease during ripening (Wenkam 1990). Ascorbic acid content of
‘Dwarf Cavendish’, ‘Rasabale’ and ‘Rajabale’ increased during ripening at 20 °C
for 21 days and then decreased slightly up to 35 days (Desai and Deshpande 1975).
In India Deekshika et al. (2015) compared the vitamin C content of mangoes and
bananas and found that mangoes had 54.78 ± 2.19 mg 100 g−1 and bananas had
20.13 ± 1.54 mg 100 g−1. Wills et al. (1984) detected 1.4 mg dehydroascorbic acid
100 g−1 in some of the bananas they tested but not in others. Vanderslice et al. (1990)
reported 3.3 mg 100 g−1 dehydroascorbic acid in bananas. These results may be
influenced by the particular test used.

Table 3.11 Ascorbic acid Ascorbic acid

content of the pulp of ripe Genotype mg 100 g−1
bananas. Modified from
‘Dwarf Cavendish’ Musa AAA 4.4
Seenappa et al. (1986)
‘Ney Poovam’ Musa AB 7.9
‘Silk’ Musa AAB 9.3

Table 3.12 Ascorbic acid Ascorbic acid

content of the pulp of unripe Genotype mg 100 g−1
bananas. Modified from
‘Bluggoe’ Musa ABB 16.0
Seenappa et al. (1986)
‘Robusta’ Musa AAA 10.3
‘Dwarf Cavendish’ Musa AAA 7.7
‘Gros Michel’ Musa AAA 6.3
52 3 Fruit Ripening

Carotenoids and Vitamin A

Vitamin A is crucial for human health, since it plays an important role in vision,
cell growth and the regulation of the immune system, but it cannot be synthesised
in sufficient quantities by the body and must be obtained from the diet. Plants do
not contain vitamin A but they contain carotenoids. Some carotenoids can be con-
verted into vitamin A during human digestion and are called provitamin A carot-
enoids (pVACs). In meat retinol is the main form of vitamin A. Measurement of
pVAC is commonly related to Retinol Activity Equivalent and is used for quantify-
ing the vitamin A value of sources of vitamin A, including both preformed reti-
noids in animal foods and precursor carotenoids in plant foods. Wall (2006)
reported that bananas are generally low in pVACs (Table 3.6), but some Fe’i
bananas that have yellow or orange pulp have been shown to contain relatively high
levels of carotenoids (Table 3.13). In ripe bananas, the major carotenoids were
lutein and carotene. ‘Dwarf Brazilian’ bananas had 96.9 mg β-carotene and
104.9 mg α-carotene 100 g−1, whereas ‘Williams’ averaged 55.7 mg β-carotene and
84.0 mg α-carotene 100 g−1. She also reported that bananas contained higher con-
centrations of lutein than of the pVACs. Average lutein concentrations were
154.9 mg 100 g−1 for ‘Dwarf Brazilian’ and 108.3 mg 100 g−1 for ‘Williams’. In
Hawai’i, ‘Apple’ bananas had an average of 96.9 μg β-carotene and 104.9 μg
α-carotene 100 g−1, whereas ‘Williams’ bananas averaged 55.7 μg β-carotene and
84.0 μg α-carotene 100 g−1.
Most carotenoids in bananas are in the peel with generally low amounts in the
pulp. Seymour (1986) found that the carotenoid content of banana peel could change
during ripening, depending on temperature. Those ripened at 35 °C had s­ ignificantly
increased carotenoids in the peel, while for in those ripened at 20 °C carotenoids
remained constant. The typical yellow colour of ripe banana peel is developed
purely because of the breakdown of chlorophyll, which masks the yellow colour in
unripe bananas. At high temperatures, the fruit remained greenish in spite of carot-
enoid synthesis, because chlorophyll content was only partially reduced. Wenkam
(1990) also found that the level of carotenoids increased during maturation and
ripening. These results confirm those of Von Loesecke (1949) and contradict those
of Gross and Flugel (1982) who found a decrease in carotenoids during the initial
phase of ripening of bananas. Measurement of the pVACs in 171 banana genotypes

Table 3.13 Carotenoid Total carotenoid content μg 100 g−1

content of different Fe’i Variety fresh weight
banana varieties. Modified
‘Aibwo’, ‘Suria’ 9400
from Sidhu and Zafar (2018)
and Englberger et al. (2006) ‘Fagufagu’ 5054
‘Ropa’ 5218
‘Gatagata’ 774
‘Toraka Parao’ 776
Changes that Occur During Ripening 53

showed that levels varied from undetectable in some fruit with white-cream pulp to
3500 μg 100 g−1 for fruit with yellow or orange pulp (Davey et al. 2009, Pereira
et al. 2011). Among Indian banana varieties Arora et al. (2008) reported ‘Red
Banana’ ranked highest in total carotenoids content for pulp (4 μg g−1 dry weight)
and β-carotene was estimated to be the highest in the peel (241.91 μg 100 g−1) and
in pulp (117.2 μg 100 g−1). Wall (2006) reported that ‘Williams’, harvested from
four locations in Hawai’i, had 8.2 mg RAE 100 g−1 for vitamin A and ranged from
6.1 to 9.3 mg RAE 100 g−1. This was higher than the 4.5 mg RAE 100 g−1 reported
by Wenkam (1990) for ‘Cavendish’. Sidhu and Zafar (2018) also showed that there
was considerable variation in carotenoids content in different banana genotypes
(Table 3.13 and Table 3.14).
The distribution of specific carotenoids in ‘Cavendish’ from the West Indies,
Central and South American were identified by Seymour et al. (1987) and were
predominantly lutein (Table 3.15). They also showed that levels of the different
carotenoids they detected could change during ripening and this could be affected
by the ripening temperature, but no consistent trends in these changes could be
detected.

Table 3.14 Carotenoid content of different banana varieties. Modified from Sidhu and Zafar 2018
and Beatrice et al. (2015)
Total carotenoid content μg 100 g−1
Variety fresh weight
‘Hung Tu’ Musa AAA 7,760
‘To’o’ Musa AA 7,765
‘Sepi’ Musa AA 10,067
‘Apantu’, ‘False Horn’ plantain Musa AAB 10,056
‘Bungaoisan’, ‘Ntanga 4’ Musa AAB 1,675

Table 3.15 Major carotenoid content in the peel of ‘Cavendish’ bananas before being initiated to
ripen and after being initiated to ripen with 1000 μL L-1 ethylene for 24 hours and ripened at either
20 or 34 °C for 5 days. Carotenoids were separated by HPLC with acetonitrile as solvent and
identified by comparison with authentic standards. Modified from Seymour et al. (1987)
Percentage total carotenoid content
Carotenoid Unripe fruit Ripened at 20 °C Ripened at 34 °C
Lutein 42.9 45.3 36.3
α and β carotene 7.1 9.8 7.3
Neoxanthin 3.8 2.6 3.2
Violaxanthin 3.5 3.3 6.8
Unknown 3.2 3.7 1.6
Unknown 1.2 10.0 7.2
Unknown 4.7 2.3 3.8
Unknown 0.3 3.0 1.7
54 3 Fruit Ripening

Folates

Folates are vitamin B9 and are a group of compounds derived from pteroylglutamic
acid. They are essential for in human development including the prevention of neu-
ral tube defects and cardiovascular and neurodegenerative diseases. Folate levels in
bananas was given as 20 μg DFE (1 microgram dietary folate equivalent = 0.6 μg
folic acid) (Table 3.6). García-Salinas et al. (2016) showed fluctuations in total
folate in bananas throughout ripening, but levels returned to their initial mature
green values when they were fully ripe. Interestingly, they found that initiating rip-
ening with exogenous ethylene increased folate by 51 %.

Other Phytochemicals

A phytochemical is a natural bioactive compound found in plant food that works

with nutrients and dietary fibre to protect against disease. Imam and Akter (2011) in
their phytochemical and pharmacological review gave the following phytochemi-
cals in various parts of bananas and plantains:
“catecholamines such as norepinephrine, serotonin, dopamine,
tryptophan, indole compounds,
pectin in the pulp,
flavonoids and related compounds (leucocyanidin, quercetin and its 3-Ogalactoside,
3-O-glucoside, and 3-O-rhamnosyl glucoside) from the unripe pulp of plantain.
serotonin, nor-epinephrine, tryptophan, indole compounds, tannin, starch, iron,
crystallisable and non-crystallisable sugars, vitamin C, B-vitamins, albuminoids,
fats, mineral salts in the fruit pulp.
acyl steryl glycosides such as sitoindoside-I, sitoindoside-II, sitoindoside-III, sito-
indoside-­IV and steryl glycosides such as sitosterol gentiobioside, sitosterol
myo-inosityl- β-D-glucoside.
a bicyclic diarylheptanoid,rel-(3S,4aR,10bR)-8-hydroxy-3-(4-hydroxyphenyl)
-9-methoxy-4a,5,6,10b-tetrahydro-3H-naphtho[2,1-b]pyran, and
1,2-dihydro-1,2,3-trihydroxy-9-(4-methoxyphenyl)phenalene, hydroxyanigoru-
fone, 2-(4-hydroxyphenyl)naphthalic anhydride, 1,7-bis(4-hydroxyphenyl)
hepta-4(E),6(E)-dien-3-one.
several triterpenes such as cyclomusalenol, cyclomusalenone, 24- methylenecyclo-
artanol, stigmast-7-methylenecycloartanol, stigmast −7-en-3-ol, lanosterol and
β-amyrin.
an antihypertensive principle, 7, 8-dihydroxy-3-methylisochroman-4-one from the
fruit peel.
cycloartane triterpenes such as 3-epicycloeucalenol, 3-epicyclomusalenol, 24-
methylenepollinastanone, 28-norcyclomusalenone, 24-oxo-29- norcycloarta-
none have been isolated from the fruit peel.
Changes that Occur During Ripening 55

cellulose, hemicelluloses, arginine, aspartic acid, glutamic acid, leucine, valine,

phenylalanine and threonine have been isolated from pulp and peel.
hemiterpenoid glucoside (1,1-dimethylallyl alcohol), syringin, (6S, 9R)-roseoside,
benzyl alcohol glucoside, (24R)-4α,l4 α,24-trimethyl-Sacholesta-8,25(27)-dien-­
3β-­o1 have been isolated from the flower.”
The ripe pulp of ‘Chinese Cavendish’, ‘Giant Cavendish’, ‘Dwarf Red’, ‘Grand
Nain’, ‘Eilon’, ‘Gruesa’, ‘Silver’, ‘Ricasa’, ‘Williams’ and ‘Zelig’ had similar
amounts of lipophilic extracts (about 0.4 % of DW). The major groups of lipophilic
components identified in these fractions were fatty acids (68.6–84.3 %) and sterols
(11.1–28.0 %), with smaller amounts of long chain aliphatic alcohols and
α-tocopherol (Vilela et al. 2014). Chanai Noysang et al. (2019) measured the phyto-
chemicals in ethanolic extracts of unripe pulp and peel of ‘Kluai Namwa’, ‘Kluai
Hom Thong’, ‘Kluai Leb Mu Nang’ and ‘Kluai Khai’ bananas. Kluai Khai pulp had
the highest amount of ethanolic extractives and the main phytochemicals identified
were alkaloids, flavonoids, tannins and polyphenols. ‘Kluai Hom Thong’ peel etha-
nolic extract had the highest antioxidant activity.
Chapter 4
Postharvest Treatments to Control
Ripening

Controlled Atmosphere Storage

CA on Pre-Climacteric Bananas

Burg and Burg (1967) showed that plant tissue was generally less sensitive to ethyl-
ene in atmospheres containing lower O2 or higher CO2 levels (Table 4.1). As
described earlier ethylene biosynthesis initiates banana ripening and the pre-­
climacteric period of ‘Cavendish’ bananas was shortened by exposure to levels of
ethylene as low as 0.1 ppm, with greater sensitivity to ethylene the more mature the
fruits at harvest (Liu 1976a), but less sensitive to ethylene in reduced O2 (4 kPa) and
increased CO2 (7 kPa) concentrations (Liu 1976b). Ahmad and Thompson (2007)
observed that the effectiveness of ethylene in hastening ripening was not reduced
with CO2 levels of less than 5 kPa and O2 levels of greater than 10 kPa. Previously,
Quazi and Freebairn (1970) showed that increased CO2 had the effect of inhibiting
the synthesis of endogenous ethylene and the respiration rate of bananas when the
pulp ripened. In an atmosphere containing 1 kPa O2, ripening was not initiated even
when exogenous ethylene was applied.
The production of ethylene was inhibited when the concentration of atmospheric
O2 was lowered to about 7.5 kPa and below, and tissues were then insensitive to
ethylene. Quazi and Freebairn (1970) showed that high CO2 and low O2 delayed the
production of ethylene in pre-climacteric bananas, but the application of exogenous
ethylene was shown to reverse this effect. Truter and Combrink (1990) found that
‘Dwarf Cavendish’ and ‘Williams’ remained hard and green for 6 weeks under
2 kPa O2 and 7 kPa CO2 at 12.5 °C. Thereafter the fruit ripened normally within
4–5 days when transferred to normal atmosphere at 16 °C. Ahmad et al. (2006)
studied the effects of CA storage (2 kPa O2 combined with 4, 6 or 8 kPa CO2) for
2 weeks on ripening of ‘Cavendish’ after removal to air. All treatments responded to

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2019 57

A. K. Thompson et al., Banana Ripening, SpringerBriefs in Food, Health,
and Nutrition, https://doi.org/10.1007/978-3-030-27739-0_4
58 4 Postharvest Treatments to Control Ripening

Table 4.1 Sensitivity at Sensitivity to

20 °C of pea seedlings to O2 kPa CO2 kPa ethylene μL L−1
ethylene in different levels of
0.7 0 0.6
O2, and CO2 with the balance
being N2 (Burg and Burg 2.2 0 0.3
1967) 18.0 0 0.14
18.0 1.8 0.3
18.0 7.1 0.6

ripening initiation with ethylene, in a similar way to fruit that had not been exposed
to CA, and produced good quality ripe fruit. All bananas, including the control,
reached colour stage 6 (Table 1.1) in 9 days after ethylene treatment. Imahori et al.
(2013) stored mature green ‘Cavendish’ in 0.5, 2, or 21 kPa O2 for 7 days at 20 oC
before ripening was initiated with ethylene. Concentrations of ethanol in mature
green fruit did not change during storage in both 21 and 2 kPa O2 atmospheres, but
increased in fruit stored in 0.5 kPa O2. They concluded that 0.5 kPa O2 is too low for
CA storage of pre-climacteric bananas but 2 kPa O2 was suitable.
At 15.5 °C ripening of bananas was retarded when they were stored in total N2,
but their flavour was poorer only if they were held in these conditions for longer
than 4 days or for over 10 days in 99 kPa N2 + 1 kPa O2 (Ahmad et al. (2006).
Klieber et al. (2002) found that storage of bananas in total N2 at 22 °C did not extent
their storage life compared to those stored in air but resulted in brown
discolouration.
The effects of CA storage on banana quality was tested by Kanellis et al. (1993)
who found that if CO2 concentrations exceeded 10 kPa, the pre-climacteric period
was prolonged, but the organoleptic qualities and the peel colour of ‘Pisang Mas’
was detrimentally affected. However, the quality of the ‘Lacatan’ (Musa AAA) and
‘Latundan’ (Musa AAB) stored in PE film bags under an atmosphere of 5 kPa O2
and 12.5 kPa CO2 at a temperature of 26–30 °C, was reported not to affect their
quality (Tiangco et al. 1987). Kanellis et al. (1993) have described how storage in
2.5 kPa O2 reduced respiration rate, slowed sugar accumulation and arrested the
normal colour changes in bananas. They concluded that the observed effects were
mediated via ethylene action rather than as a direct effect on respiration.

CA Effects After Ripening Initiation

Hesselman and Freebairn (1969) showed that ripening of bananas, which had
already been initiated to ripen by exogenous ethylene, was slowed in a low O2 atmo-
sphere. Wade (1974) showed that bananas could be ripened in atmospheres of
reduced O2, even as low as 1 kPa, but the peel failed to degreen, which resulted in
ripe fruit which were still green. Similar effects were shown at O2 levels as high as
15 kPa. Since the degreening process in ‘Cavendish’ is entirely due to chlorophyll
degradation (Seymour et al. 1987a) the controlled atmospheres storage effect was
Hypobaric Storage 59

presumably due to suppression of this process. Maneenuam and Doorn (2007)

showed that ‘Sucrier’ that had been initiated to ripen and then exposed to 90 ± 2 kPa
O2 had significantly increased peel spotting and the activity of PAL and PPO were
lower compared to the controls (18 ± 2 kPa O2). Quazi and Freebairn (1970) showed
that high CO2 and low O2 delayed the increased production of ethylene associated
with the initiation of ripening in bananas, but the application of exogenous ethylene
was shown to reverse this effect. Wade (1974) showed that bananas could be ripened
in atmospheres of reduced O2, even as low as 1 %, but the peel failed to degreen,
which resulted in ripe fruit which were still green. Similar effects were shown at O2
levels as high as 15 %. Since the degreening process in Cavendish bananas is
entirely due to chlorophyll degradation (Seymour et al. 1987, Blackbourn et al.
1990), the controlled atmosphere storage treatment was presumably due to suppres-
sion of this process. Hesselman and Freebairn (1969) showed that ripening of
bananas, which had already been initiated to ripen by ethylene, was slowed in low
O2 atmospheres.

Hypobaric Storage

A considerable amount of work has been done on the effects of reduced pressure in
the storage atmosphere of bananas by Stanley Burg. In his review Burg (2004)
reported that at 13.3–14.4 °C ‘Valery’ and ‘Gros Michel’ ripened in 10 days under
atmospheric pressure but remained green for 40–50 days under 152 mm Hg, but
mould could develop during this protracted period, which could damaage the fruit.
Burg (2004) also reported that ‘Valery’ stored at 13.3 °C under 6.4, 9.5 or 12.7 kPa
remained green for more than 105 days and ripened normally when they were trans-
ferred to atmospheric pressure and exposed to exogenous ethylene, with no deleteri-
ous effects on flavour or aroma. Burg (2004) also stored bananas under 49, 61, 76,
101, 122 or 167 mm Hg for 30 days and found they lost 1.1–3.6% in weight with the
higher losses at the higher pressures because of their higher respiration rates. All the
fruit were still green and there were no apparent differences between the different
hypobaric conditions. Burg and Burg (1965) showed delays in ripening of bananas
when they were stored under 125–360 mm Hg compared to those under atmospheric
pressure.
Apelbaum et al. (1977) stored ‘Dwarf Cavendish’ at 14 °C under 81, 253 or
760 mm Hg pressure and found that they began to turn yellow after 30 days under
atmospheric pressure, 60 days under 253 mm Hg but they still remained dark green
under 81 mm Hg. When they were subsequently transferred to atmospheric pressure
at 20 °C and exposed to exogenous ethylene they all ripened to a good flavour, tex-
ture and aroma. Bangerth (1984) successfully stored ‘Cavendish’ at 14 °C under
51 mm Hg for 12 weeks and found that when they were subsequently ripened in
50 μl L−1 exogenous ethylene they were the same quality as freshly harvested fruit
or those that had been stored under atmospheric pressure.
60 4 Postharvest Treatments to Control Ripening

Research has also been done on the effects of reduced pressure storage on the
response of bananas to exposure to exogenous ethylene. Bangerth (1984) found that
there was no diminished response to ethylene in storage under 51 mm Hg, whatever
the storage temperature tested. Liu (1976c) stated that at 14 °C the duration of the
pre-climacteric period of bananas, even those that had been exposed to ethylene,
was prolonged under low pressure (0.1 atmospheric pressure) or at a low concentra-
tion of O2. As well as delaying the initiation of ripening, hypobaric storage has also
been shown to delay the speed of ripening of bananas that have been initiated to
ripen. Quazi and Freebairn (1970) found that bananas that had been initiated to
ripen by exposure to exogenous ethylene did not ripen during hypobaric storage but
began to ripen within 1–2 days after transfer to ambient atmospheric pressure and
had good eating quality and texture. Liu (1976) initiated ‘Dwarf Cavendish’ to ripen
by exposure to 10 μL L−1 ethylene at 21 °C and then stored them at 14 °C for 28
days under hypobaric storage of 51 and 79 mm Hg or controlled atmosphere storage
of 1% O2 + 99% N2. All fruit remained green and firm and continued to ripen nor-
mally after they had been removed to ripening temperature in atmospheric
pressure.

Modified Atmosphere Packing

MAP has been used successfully to slow the ripening of bananas after they have
been initiated to ripen and to delay their ripening initiation (Fig. 4.1). This effect is
primarily due to changes in the O2 and CO2 in and around the fruit but, as reported
by Thompson et al. (1974), reducing fruit weight loss can also contribute to this
effect. This can also be shown with the effect of moist coir and perforated PE bags
that substantially delayed ripening, but were less effective that non-perforated PE
bags that would both reduce weight loss and substantially change the O2 and CO2
around the fruit (Table 4.2).
Nair and Tung (1988) reported that ‘Pisang Mas’ had an extension in their post-
harvest life of 4–6 weeks at 17 °C when they were stored in evacuated collapsed PE
film bags by applying a vacuum not exceeding 300 mm Hg. There are many other
reports of the positive effects of MAP on different varieties of banana in different
films from different countries including: Shorter et al. (1987), Tiangco et al. (1987),
Tongdee (1988), Abdullah and Pantastico (1990), Marchal and Nolin (1990), Satyan
et al. (1992), Wei and Thompson (1993), Chamara et al. (2000) and Chauhan et al.
(2006).
Vacuum packing is where bananas are placed in a plastic bag and the air is
extracted with a vacuum pump and the bags are sealed. MAP is commonly used for
the international banana trade (Fig. 4.2). Banavac® is a patented system which uses
large 0.04 mm thick PE film bags in which typically 18.14 kg of green bananas are
packed then a vacuum is applied and the bags are sealed and transported in cartons
(Badran 1969). Nair and Tung (1988) reported that 'Pisang Mas' bananas had an
extension in their postharvest life of 4–6 weeks at 17 °C when they were stored in
Modified Atmosphere Packing 61

Fig. 4.1 Changes in skin colour (1 = dark green and 6 = fully yellow) of five finger clusters of
‘Apple’ bananas stored at 13 °C in different thickness of polyethylene film bags after being initi-
ated to ripen. Source: Bowden 1993

Table 4.2 Effects of Days to Weight loss

wrapping and packing Packing material ripeness at ripeness
material on time to ripening
Not wrapped 15.8 17.0 %
and weight loss at Ripening
Index 7 of plantains stored at Paper 18.9 17.9 %
tropical ambient conditions Moist coir fibre 27.2 (3.5 %)a
of 26–34 °C and 52–87 % Perforated PE 26.5 7.0 %
RH (modified from PE 36.1 2.6 %
Thompson et al. 1972) LSD (p = 0.05) 7.28 2.81
a
The fruit actually gained 3.5% in weight

evacuated collapsed polyethylene film bags by applying a vacuum not exceeding

300 mm Hg.
Ahmad and Thompson (2007) tested the effect of MAP on the ripening and qual-
ity of ripe fruit. Bananas ripened in 0.05 mm of PE film bags had a shelf-life
extended by 5 days, as well as a slower the rate of softening and a more attractive
fresh appearance than those not in plastic bags. Organoleptic panellists preferred the
bananas ripened in PE bags to those not in PE bags because of their better flavour
and appearance. Bowden (1993) showed that the effect of MAP on ripening (changes
in yellowing of the peel) of bananas that had been initiated to ripen was related to
62 4 Postharvest Treatments to Control Ripening

Fig. 4.2. ‘Gros Michel’ vacuum packed in polyethylene film ready for export

the thickness of plastic and therefore the gas composition within the film (Fig. 4.1).
Yao et al. (2014) stored plantains sealed in polyethylene film bags of different thick-
nesses at 28 ± 2 °C and found that the concentration of O2 was reduced to 3.5 kPa
after 14 days in 20 and 30 μ bags, to 0.5 kPa in 40, 50 and 60 μ bags and to 0.3 kPa
in 70 and 80 μ thick. Film thickness also affected weight loss, for example, at 13 °C
Wei and Thompson (1993) showed that weight loss of ‘Apple’ bananas after 4 weeks
storage in PE film was 1.5% in 50 μm, 1.8 % in 37.5 μm and 2.1 % in 25 μm
while for fruit stored without packaging it was 12.2 %.

Ethylene Absorbents

Ethylene absorbents have been used, usually inside MAP to extend the postharvest
life of many types of fruit. Various brands have been marketed commercially e.g.
Green Keeper (GK) and Ethysorb, which contain potassium permanganate. More
recently sachets containing palladium (It’s Fresh!) has been successfully used as
ethylene absorbents. Kulkarni et al. (2011) stored ‘Robusta’ under active and pas-
sive modified atmosphere packaging at 12 ± 1 °C and 85–90 % RH for 2 seasons
and found that after 3 weeks a steady state of about 8.6 and 8.2% of CO2 and 2.8 and
2.6 % of O2 in passive MAP and MAP+GK packages, respectively were established
(Fig. 4.3). GK is a brand of sachet containing potassium permanganate. Weight loss
was 0.8 % after 7 weeks of storage, as against 5 % in openly kept green bananas
after 3 weeks. Both MAP and MAP+GK treatments delayed colour, texture, pulp to
Ethylene Absorbents 63

Fig. 4.3 Gas composition of MAP and MAP+GK for ‘Robusta’ in LDPE films. GK = KMnO4
sachets as Green Keeper. Modified from Kulkarni et al. (2011)

peel ratio and TSS content compared to controls that were stored non-wrapped.
Results indicated that the shelf-life of fruits packed under MAP and MAP+GK
could be extended by up to 5 and 7 weeks, respectively as compared to 3 weeks for
openly kept control fruits. Sensory quality of fully ripe fruits of both passive MAP
and MAP+GK treatments after ripening was very good.
Ethylene absorbents have also been used in banana transport. Scott et al. (1971)
found that during transport an alternative to refrigeration was sealing hands of green
fruit in polythene bags with an ethylene absorbent (potassium permanganate). The
fruit remained in a green, firm condition for up to 18 days at ambient temperatures,
during transport from North Queensland in Australia to Auckland in New Zealand.
This technique has also been successfully demonstrated to delay the ripening of
whole bunches of bananas during shipment (Wills et al. 1998). They estimated that
about 2 weeks additional storage life was obtained by packing potassium perman-
ganate in sealed bags with the fruit. Chauhan et al. (2006) stored unripe ‘Pachbale’
bananas at 13 ± 1 °C in modified atmospheres consisted of PE film pouches fol-
lowed by Ethrel induced ripening at 30 ± 1 °C. The PE packaging used was: not
flushed or flushed with 3 % O2 and 5 % CO2 or partial vacuum (400 mmHg) giving
a shelf-life extension to 15, 24 and 32 days, respectively as against 12 days for the
control. Application of an ethylene scrubber in combination with silica gel as a des-
iccant and soda-lime as a CO2 scrubber further enhanced the shelf-life to 18, 28 and
36 days, respectively under the different types of modified atmospheres specified.
One supermarket in Thailand used ethylene absorbing sachets in their MAP
bananas in an attempt to increase their shelf-life both whilst being offered for sale
64 4 Postharvest Treatments to Control Ripening

on their shelves and for customers at home. The temperature is relatively high in
both places and even under air-conditioning the temperature can be about 25 °C. It
was reported that this treatment was unsuccessful in extending the shelf-life of the
bananas (Patcharin Chitaurjaisuk personnel communication 2019) and it was not
used. It was suggested that this was not surprising since these bananas would have
been initiated to ripening by ethylene treatment before they were packed and there-
fore absorbing any ethylene in the pack would have no effect on speed of ripening.
Scott et al. (1971) found the storage life of pre-climacteric bananas could be
extended by at least 2 weeks by packaging them in sealed polyethylene bags with an
ethylene absorbent. They showed that the inclusion of potassium permanganate in
sealed packages of bananas reduced the mean level of ethylene from 395 down to
1.5 μL L−1 and reduced brown heart from 68 to 36 % in stored pears. Where potas-
sium permanganate was included in the bags containing bananas the increase in
storage life was 3–4 times compared to non-wrapped fruit and they could be stored
for 6 weeks at 20 or 28 °C and 16 weeks at 13 °C (Satyan et al. 1992). In tests with
pre-climacteric ‘Harichhaal’ (Musa AAA) bananas Kumar and Brahmachari (2005)
found that potassium permanganate-soaked paper in sealed PE bags was the most
effective packaging treatment for extending their storage life among those that they
tested.

Chemicals

Metal Ions

Domínguez et al. (1998) showed that exposing bananas to cobalt ions or silver ions
inhibited ethylene biosynthesis and both were found to inhibit both ethylene pro-
duction and ripening, even though silver is believed to act directly on ethylene
action only and cobalt ions inhibited induced respiration and ethylene production.
Silver ions effectively inhibited both the initiation and the continuation of tomato
ripening (Davies et al. 2006). They showed that the application of silver thiosul-
phate to mature green tomatoes prevented the appearance of several novel proteins
associated with ripening, including polygalacturonase.

1-Methylcyclopropene

1-MCP is an ethylene antagonist that has been used for controlling ripening and
senescence of fruit and vegetables by inhibiting both ethylene biosynthesis and
action by irreversibly binding to ethylene-receptors and strongly inhibiting the auto-
catalytic production of ethylene (Lurie 2008). Pathak et al. (2003) reported that
1-MCP application directly affected ACS transcript accumulation and inhibited the
autocatalytic pathway (System 2) in ‘Dwarf Cavendish’ bananas but did not affect
Chemicals 65

System 1 ethylene production. Kesar (2010) found that exposure of unripe mature
bananas to exogenous ethylene induced the expression of the gene MaPR1a, and
1-MCP exposure prior to ethylene treatment inhibited its expression. Ruiling Liu
et al. (2016) suggested that the mode of action of 1-MCP was related to the expres-
sion of acid transport genes, including MdVHA-A, MdVHP and Ma1. This was
because 1-MCP maintained fruit acidity in apples by regulating the balance between
malate biosynthesis and degradation. In addition, Trivedi and Nath (2004) addressed
the effects of 1-MCP on fruit firmness maintenance and found that treated bananas
had low expression of Maexp1, an ethylene induced expansin gene. 1-MCP applica-
tion also lowered the solubilisation of pectin due to suppression of cell wall hydro-
lases such as pectin methylesterase, PG, cellulase and pectate lyase activities in
bananas (Lohani et al. 2004).
Although 1-MCP treatment can delay the ripening process in climacteric fruit,
including banana, the efficiency of 1-MCP treatment is limited according to the
maturity of fruit at harvest, 1-MCP concentration, temperature and durations of
exposure (Bagnato et al. 2003; Pelayo et al. 2003; Moradinezhad et al. 2008). Harris
et al. (2000) suggested that the efficiency of 1-MCP in prolonging shelf-life of
bananas varied according to fruit maturity at harvest and in bananas it was less
effective at advanced maturity at harvest (Moradinezhad et al. 2008). They also
found that 1-MCP treatment was more effective in the fruit in the early part of the
climacteric and in hands from the top of bunch rather than from the bottom of the
bunch.
Bagnato et al. (2003) reported that the shelf-life of ethylene treated ‘Cavendish’
bananas was delayed by 300 μL L−1 1-MCP treatment which it was longer than non-­
treated fruit for 3 days. Jiang et al. (1999a and b) showed that banana sealed in
0.03 mm thick PE film bags containing 1-MCP at either 0.5 or 1.0 μL L−1 delayed
the ripening by about 58 days compared to non-wrapped, non-treated fruit. The
analysis of ethylene and respiration rate within the PE bags confirmed that 1-MCP
supressed both ethylene biosynthesis and respiratory rate in the banana fruit.
Subsequently Jiang et al. (2004) showed that exposure of bananas to mL L−1 1-MCP
slowed ripening but also enhanced chilling injury that was associated with increased
membrane permeability. Pinheiro et al. (2005) had studied the effect of 1-MCP con-
centration on postharvest quality of ‘Apple’ bananas during ripening. They found
that 1-MCP treatment for 12 h at the concentration of 50 nL L−1 was the best con-
centration in extending shelf-life based on total sugar, soluble pectins, firmness and
external appearance at the end of storage compared to higher concentrations of
1-MCP. In ‘Kluai Khai’ bananas (Musa AA), 1-MCP treatment at 250 ppb for 24 h
lowered respiratory rate and ethylene evolution as well as delaying ripening and
also prolonged their shelf-life to some 20 days during storage at 20 °C compared to
the fruit treated with lower concentrations (Jansasithorn and Kanlayanarat 2006).
Kesar (2010) found that exposure of unripe mature bananas to exogenous ethyl-
ene induced the expression of the gene MaPR1a, which increased with ripening and
1-MCP treatment prior to ethylene exposure, thus inhibiting gene expression.
Ruiling Liu et al. (2016) suggested that the mode of action of 1-MCP was related to
the expression of acid transport genes, including MdVHA-A, MdVHP and Ma1
66 4 Postharvest Treatments to Control Ripening

because 1-MCP maintained fruit acidity by regulating the balance between malate
biosynthesis and degradation. 1-MCP delayed the postharvest loss of malate and
citrate.
With ‘Cavendish’ bananas harvested at the mature-green stage, exposure to
0.01–1.0 μL L−1 1-MCP for 24 h delayed peel colour change and fruit softening,
extended shelf-life and reduced respiration rate and ethylene production (Jiang et al.
1999). They obtained similar results with bananas sealed in 0.03 mm thick PE bags
containing 1-MCP at either 0.5 or 1.0 μL L−1, but delays in ripening were longer at
about 58 days. Analyses of ethylene and CO2 concentrations within the PE bags
confirmed that 1-MCP suppressed both ethylene evolution and respiration rate.
Bananas treated with 1-MCP were shown to result in uneven degreening of the
peel. De Martino et al. (2007) initiated ‘Williams’ to ripen by exposure to 200 μL L−1
of ethylene for 24 h at 20 °C. These bananas were then treated with 200 nL L−1
1-MCP at 20 °C for 24 h then stored in > 99.9 kPa N2 + < 0.1 kPa O2 in perforated
PE bags at 20 °C. No differences in the accumulation of acetaldehyde and ethanol
were detected during storage compared to fruits not treated with 1-MCP. Peel
degreening, the decrease in chlorophyll content and chlorophyll fluorescence, were
delayed after the 1-MCP treatment. There was some general browning throughout
the 1-MCP treated peel in both the green and yellow areas of the ripening peel. They
concluded that it appears the 1-MCP treated peel, 24 h after the ethylene treatment,
may still undertake some normal senescence that occurs during banana ripening.
Green mature ‘Apple’ bananas were treated with 1-MCP at 0, 50, 100, 150 or
200 nL L−1 for 12 h and then stored at room temperature of 20 ± 1 °C and 80 ± 5 %
RH. 50 nL L−1 1-MCP was the best concentration in extending the shelf-life based
on total sugars, soluble pectins, firmness and external appearance in the end of stor-
age. The 50 nL L−1 1-MCP delayed the onset of peel colour change by 8 days while
at 100, 150 and 200 nL L−1 1-MCP the delay was by 10 days compared to non-­
treated (Pinheiro et al. 2005). ‘Khai’ bananas were treated with 1-MCP at 50, 100
or 250 ppb for 24 h then stored at 20 °C. 1-MCP delayed ripening and prolonged
storage to some 20 days with the higher concentration being more effective, but
bananas treated with 250 ppb 1-MCP had the lowest respiration rate and ethylene
production. However, there is no significant difference among treatments for colour
changes (Jansasithorn and Kanlavanarat 2006). Joyce et al. (1999) found that banana
ripening induced by propylene, could be delayed by exposure to 15 ml−1 L−1 of
1-MCP at 20 8 °C for 12 h. However, the 1-MCP treatment was less effective as
propylene-induced ripening progressed, although the eating-ripe condition of fruits
was maintained for a longer time than the non-treatment. Similarly, Yueming et al.
(1999) found that 1-MCP applied at concentrations in the range of 0.01–10 ml−1 L−1
at 20 °C for 12 h 1 day after ethylene treatment slowed the ripening of bananas, but
it was ineffective when applied 3 or 5 days after ethylene treatment.
1-MCP delayed or slowed ripening of pre-climacteric and climacteric bananas.
1-MCP treated pre-climacteric fruit eventually ripened, possibly due to synthesis of
new ethylene receptors. 1-MCP treatment became less effective as ripening pro-
gressed, especially after peak ethylene production. Nevertheless, 1-MCP treatment
extended the shelf-life of fruit when applied at later ripening stages, which may be
Chemicals 67

a commercial advantage. Thus, 1-MCP promises improved control of banana fruit

ripening when applied before or after exposure to exogenous ethylene (Joyce et al.
1999). Deaquiz et al. (2014) tested the interaction between 1-MCP (600 mg L−1)
and Ethrel (3 mL L−1) on ripening of yellow pitahaya fruits (Selenicereus megalan-
thus) and concluded that the 1-MCP reduced ethylene action and slowed the ripen-
ing. A patented system, marketed as RipeLock™ technology, treats bananas, after
they have been initiated to ripen, with 1-MCP and packs them in micro-perforated
film. The company claim that this combination of treatments “extends banana yel-
low life 4–6 days over any existing technologies.” (Anonymous 2019).

Salicylic Acid

Salicylic acid is a plant growth regulator that has been shown to stimulate defence
mechanism against both biotic and abiotic stresses and can inhibit ethylene biosyn-
thesis. It is widely accepted as a postharvest treatment for fruit and vegetables
(Asghari and Aghdam 2010; Supapvanich and Promyou 2013). Its mode of action
was reported to induce the activity of phenylananine ammonia lyase (PAL) activity
triggering the phenylpropanoid pathway resulting in the accumulation of phenolic
compounds and antioxidant activity in plants. It also induces heat shock proteins
and retards lipid peroxidation of membrane leading to increased tolerance to chill-
ing temperatures (Supapvanich and Promyou 2013; Aghdam et al. 2014). Pan et al.
(2007) also reported that treatment with salicylic acid alleviated chilling injury
symptoms by retarding relative electrical conductivity of tissue during storage of
bananas. They also found that salicylic acid retarded respiration rate, the change of
banana peel colour and the increases in PPO activity during storage. Thus, posthar-
vest treatment with salicylic acid has been used to control fruit ripening and physi-
ological disorders caused by stress during storage. Srivastava and Dwivedi (2000)
reported that postharvest immersion of ‘Harichhal’ bananas in salicylic acid at 500
or 1000 μM for 6 h could delayed their ripening. Fruit softening (caused by major
cell wall hydrolases such as PG, xylanase and cellulose), pulp to peel ratio, reducing
sugars content, invertase activity, respiration rate and the major antioxidant enzymes
(catalase and peroxidase) were all lower in salicylic acid treated bananas compared
to control. Anuchai et al. (2018) also showed that immersion for 30 mins in 2 mM
salicylic acid could maintain visual appearance and delayed senescence of ‘Hom
Thong’ bananas.

Gibberellic Acid

Vendrell (1970) reported that postharvest immersion in aqueous solution of GA3 at

10−6 to 10−2 M could delay ripening and yellowing of the peel of bananas. However,
Rossetto et al. (2003) found that exogenous application of GA3 to bananas impaired
68 4 Postharvest Treatments to Control Ripening

the onset of starch to sugars conversion and delayed sucrose accumulation by at

least 2 days, which they attributed to the disturbance of sucrose phosphate synthase
activity but GA3 application did not affect their ethylene synthesis or respiratory
rate. Archana and Sivachandiran (2015) reported that GA3 immersion of ‘Kathali’
bananas at 500 or 750 ppm prolonged their storage life, delayed peel colour change
and sugar accumulation, reduced the rate of increasing pH and the loss of fresh
weight.

Diazocyciopentadiene

The application of DACP to tomato fruit, while still on the plant, was shown to
retard the development of red color by blocking the ethylene receptor sites (Sisler
and Blankenship 1993). DACP is a weak inhibitor of ethylene responses, but upon
irradiation with visible light gives rise to much more active components (Sisler
1991), but it is prohibited for commercial use (Sisler and Lallu 1994).

Indole-3-Acetic Acid

Auxins are plant hormones that can retard ripening and senescence of fruit due to
suppression of ethylene. The effect of IAA on banana ripening is associated with the
reduction of the climacteric rise in respiration rate and starch to sucrose formation
possibly by affecting the activity of β-amylase (Purgatto et al. 2001; 2002). Lohani
et al. (2004) found that pre-treating ‘Dwarf Cavendish’ bananas with either 1-MCP
or IAA, suppressed the effect of exogenous ethylene application on the activities of
cell wall hydrolases such as pectin methylesterase, PG, pectate lyase and cellulose
over the period of 7 days after ripening was initiated by ethylene. Fernández-Falcón
et al. (2003) reported that spraying ‘Dwarf Cavendish’ bananas in the field with
IAA could induce their resistance to Panama disease.

Abscisic Acid

ABA is a plant hormone regulating fruit ripening and senescence in both climacteric
and non-climacteric fruits (Leng et al. 2014). In climacteric fruit, endogenous ABA
levels increased before ripening initiation and then decreased as the fruit ripened. In
both tomatoes and bananas there was an increase in ABA prior to the increase in
ethylene synthesis during ripening, and exogenous application of ABA induced eth-
ylene production through biosynthesis gene expression (Jiang et al. 2000; Zhang
et al. 2009a). However, in non-climacteric fruit, ABA levels increased from matura-
tion to harvest (Setha 2012; Leng et al. 2014). Treatment of bananas with ABA by
Chemicals 69

vacuum infiltration increased respiration rate and hastened ripening, effects that
were apparently independent of ethylene. Suppression of ABA led to a delay in fruit
ripening (Sun et al. 2012). Lohani et al. (2004) showed that in bananas ABA can act
synergistically with ethylene to promote softening. Pre-treating ‘Dwarf Cavendish’
bananas with ABA stimulated activities of PME, PG, pectate lyase and cellulose
and was most evident for pectate lyase. So, ABA has been shown to be able to
increase softening of the bananas with or without ethylene. Moreover, ABA, like
indole acetic acid, induces an increase in enzyme activity such as that of catalase,
peroxidase, acid phosphatase, phenoloxidases, unlike gibberellins (Vendrell 1969).

Lysophosphatidylethanolamine

LPE, a natural glycerolipid, has been approved for use as plant growth regulator for
horticultural application in some countries. It was first patented in 1992 (US
5126155) where it is claimed to enhance fruit ripening and delay their senescence
(Amaro and Almeida 2013). Postharvest treatment with LPE has been applied to a
range of fruits such as tomato (Farag and Palta 1993), red pepper (Kang et al. 2003),
cranberries (Ozgen et al. 2005) and Thompson seedless grapes (Hong et al. 2007).
In banana fruit, Ahmed and Palta (2015) reported that postharvest dip treatment
with LPE could increase marketability of ‘Cavendish’ bananas by improving shelf-­
life of the fruit by 1–2 days. The ripening and senescence of the LPE treated bananas
were delayed due to the reduction of respiration rate and also by maintaining mem-
brane integrity and decreasing in breakdown of starch and cell walls. To improve
water solubility of LPE, the combination of LPE and lecithin was applied to bananas
by Ahmed and Palta (2016). They found that the combination of 200 mg L−1 LPE
and lecithin could increase marketability of the fruit for 7 days after treatment and
gave better quality, involving higher pulp firmness, lower ion leakage from peel and
thicker peel as compared to LPE alone. Improved membrane integrity with a reduc-
tion in electrolyte leakage and ethylene biosynthesis was also observed in banana
peel treated with LPE (Ahmed and Palta 2011).

Aminoethoxy-Vinylglycine

AVG is an ethylene antagonist inhibiting the conversion of methionine to ACC

(Yang et al. 1979) and was shown to retard ethylene biosynthesis in bananas through
inhibiting ACC content and ACS activity (Cuadrado et al. 2008). Domínguez et al.
(1998) showed that banana slices, which had been infiltrated with AVG, were initi-
ated to ripen with exogenous ethylene showed stronger effects than expected. AVG
is marketed as ReTain, which contains 15 % w/w AVG, and has been registered for
use on apples in several countries. It is applied by spraying directly on fruit in the
field (Lurie 2008; Toan et al. 2011). Toan et al. (2010) suggested that 0.8 g L−1 of
70 4 Postharvest Treatments to Control Ripening

ReTain sprayed on ‘Cavendish’ bananas 2 weeks before harvesting, inhibited ethyl-

ene biosynthesis and lowered respiratory rate and postharvest losses. The shelf-life
of the ReTain treated bananas was extended to 16 days compared to 8 days for the
non-treated fruit. Toan et al. (2011) also suggested that the most appropriate of AVG
application for bananas was from 74 to 78 days after formation of the last hand of
the bunch.

Nitrous Oxide

NO as a key multifunctional-signalling molecule in plants mediating multiple phys-

iological and biochemical responses to biotic and abiotic stresses. It has been used
to maintain postharvest quality and extend shelf-life of several fruit and vegetables
(Wang et al. 2015a). It also induces plant defence against oxidative stresses by
reducing reactive oxygen species accumulation (Wu et al. 2014). Manjunatha et al.
(2012) reported that 1 mM sodium nitroprusside, a NO donor, delayed the peak in
respiration rate and sugar accumulation and lowered PPO and PAL activity during
ripening of ‘Cavendish’ bananas. Cheng et al. (2009) found that NO reduced ethyl-
ene evolution in banana slices during ripening due to the inhibition of ACO activity
and the suppression of MA-ACO1 gene transcription. They also found that NO treat-
ment retained higher contents of acid-soluble pectin and starch and retarded cell
wall hydrolases activity, which may account for the reduction of fruit softening.
Wang et al. (2015a) found that NO treatment delayed ripening of bananas by 2 days
compared to non-treated fruit and preserved chlorophyll content by inhibiting chlo-
rophyll degradation enzymes such as chlorophyllase and Mg-dechelatase activity
after cold storage. The tolerance to chilling injury in bananas during cold storage
was enhanced by NO treatment through promoting both the enzymatic and non-­
enzymatic antioxidant systems (Wu et al. 2014; Wang et al. 2015a) and maintaining
high levels of energy status and energy metabolism (Wang et al. 2015b). Palomer
et al. (2005) stored bananas at 20 °C and showed that ripening was delayed after NO
treatment, as judged by ethylene biosynthesis, respiration rate, colour change, acid-
ity and softening. However, they found that these effects were not detectable at
20 % NO concentration, but steadily rose at increasing concentrations above 40 %
and appeared to be saturated at 80 %. They concluded that the effects of NO in slow-
ing ripening could be due to its anti-ethylene activity without detrimentally affect-
ing banana quality.

Maleic Acid

Maleic acid (cis-butenedioic acid) is a colourless crystalline solid with a double

bond in the cis (Z) configuration that has a role as a plant metabolite and can be
conjugated to free base compounds or drugs to improve their stability, solubility and
Coatings 71

dissolution rate. Little work could be found on its effect on fruit ripening, but
Copisarow (1935) reported that maleic acid can inhibit ripening in several fruit,
including bananas, as well as protecting fruit from decay.

Coatings

Ncama et al. (2018) reported that coating fruit has been successful, not only in
reducing water loss and delaying senescence, but also in increasing the antimicro-
bial properties of the coated produce. Chitosan, gum arabic and several essential
oils, have been successfully used as fruit coatings (Thompson et al. 2016). Coatings
in the form of natural waxes or synthetic products have been shown to extend the
storage life of bananas and plantains by slowing gaseous exchange between the fruit
and the atmosphere, and thus delaying the onset of the climacteric. Water loss and
respiratory activity were reduced and the duration of the pre-climacteric period of
bananas and plantains was increased (Banks 1984; Marchal et al. 1988). Semperfresh
and Tal Prolong are names of commercial coatings based on sucrose esters of fatty
acids combined with CMC, which has proved effective with bananas. Semperfresh
and Tal Prolong are applied by dipping the fruit in a dilute suspension. When
allowed to dry they form a very thin, invisible layer which acts to modify the inter-
nal atmosphere of the fruit, particularly causing a fall in the O2 concentration within
the fruit, with much less effect on the internal concentration of CO2. Experiments in
which radioactively labelled sucrose ester coatings have been applied to bananas
show that these coatings do not migrate into the fruit, but remain on the surface
(Bhardwaj et al. 1984). The mode of action of a sucrose-ester coating on the physi-
ology of the ripening banana has been analysed in detail by Banks (1984). In a study
of the effect of Semperfresh on the preservation of bananas transported under refrig-
erated conditions and stored under non-refrigerated conditions (20 °C), Marchal
et al. (1988) found ripening to be delayed without any adverse effect on flavour.
Similarly, experiments carried out over a period of three years under tropical condi-
tions (Al-Zaemey et al. 1989) showed that coating plantains (Musa AAA ‘Dominico’,
‘Harton’ and ‘Horn’) and bananas (Musa AAA ‘Giant Cavendish’) with Semperfresh
slowed the major changes associated with ripening (Table 4.3), without affecting
the taste of the fruit. A coating similar to CMC was described by Deng et al. (2017).
This was cellulose nanofiber-based emulsion (0.3% cellulose nanofiber +1 % oleic
acid +1 % sucrose ester fatty acid, w/w on a wet basis), which they found reduced
respiration rate and ethylene production in bananas, thus delaying their ripening.
The coated bananas ripened normally during 10 days ambient storage; the coating
simply delayed the ripening processes. Maqbool et al. (2010) dipped bananas in a
composite coating of 10 % gum arabic +1 % chitosan. They showed that after
33 days storage coated fruit had 24 % lower weight loss and 54 % lower TSS than
those not coated and were firmer, had higher total carbohydrates and reducing sug-
ars lower than those not coated. The effects of coating bananas with coconut oil
(140 or 200 mL) combined with 25 g beeswax and 50 mL sunflower oil during
72 4 Postharvest Treatments to Control Ripening

Table 4.3 Effects of concentration of Semperfresh on plantains that had been ripened to fully
yellow at 29 °C. Modified from Al-Zaemey et al. (1989)
Semperfresh %
0 1 1.5 2 LSD p = 0.05
−1
Weight loss % day 0.62 0.55 0.38 0.26 0.14
Days ripening 3.3 5.3 6.3 6.7 0.36
Peel firmness kg 8 mm−1 6.4 5.6 5.6 4.4 0.35
TSS 18.3 17.5 17.5 18.5 ns

35 days storage compared to non-coated fruit showed better retention of colour and
appearance as well as of ascorbic acid (Mladenoska 2013). Truter and Combrink
(1990) showed that the storage life of ‘Dwarf Cavendish’ and ‘Williams’ was
extended when coated with Semperfresh. Banks (1984) reported that coating
bananas with TAL Pro-long modified their internal atmospheres by reducing the
permeability of the fruit skin to gases. Permeability of control fruit to CO2 was
greater than that of O2 and ethylene, and this differential permeability was enhanced
by coating. This resulted in a depression of the fruit’s internal O2 content which
affected ripening without a concomitant increase in the levels of CO2 which could
have proved toxic. The skins of coated fruit lost chlorophyll more slowly than con-
trols and there was a small effect on the accumulation of monosaccharides in the
fruit pulp. These effects were associated with depressed rates of respiration and
ethylene production in the coated fruit, but the accumulation of acetaldehyde and
ethanol was no more rapid than in controls.

Irradiation

Irradiation can have an effect similar to that of coating, but ripening and the organo-
leptic quality were modified to a certain extent depending on the dose (Thomas
1986). At 13 °C the length of storage was doubled (from 14 to 29 days) by an irra-
diation of 0.85 kGy, the development of the colour and of softening was blocked,
but peel splitting and a loss of aroma are observed (Brodrick and Strydon 1984).
PVA is a biodegradable polymer (Matsumura et al. 1999) and tannin it is a renew-
able natural material (Moric et al. 2014). Bananas were coated with thin films
with different ratios of plasticized PVA, CMC and Tannin solutions. Gamma irra-
diation improved their thermal properties, which provides suitable coating material
based on these natural biodegradable polymers for food preservation (Senna et al.
2014). Zaman et al. (2007) treated mature green bananas with 0.30, 0.40 or 0.50 kGy
for 5 mins and stored them at room conditions of 25 ± 2 °C and 80 ± 5 % RH. The
control bananas ripened within 6 days while the gamma irradiated bananas ripened
within 26 days. A minor decrease in the ascorbic acid content was the only adverse
effects observed in irradiated bananas and no other major changes occurred in nutri-
tional and organoleptic qualities with no differences detected between the three
levels of irradiation.
I rradiation 73

Extensive work on tropical fruits such as banana, mango and papaya have clearly
established that the maturity of the fruits at harvest, the time delay between harvest
and irradiation, and the physiological state of the fruit as related to its position in the
climacteric at the time of irradiation all can influence the radiation-induced delay of
ripening (Thomas 1986). Appreciable delays in ripening and consequent enhance-
ment of shelf-life in bananas and plantains has been reported when pre-climacteric
fruits were treated at doses in the range 0.2–0.4 kGy; the extent of delay of ripening
is dependent on fruit maturity at harvest and on storage temperature (Maxie et al.
1968, Kao 1971, Thomas et al. 1971, Thomas 1986). Irradiation delayed the onset
of the respiratory climacteric and the intensity of the respiratory peak, as evidenced
by CO2 evolution (Maxie et al. 1968, Thomas et al. 1971). Maximum delay of ripen-
ing has been observed with fruits of lower maturity (threequarters to full three-­
quarters); as fruit maturity increased, a progressive decrease in ripening delay and
shelf-life occurred (Kahan et al. 1968, Thomas et al. 1971). The optimal radiation
dose for inhibition of ripening, and the maximum dose the fruits can tolerate with-
out exhibiting phytotoxicity or radiation-induced injury, seem to differ among cul-
tivars and even for the same cultivar grown in different geographic areas. However,
irrespective of cultivar differences, doses exceeding 0.5 kGy invariably resulted in
browning or blackening of the skin in pre-climacteric bananas, while doses of 1 kGy
and more often caused splitting and softening of fruits. Fruits that are already in the
climacteric state can tolerate doses as high as 2 kGy without any external manifesta-
tion of radiation injury. The browning of skin occurring in pre-climacteric fruits
exposed to higher doses has been attributed to increased PPO activity (Thomas
1986; Surendranathan and Nair 1972).
Following the observation that irradiated bananas, when exposed to ethylene, take
longer periods to reach a comparable stage of ripeness than do non-irradiated fruits,
it has been postulated that the inhibition of ripening caused by irradiation involves a
decreased sensitivity to the ripening action of ethylene (Maxie et al. 1968, Thomas
et al. 1971). Gamma radiation. Alterations in carbohydrate metabolism by regulating
certain key enzymes of the glycolytic pathway, TCA cycle and gluconeogenic path-
way, and the resultant interference with ATP production required for various syn-
thetic processes during ripening have been reported as some of the main causes for
the delay of ripening in irradiated bananas (Surendranathan and Nair 1973 and 1980).
Predominance of the pentose phosphate pathway, accompanied by a gradual activa-
tion of fructose-1-6- diphosphatase, decreased succinic dehydrogenase activity and
the operation of glyoxalate shunt pathway, as evidenced by increases in isocitrate
lyase and malate synthetase, have been reported in ‘Dwarf Cavendish’ bananas irra-
diated in the pre-climacteric stage (Surendranathan and Nair 1972, 1976). An impair-
ment of succinic dehydrogenase activity has been observed in mangoes following
irradiation (Thomas et al. 1971). In mature mango fruits of the cultivar ‘Haden’, the
increase in NADP-malic enzymatic activity, usually observed during ripening, was
significantly diminished but not delayed by irradiation at 0.75 kGy (Dubery et al.
1984). An exponential decrease in the activity and significant differences in some of
the allosteric properties and kinetic parameters Vmax and Km of NADP malic
enzyme purified from mango fruit have been reported when irradiated in vitro
74 4 Postharvest Treatments to Control Ripening

(Dubery et al. 1984). However, it is unlikely that these effects observed in vitro are
the same as in intact fruits irradiated with the same doses.
An increase in ascorbic acid level was observed in ‘Gros Michel’ bananas irradi-
ated at 0.25–0.5 kGy, which was attributed to increased extractability (Maxie et al.
1968), whereas in five banana cultivars grown in India, irradiation at levels optimal
for delaying ripening (0.3 kGy) did not significantly affect ascorbic acid levels
(Thomas et al. 1971). In ‘Alphonso’ mangoes grown in India, the temperature at
which fruits were stored and ripened was found to influence the changes in ascorbic
acid levels during ripening. Increased niacin levels were observed in ‘Gros Michel’
bananas irradiated at 0.3 and 0.5 kGy, whereas the thiamin content was unaltered at
these doses (Maxie and Sommer 1968).

Temperature

In 1928 the Low Temperature Research Station was established in St Augustine in

Trinidad at the Imperial College of Tropical Agriculture. The initial work was con-
fined to “…… improving storage technique as applied to Gros Michel
[bananas]………..for investigating the storage behaviour of other varieties and
hybrids which might be used as substitutes for Gros Michel in the event of that
variety being eliminated by the epidemic spread on Panama Disease” (Wardlaw and
Leonard, 1940).
Symptoms of chilling damage to bananas include a reddish-brown streaking in the
skin associated with the vascular bundles, reduced latex flow, slow colour develop-
ment during ripening, a dull or greyish yellow colour in ripe fruit and brown skin in
advanced stages when they were exposed to 4–6 °C. More subtle changes are the
reduced production of volatiles in the ripening fruit and disturbances to the evolution
of CO2 and ethylene. A delay in the climacteric rise in CO2 occurs and multiple peaks
appear instead of one (Murata 2006). Beyond 25 °C the duration of the pre-­climacteric
is greatly shortened, and fruit quality is altered, because of modifications to
fruit metabolism during ripening. Above 35 °C the development of the peel and of
the pulp is desynchronized, with softening of the pulp proceeding faster than the
colouring of the peel. This results in fruit with a soft pulp but a green peel. The fruits
are then in a condition known as ‘cooked/boiled green’. Above 48 °C the climacteric
is not triggered and ripening is effectively blocked. If the temperature is too low it
can cause chilling injury even at 13 °C (Fig. 4.4). “Under peel ­discoloration” is a
mild form of chilling injury that occurs to bananas when they are stored at just below
the critical temperature/time combination. Longer exposures and lower temperatures
can result in more severe injury symptoms, as described above, including failure to
ripen, dull or greyish to the peel of ripe fruit and in severe cases the peel turns dark
brown or black, and even the flesh can turn brown and develop an off taste.
Harvey (1928) reported that “relatively high temperatures speeds up ripening
when they are allowed to ripen on the plant”. He reported that for green ‘Gros
Michel’, 4 or 5 days may be required for ripening at 65–70 °F, but at 80 °F this time
Temperature 75

Fig. 4.4 Chilling injury in ‘Cavendish’ in the form of under peel discoloration (UPD). (Photographs
Allen Hilton, Cranfield University)

may be decreased, but at 60°F. ripening will be much slower. Generally, it has been
found that ripening will be better throughout the pulp if the temperature is not much
above 65 °F. At that time Harvey (1928) reported that “high temperature has a ten-
dency to shrivel the fruit. However. jobbers frequently use high temperatures when
fruit is in demand because the fruit reaches a marketable color much more quickly
than at 65 °F. High temperature with attendant high humidity favors the develop-
ment of molds and the blackening of fruits, and hastens the rotting of the fruit stalk.
A blackened skin gives a bunch of bananas a poor appearance. The rotting of stalks
becomes a serious matter when the decay of the fingers allows the fruits to break
loose from the hand. Reynolds suggests the following temperatures for the control
of ripening rooms: 72 °F. (or over) Danger of cooking 68 °F. Fast or forced ripening
66–62 ° F. Normal ripening 60 °F. Slow ripening 58 °F. Holding green bananas
56 °F. Holding ripe bananas. There is some clanger of chilling even ripe fruits at
temperatures below 54 °F. Green bananas may show effects of chilling at slightly
higher temperatures. Incidentally, the banana will not freeze until the temperature
gets to a few degrees below the freezing point, 32 °F. However, badly chilled fruits
that have not been frozen are liable to turn black. If only slightly chilled they may
show a gray-green appearance after ripening. In ripening tomatoes, the use of high
temperatures (above 90 °F.) is to be avoided as far as possible, because the fruit rots
badly. Usually there are enough fungus spores on the fruits to cause rotting if condi-
tions are favourable. It is a problem, then, with the use of heat alone, to avoid rotting
discoloration, shrivelling, and poor flavor at high temperature, or to avoid the long
time required at low temperature. The time required to ripen ‘Cavendish’ bananas
by heat alone is so great as to exclude this genotype from commercial use.”
Since bananas suffer from chilling injury and recommended storage tempera-
tures are essential the minimum temperature that can be used before chilling injury
occurs, temperatures close to that minimum would be the best for maximizing their
postharvest life. The skins of fruit stored at below about 13 °C gradually darkened
with increasing relative electrolyte leakage. Under peel discolouration is a milder
form of chilling injury and can also occur during growth when night temperatures
drop to below 13 °C and is caused by latex coagulation in the peel laticifers and
subsequent browning of the latex by phenolic oxidation (Fig. 4.4). Chilling injury
of bananas stored below 13 °C was also reported to be associated with a decrease in
ethylene binding. The ability of tissue to respond to ethylene is evidently reduced,
thereby resulting in failure to initiate ripening (Yueming Jiang et al. 2004). However,
76 4 Postharvest Treatments to Control Ripening

chilling injury is time as well as temperature dependent that is why some authorities
give maximum time or a journey time. Recommendations for optimum storage of
green bananas include the following:
‘Giant Cavendish’
12.8–14.4 °C with 85–90 % RH (Pantastico 1975)
13–14 °C (Thompson and Burden 1995)
14 °C for ‘Grand Naine’ (Robinson and Saúco 2010)
‘Dwarf Cavendish’
11.5–11.7 °C (Wardlaw 1937)
‘Gros Michel’
11.5–11.7 °C (Wardlaw 1937)
11.7 °C (Hardenburg et al. 1990)
‘Lacatan’
14–14.4 °C (Wardlaw 1937)
‘Latundan’
14.4–15.6 with 85–90 % RH for 3–4 weeks (Pantastico 1975)
Non-specified varieties
13 °C (Purseglove 1975)
12.8–15.6 °C with 85–90 % RH for 4 weeks (Pantastico 1975)
12–14 °C and 90–95 % RH for 2–3 weeks (Mercantilia 1989)
13 °C and 85–90 % RH for 10–20 days (Mercantilia 1989)
13 °C and 85–90 % RH for 10–20 days (Snowdon 1990)
13–15 °C and 85–90 % RH for 1 month (Snowdon 1990)
13.3–14.4 °C and 90–95 % RH for all cultivars except Gros Michel (Hardenburg
et al. 1990)
14.4 °C and 85–95 % RH for 7–28 days (SeaLand 1991)
12–14 °C and 85–95 % RH for 15 days (Tchango et al. 1999)

Humidity

Water deficit in plant tissues may stimulate ethylene production and as a conse-
quence there can be an increase of tissue respiration rate (Yang and Pratt 1978).
Exposing fruit postharvest to low humidity may also result in sufficient stress to the
fruit to initiate ripening, as was shown by for plantains (Table 4.4).
Storage humidity was shown by Wardlaw and Leonard (1940) to affect respira-
tion rate of ‘Gros Michel’ bananas in that at 29.4 °C the climacteric was earlier for
those in 100 % RH compared to those in 70 % RH (Fig. 4.5). In experiments it was
Humidity 77

Table 4.4 The effects of the humidity of the incoming air in a flow through system on the ripening
of freshly harvested plantains in ambient conditions (23–33 °C) in Jamaica. Penetrometer force
was the force required to inject a 5 mm Magness and Taylor pressure tester probe (adapted from
Thompson et al. 1972)
Penetrometer
Storage Colour Weight loss Total soluble Pulp:peel force
humidity score % solids ratio Skin Pulp
100 % RH 2.3 0.9 10.6 1.9 12.0 6.6
0 % RH 7.0 18.4 24.9 3.5 8.2 1.8

280
260
240
Respiration rate (mg. per kg. per hr.)

220
200
High humidity

180
160
Low humidity

140
120
100
80
60
40
20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17
Days at 85°F. (29.4°C.)

Fig. 4.5 Effects of humidity on respiration rate of ‘Gros Michel’. Source: Wardlaw and Leonard
1940

shown that plantains stored in cartons without a packing material lost weight rapidly
during ripening and became blackened and shrivelled. Those packed in dry coir dust
lost less weight, and those in moist coir dust gained weight; no shrivelling was seen
following the latter two treatments and there was less skin blackening (Table 4.2).
The fruits in coir dust remained green for long periods before ripening rapidly
(Thompson et al. 1974). However, they took up moisture and would eventually split
if stored for too long.
Littmann (1972) found that water stress decreased the pre-climacteric life of
‘Giant Cavendish’ bananas and showed a depression in the climacteric peak of respi-
ration rate. Finger et al. (1995) reported that fresh mass loss of 5 % and higher pro-
moted shortening of the pre-climacteric life of bananas and induced a decrease of
maximum rates of respiration and ethylene production during climacteric ripening.
Pre-climacteric ethylene production was stimulated by water stress. Fruit with a
mass loss of 20 % ripened abnormally with a decreased pulp softening and excessive
78 4 Postharvest Treatments to Control Ripening

brown colour of skin. Akkaravessapong et al. (1992) examined the effect of approxi-
mately 50 % RH, approximately 70 % RH and approximately 90 % RH on the sus-
ceptibility of bananas to mechanical damage from 2 days after harvesting until 5 days
after ripening had been initiated. Humidity did not influence susceptibility to
mechanical injury, the tissues damaged at 50 % RH dried to a black colour while
those damaged at 90 % RH remained light brown. George and Marriott (1985) found
that in plantains an increase of weight loss of 0.5 % per day resulting from low stor-
age humidity was typically accompanied by a decrease of 65 % in the potential stor-
age life. They also found that treatment with gibberellin extended the pre-climacteric
period when subsequent storage was under high humidity (unstressed conditions),
but had no effect under low humidity (water stressed conditions). Ullah et al. (2006)
observed that high humidity delayed the ripening of bananas and consequently
increased their shelf-life. Storage of bananas submerged in water delayed the climac-
teric phase of bananas in a similar way to controlled atmosphere storage but they did
not complete their ripening processes properly. Storage of bananas submerged in
water resulted in splitting of bananas due to excessive intake of water. Microbial
spoilage of water-stored bananas emerged as serious problem against successful
application of this technology. Reduced dissolution of O2 into water affected the
ripening process, and in order to attain the best quality and increased shelf-life, water
storage was found to be not suitable for bananas but storage in 100 % RH showed
better results.
The effects of storage in polyethylene film bags on the postharvest life of fruits
may also be related to moisture conservation around the fruit as well as the change
in the CO2 and O2 content. This was shown by Thompson et al. (1972) for plantains
where fruit stored in moist coir or perforated polyethylene film bags had a longer
storage life than fruits that had been stored unwrapped. But there was an added
effect when fruits were stored in non-perforated bags, which was presumably due to
the effects of the changes in the CO2 and O2 levels (Table 4.2). So, the positive
effects of storage of fresh pre-climacteric fruits in sealed plastic films may be, in
certain cases, a combination of its effects on the CO2 and O2 content within the fruit
and the maintenance of high moisture content. The effect of moisture content is
more likely to be a reduction in stress of the fruit that may be caused by a rapid rate
of water loss in non-wrapped fruit. This in turn may result in increased ethylene
production to internal threshold levels that can initiate ripening.
George and Marriott (1982) reported that the lower the humidity, the greater the
loss of water and the shorter is the duration of the pre-climacteric period, with plan-
tains being more sensitive than other Musa clones tested. Low humidity brought
forward the production of ethylene and the climacteric rise in respiration, but its
effect on the level of ethylene production varied with genotype. For example, in
plantains (Musa AAB ‘Apem’) the production of ethylene and the level of ACC
were increased (George and Marriott 1985), while Nair and Tung (1988) observed
the opposite for ‘Pisang Mas’ (Musa AA). For all the clones studied, a reduction in
humidity did not alter the rate of respiration, but affected the pulp and peel ratio,
peel colour, pulp softening and in TSS (Broughton et al. 1978; George and Marriott
1985; Nair and Tung 1988). These effects were shown for plantains (Table 4.2).
Chapter 5
Initiation of Ripening

Burg and Burg (1967) reported that carbon monoxide, butylene, propylene and acet-
ylene all had biological activity similar to that of ethylene when tested with the pea
straight growth test. They also stated that a pre-requisite for effectiveness in initiat-
ing banana ripening appears to be that they must be unsaturated, that is, have a
double or triple bond between carbon atoms. Dhembare (2013) also named several
chemicals that had been used in banana ripening including calcium carbide, acety-
lene gas, carbon monoxide, potassium sulfate, Ethrel, potassium dihydrogen artho-
posphate, putrisein, oxytocin and photoporphyrinogen. However, gases other than
ethylene that have been shown to initiate ripening of bananas were all considerably
less effective than ethylene. Kader A.A. (1992 quoted by Siddiqui and Dhua 2010)
reported the comparative effectiveness of ethylene and other compounds in initiat-
ing fruit ripening where ethylene = 1, propylene = 130, vinyl chloride = 2370, car-
bon monoxide = 2900, acetylene = 12,500 and 1-butene = 140,000, although it is
difficult to reconcile some of these figures with published experimental work.
Ripening of individual bananas may vary within a bunch and even within a hand
(Fig. 5.1), therefore for most commercial practice fruit are harvested at the pre-­
climacteric stage and initiated to ripen by application of a chemical exogenously. In
some local markets it is actually preferred that all the bananas in one hand do not
ripen at the same time, because this allows them to be eaten in sequence since many
people like to eat only one or two bananas each day so initiating them to ripen all at
the same time is a disadvantage.
In the experience of the authors many people believe that fruits that are not initi-
ated to ripen exogenously by a chemical taste better. This has been studied and
results have proved inconclusive. Limited tests carried out with academic Food
Technologists in Thailand also proved inconclusive, but generally slightly favoured
those that had been initiated to ripen exogenously (Jiraporn and Thompson 2019
unpublished). Sonmezdag et al. (2014), based on sensory analysis, reported that
banana not exposed to exogenous ethylene were preferred because of their better
fruity aroma and general impression attributes.

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2019 79

A. K. Thompson et al., Banana Ripening, SpringerBriefs in Food, Health,
and Nutrition, https://doi.org/10.1007/978-3-030-27739-0_5
80 5 Initiation of Ripening

Fig. 5.1 ‘Namwa’ (Musa AAB) ripened naturally on sale at the road side (a) and in a supermarket
(b) in Bangkok in March 2019.

A comparison of the quality of bananas ripened with or without exogenous eth-

ylene to the same skin colour was made by a descriptive analysis panel (Scriven
et al. 1989). They found the pulp of the fruit ripened without exogenous ethylene
was less green, more fruity and softer than the exogenous ethylene ripened fruit.
The fingers of the same hand of a bunch may also initiate to ripen at different times.
Also, the production of endogenous ethylene by fruits placed in the same conditions
varies between genotype. For example, Karikari et al. (1979) showed that under
conditions where ‘Cavendish’ (AAA) produced 2.1–2.5 μL−1 kg hr.−1 ethylene, tet-
raploids (Musa AAAA) produced 6.7 μL−1 kg hr.−1 ethylene and ‘Apem’ (AAB)
produced 34.8 μL−1 kg hr.−1 ethylene. When a local Thai farmer, who grows bananas
on a small scale, was asked which method he uses for initiating ripening he replied
that “there are 2 reasons for non-using ethephon to activate banana ripening. The
first, using ethephon makes more rapid spoilage on banana than calcium carbide. He
said, it may come from water for preparation ehtephon solution. Secondly, using
ethephon takes shorter time than calcium carbide for banana ripening. It is too fast
for sale. When he would like to sale banana, he needs time for waiting customers. If
the banana goes ripening too fast, it will go to spoilage too fast as well. This situa-
tion is not good for him (sale person)” (translated by Jiraporn Sirison). Jiraporn and
Thompson (unpublished, 2019) ripened ‘Namwa’ bananas at 29–31oC and 71–86%
RH either with ethylene or left them to ripen “naturally”. Those initiated with eth-
ylene were fully ripe and an even yellow colour after 4 days while those ripened
“naturally” took 8 days to ripen and still had green tips on each finger, which per-
sisted even when they were over-ripe and very soft after 11 days.

Ethylene

Ethylene is used in fruit ripening rooms, which is a colourless gas with a sweetish
odour and taste, has asphyxiate and anaesthetic properties and is flammable. Its
flammable limits in air are 3.1–32% volume for volume and its autoignition
Ethylene 81

temperature is 543 °C. Care must be taken when the gas is used for fruit ripening to
ensure that levels in the atmosphere to not reach 3.1%. As added precautions all
electrical fitting must be of a “spark-free” type and warning notices relating to
smoking and fire hazards must be displayed around the rooms.
Ethylene was first identified by Neljubow (1901) and has been known to have
physiological effects on crops for over 100 years. F.E. Denny used ethylene for rip-
ening bananas, tomatoes and pears and subsequently took out a patent (Denny
1923). Ethylene was first identified as a volatile chemical produced by ripening
apples by Gane (1934). It was assumed that ethylene was produced only in climac-
teric fruits during the ripening phase, but with the development of chromatographic
analytical techniques, it is clear that all crops are able to produce ethylene under
certain conditions (Hulme 1971). In his review, Curry (1998) reported that ethylene
not only initiates ripening of climacteric fruit but can also induce fruit abscission,
induce flowering, promote seed germination, break dormancy, promote root initia-
tion, delay sprouting and promote plant dwarfing.
Bananas are sensitive to physiological levels of ethylene as low as 0.3–0.5 μL L−1
if the O2 and CO2 levels are similar to those found in outside fresh air (Peacock
1972). Biale (1960) and Gane (1936) reported that 0.1–1 μL L−1 levels of ethylene
were required to initiate ripening. The main factors affecting response to exogenous
ethylene are: fruit maturity, time after harvest when ethylene exposure began, tem-
perature and the length of exposure to ethylene. ‘Cavendish’ bananas have been
shown to be more sensitive to ethylene at 35 °C than at 20 °C (Seymour et al. 1985).
Traditionally in commercial rooms the fruits were maintained in an atmosphere of
1000 μL L−1 ethylene for 24 h, generally between 16 and 18 °C and humidity as
high as possible (around 95% RH) (Thompson and Seymour 1982; Nelson nd). As
was described earlier, the various changes that occur during ripening might not all
occur simultaneously. For example, in pears a differential effect on the initiation of
individual ripening processes has been found. Pears required 0.08 μL L−1 ethylene
to initiate softening while initiation of the respiratory climacteric required
0.46 μL L−1 (Wang and Mellenthin 1972).
The effect of endogenous ethylene on ripening is universally accepted to follow
a time–concentration relationship with the higher the concentration of ethylene and
the longer the exposure time to ethylene, the faster the initiation of ripening (Wills
et al. 1998). Recommended concentrations for the induction of ripening of avoca-
dos, bananas, honeydew melons, kiwifruit, mangoes, stone fruits and avocados are
between 10 and 100 μL L−1 (Saltveit 1999). Knee (1985) reported a threshold level
of 0.1–0.5 μL L−1 to initiate the ripening of avocado, banana, honey dew melon and
pear. However, studies on kiwifruit have found that ripening was initiated by levels
of 0.01 μL L−1 (Mitchell 1990). Peacock (1972) speculated that for bananas there is
no effective threshold level of ethylene that induces ripening, although he examined
short term exposure to ethylene levels >0.1 μL L−1 rather than longer exposure to
concentrations <0.1 μL L−1. ‘Cavendish’ and ‘Lady Finger’ bananas were stored at
15, 20 and 25 °C in an atmosphere containing 0.001, 0.01, 0.1 and 1.0 μL L−1 ethyl-
ene in air and as expected, green life increased as the temperature and ethylene
concentration decreased (Wills et al. 2014). Postharvest exposure to ethylene can
82 5 Initiation of Ripening

have negative effects on crops including: apples – scald, aubergines - brown spots,
potatoes – sprouting, grapes – mould, onions and garlic – odour, broccoli – yellow-
ing, carrots – bitterness and leafy vegetables - loss of green colour.

Cylinders

Large gas cylinders containing ethylene under pressure can be used in rooms to
initiate banana ripening (Fig. 5.2). One method of application is to calculate the
volume of the ripening room and from a cylinder of ethylene gas, enrich the ethyl-
ene level until it reaches the required amount. This is done with a stop watch and
flow meter attached to the pipe connecting the room with the gas cylinder (Fig. 5.3).
The problems associated with this method are that ethylene can be explosive in air
and, although only 0.1% is commonly used a mistake can easily be made and ethyl-
ene concentrations may reach explosive levels. The second problem is that if the
ripening room is not completely gas tight then the ethylene may leak and the level
fall below the threshold that is required to initiate ripening. The former problem is

Fig. 5.2 Large cylinders of ethylene outside a banana ripening room in Brazil
Ethylene 83

Fig. 5.3 Initiation of

ripening using an ethylene
cylinder

partially addressed by diluting the ethylene in the cylinders with nitrogen giving
about a 5% ethylene concentration which gives a potential greater margin of error if
a mistake is made when the gas is being applied. Also, the ethylene gas may be
applied in small cylinders which are called ‘lecture tubes’. These contain a small
amount of pure ethylene (often 35 L) under pressure. The whole of the contents of
the tube is thus released into the room. If, for example, the ripening room had a
volume of 70 m3 and a desired ethylene concentration of 1000 μL L−1 was required
then two 35 L lecture tubes would be discharged into the room.
Ripening of bananas in Australia was reported to be at 13–14 °C for 2–7 days,
with the length of time depending on the distance of the wholesaler from the port of
entry and on market demand. To ripen the bananas, the polythene bag in which the
fruit was stored, is opened and ethylene gas applied to the fruit. Controlled ripening
is undertaken at 14.5–21 °C. Humidity is generally maintained at 85–95% RH in the
early stages of ripening and, once a trace of colour appears, it is reduced to 70–80%.
Ripening rooms are usually gassed with ethylene on each of two successive days
(Story and Simons 1999).
84 5 Initiation of Ripening

Catalytic Generators

Ethylene generating equipment, which gives slow release of ethylene over several
hours, is now available; it addresses both the problems of room leakage and mini-
mizes the hazards of use. Suitable equipment is manufactured commercially which
gives a constant stream of ethylene over a 16 h period (Fig. 5.4). The number of
ethylene generators required depends on the size of the room. Catalytic generators
probably generating ethylene by heating ethanol in a controlled way in the presence
of a copper catalyst. Care should be exercised in doing this because of the inflam-
mability of the alcohol. They are patented and careful controls need to be estab-
lished to ensure when such a process is set up it is completely safe.

Ethrel

Compounds, in which the fruit can be dipped and decompose in or on the fruit to
release ethylene can have the advantage of easy application. Etacelasil (2-­chloroeth
yl-­tris-[ethoxymethoxy] silane) or ACC (1-aminocyclopropane-1-carboxylic acid)
have not been used practically for the application of ethylene but 2- chloroethyl
phosphonic acid is marketed as Ethrel and also called Ethephon. Reid (2002)
reported that the use of Ethrel in the USA is approved on some crops in certain
States and circumstances including postharvest fruit ripening of bananas, tomatoes
and peppers. Ethrel reacts with water releasing ethylene but it is also hydrolysed in
plant tissue to produce ethylene, phosphate and chloride.

OH -
Cl - CH 2 - CH 2 - PHO3 - ® CH 2 = CH 2 + PH 2 O 4 - + Cl -

Fig. 5.4 Ethylene generator being used in a ripening room. Reproduced with permission of Greg
Akins, Catalytic Generators Inc., Norfolk, VA. USA
Ethylene 85

In practice ethylene can also be released from Ethrel by mixing it with a base such
as sodium hydroxide. For example, Ethrel C (a proprietary brand) will release 93 g
from 1 L or 74.4 L of ethylene gas per litre of Ethrel. It has been used in this way to
initiate fruit to ripen by placing containers of Ethrel in a gas tight room containing
the fruit and then adding the base to the containers. This method was used in com-
mercial ripening rooms in the Yemen, using mainly ‘Dwarf Cavendish’ and some
‘Poyo’. Buckets of dilute sodium hydroxide solution were placed throughout the
ripening rooms. When all these were in place measured amounts of Ethrel are added
to each bucket, which gave an instant release of ethylene gas into the room. The
mixture was 80 mL Ethrel and 1 L of sodium hydroxide for every 300 m3 of room,
in theory the temperature of the pulp of the banana should be 20–22 °C, but in prac-
tice the pulp temperature was 29–30 °C. The rooms remained closed for 3 days, but
rubber gaskets on the doors were pulled away in places and had been damaged by
fork lift trucks so they were far from gas tight. On removal after 3 days the fruit
were quite soft, but still green and lacking in flavour. A comparison was made
between using this method and initiating ripening with calcium carbide. The Ethrel
method was found to be some 50 times more expensive than using calcium carbide
but was more effective than calcium carbide (Thompson 1985).
Venkata Subbaiah et al. (2013) ripened bunches of ‘Grand Naine’, half the fruit
were dipped in Ethrel and the other half not dipped, at room temperature (25–29 °C
and 79–89% RH). The Ethrel treated fruit ripened in 4 days with excellent quality
attributes, whereas non-treated fruit took 10 days to ripen. Ram et al. (2009) dipped
unspecified genotypes of bananas in various concentrations of Ethrel (1000, 2000,
3000 or 4000 ppm) and found little difference between the concentrations, and fruit
dipped in 1000 ppm for 5 minutes started to soften in 3 days (compared to 9 days
for the control) and became ready to consume in 5 days with shelf-life of 8 days.
They were considered to have an excellent flavour. In Thailand Ethrel is used for
fruit ripening as well as calcium carbide. Ethrel was purchased locally (March
2019) for THB 350 per litre. Two overflowing capfuls were added to half a bucket
of water (Fig. 5.5) and the hands of bananas were dipped in the bucket so that they
were fully immersed. The amounts used by local traders were measured, and it was
found that they added about 30 mL of Ethrel to 7.5 L of water which gave an
approximately 250 ppm solution. The recommendation on the bottle of Ethrel for
ripening bananas was 500–1500 ppm, so they were using considerably less than the
recommended level. After dipping the fruit were placed on a table to ripen at a con-
stant temperature of 29–31 °C (Fig. 5.6). Colour changes were perceived in the
Ethrel treated hands after 2 days. After 4 days the non-treated hands appeared to be
exactly as they were 4 days earlier with latex exuding from the skin when they
were peeled and being hard, difficult to peel and tasting bitter. The Ethrel treated
hands at the same time were of excellent texture, flavour and easy to peel (Fig. 5.7).
After the non-treated fruit were tested 11 days after harvesting, they were overripe
and very soft with the fingers falling away from the crown, but they were still good
to eat. However, there was still a very small amount of green at the distal end of the
peel of fingers. This is in contrast with the fruit initiated to ripen with Ethrel that had
not green in the peel when they were fully ripe and soft (after 4 days) (Fig. 5.8).
Fig. 5.5 Preparing Ethrel for dipping bananas in Thailand

Fig. 5.6 ‘Kluai Hom Thong’ Musa AAA were used in this test at 5.00 pm on 21 March 2019 on
left is a banana hand 1 h after being dipped and on the right a hand that had not been dipped

Fig. 5.7 ‘Kluai Hom Thong’ Musa AAA 25 March 2019 8.00 am 4 days after Ethrel treatment.
The one on the left appeared to be exactly as it was 4 days ago with latex exuding from the skin
when it was peeled and being hard, difficult to peel and tasting bitter. The one on the right was of
excellent texture, flavour and easy to peel
Ethylene 87

Fig. 5.8 ‘Kluai Hom Thong’ Musa AAA harvested 21 March. Not treated to initiate ripening and
stored at about 27–31 °C and 71–86% RH and photographed on 28 March 2019 8.00 am 7 days
after harvest, 29 March 2019 8 days after harvest and 30 March 2019 9 days after harvest, 31
March 2019 10 days after harvest, 1 April 2019 11 days after harvest

Ethrel can contain impurities including 1,2-ethanediylbis (phosphonic acid) and

monochloroethyl ester of (2-chloroethyl)-phosphonic acid (Segall et al. 1991, EPA
1988). The latter may degrade to monochloroacetic acid (EPA 1988). Islam et al.
(2016) reported that both Ethrel and ethylene glycol samples they tested contained
trace amounts of sulphur compounds. Ethrel has several many uses and has been
used commercially to initiate flowering in pineapples, anti-lodging in cereals, stim-
ulation of latex flow in rubber, enhanced degreening of citrus fruits as well as initia-
tion of ripening in climacteric fruit.

Encapsulation

Ho et al. (2011) developed molecular encapsulation of ethylene into α-cyclodextrin

and tested ethylene release from the capsules at 52.9, 75.5 or 93.6% RH and 45, 65,
85 or 105 °C. Kinetics analysis, based on Avrami’s equation (describes the kinetics
of crystallisation), showed that the release of ethylene from the capsules increased
as humidity and temperature were increased. They subsequently investigated using
88 5 Initiation of Ripening

deliquescent calcium chloride and magnesium chloride to release ethylene from an

ethylene-α-cyclodextrin inclusion complexes (IC) powder at humidities between
11.2 and 93.6% RH at 18 °C (Ho et al. 2015). They found that when the IC powder
and deliquescent salts were mixed at a ratio of 1:5, respectively, the CaCl2 and
MgCl2 started to deliquesce at 32.7% RH when the IC powder dissolved in the con-
centrated salt solutions to release ethylene. Increasing the humidity accelerated the
release rate. Maximum release of ethylene gas was achieved after 24 h at 75.5 and
93.6% RH for both IC powder-deliquescent salt mixtures. The deliquescent salts
proved to be a simple option for releasing ethylene gas from the IC powder and is
being marketed as Ripestuff™ (University of Queensland 2017). The powder is an
ethylene-α-cyclodextrin inclusion complex.
When tested on mangoes (Ho et al. 2016) in the laboratory, for in-transit ripening
over two seasons they found that ethylene gas started to be released from the IC
powder in 2 h and complete release was achieved in 24 h. Assessments of fruit
colour and firmness showed that encapsulated ethylene and commercial grade eth-
ylene from pressurised cylinder similarly shortened the ripening time to 9–10 days
(after harvest) for treated fruit as compared with 15 days for untreated mangoes.

Acetylene

Acetylene gas can be used in place of ethylene and has the advantage of being inex-
pensive and readily available in many developing countries. As reported above, eth-
ylene generated from Ethrel can cost 50 times more than acetylene generated from
calcium carbide (Thompson 1985).
Thompson and Seymour (1982), Smith et al. (1986) Smith and Thompson
(1987), tested the effects of acetylene on initiation of ripening of pre-climacteric
‘Giant Cavendish’ bananas. They used pure acetylene, not calcium carbide, and
obtained the required concentrations of acetylene from cylinders of high purity mix-
tures of acetylene in air, specially prepared by British Oxygen Company, at various
concentrations. Acetylene was shown to be effective in initiating ripening of
bananas. Ethylene initiated ripening at 19 °C after exposure for 24 h at 0.01 mL L−1,
but 1 mL L−1 of acetylene was required to give the same effect at that temperature
(Thompson and Seymour 1982). Smith et al. (1986) showed that the effects of acet-
ylene on bananas was proportional to temperature and exposure time (Fig. 5.9)
where exposure of fruit to 1 mL L−1 of acetylene for 4 h did not initiate ripening at
between 20 and 30 °C but it did initiate ripening at 35 °C.
With 8 h exposure to acetylene, fruit were not initiated at 20 °C, were partially
initiated at 25 °C and were fully initiated at 30 °C. Smith et al. (1986) observed an
interaction between the pulp temperature of bananas and their response to acetylene
in that, over a temperature range of 14–35 °C, the higher the temperature the shorter
the exposure time, or the lower the concentration of acetylene was required. The
period of exposure to acetylene was also shown to affect initiation of ripening, with
longer exposures, up to 24 h, requiring a lower concentration of acetylene or a lower
Acetylene 89

Fig. 5.9 Effects of exposure for either 4 or 8 h to 1 L per cubic metre of acetylene at various tem-
peratures on the ripeness of bananas after 6 days at 20 °C. Where a = peel colour score; b = pulp
firmness. Smith et al. (1986)

temperature to initiate ripening (Fig. 5.10). Bananas exposed to acetylene at the

lower temperatures (14–16 °C) were shown to ripen normally with respect to pulp
TSS and pulp texture (rupture force), although the co-ordination of the degreening
response was lost in some fruit exposed to acetylene for 72 h. They found, however,
that the temperature at which bananas were initiated to ripen did not affect their
flavour.
Burg and Burg (1967) reported that acetylene had a lower biological activity than
ethylene, when tested with the pea straight growth test. However, they found that in
order to initiate ripening of bananas acetylene should be applied at 2.8 mL L−1. In
mangoes it was found that at 25 °C and with a 24 h exposure time, at least 1 mL L−1
of acetylene or 0.01 mL L−1 of ethylene was required to initiate ripening (Medlicott
et al. 1987). They also found that treatment with 0.01 mL L−1 acetylene resulted in
limited softening of mangoes but had no effect on the other ripening changes
analysed.

Calcium Carbide

Calcium carbide reacts with water to produce acetylene:

CaC2 + 2 H 2 O ® Ca ( OH )2 + C2 H 2
90 5 Initiation of Ripening

Fig. 5.10 The effects of exposure at 20 °C for 6 days to 1 mL L−1 of acetylene at different tem-
peratures on the ripening of bananas. Source Smith et al. 1986

In many third world countries CaC2 is used as described above. Acetylene is used
commercially to initiate ripening in many less developed countries in the form of
CaC2 because it is cheaper than ethylene sources and easier to apply in ripening
rooms. Calcium carbide is a by-product of the iron and steel industry and the mate-
rial available contains impurities. Technical grade calcium carbide of regular chip
size of 4–7 mm conforming to British standard BS 642 (1965) is the type most com-
monly used. If it were pure calcium carbide then 1 kg would produce 300 L of
acetylene gas. The gas is released when the calcium carbide is exposed to moisture.
The reaction can be violent so the way it is commonly applied is to wrap small
amounts (just a few grams) in twists of newspaper and put these among the bananas
to be ripened (Fig. 5.11). The high humidity reacts with the calcium carbide giving
a slow release of acetylene. Where large quantities of acetylene are required quickly
the small amounts of calcium carbide can be dropped carefully into large buckets of
water. Morton (1987) reported that in 1874 in the Azores it was accidentally discov-
ered that smoke would bring pineapple plants into bloom in 6 weeks and in 1936
acetylene were employed to expedite uniform blooming. Some growers deposited
calcium carbide in the crown of each plant that released acetylene when it got wet.
There is some indication that different varieties of banana require different
amounts of calcium carbide to initiate ripening. Acedo and Bautista (1993) showed
that ‘Latundan’ (Musa ABB) and ‘Saba’ (Musa BBB) bananas exposure of 5 g of
calcium carbide for 1 day “effectively enhanced ripening”. ‘Latundan’ harvested at
full maturity required a lower level of calcium carbide then ‘Saba’ harvested at the
full ¾ stage.
In Thailand farmers buy a small amount of calcium carbide in a local shop (per-
haps 100–150 g) that has been broken down into small packages by the shop owner.
Acetylene 91

Fig. 5.11 Calcium carbide (27 g) wrapped in paper and inserted in a box of ‘Namwa’ bananas
(4.2 kg) to initiate them to ripen in Thailand

Farmers are charged 10 TBT for a package, but it is possible to buy a kg sealed in a
plastic bag for 60 TBT. The farmer then puts a small amount (20–30 g) in a paper
with about 60 banana fingers. This ratio will vary depending on the size of the fin-
gers and whether the carton provides an effective barrier to loss of acetylene, that is,
whether it is well padded with paper and cardboard. The judgement of all these fac-
tors is based on individual farmers’ experience. It is impossible to calculate the dose
of acetylene received by individual fingers, but from observations of bananas that
have been initiated to ripen in this way, it works,

Sensory Analysis

Results reported by Nura et al. (2018) for sensory analysis, show highest sensory
score was recorded for bananas exposed to 25 g CaC2 kg−1 and lowest sensory
score was recorded in fruit that had not been exposed to any chemical for ripening
initiation (control samples). Acceptability was found to increase with increasing in
CaC2 concentration. The results for sensory analysis agreed with the finding of
Sarananda (1990) who recommend the use of acetylene generated from calcium
carbide in ripening of ‘Embul’ bananas. Amarakoon et al. (1999) also recom-
mended the use of 1 g CaC2 kg−1 to ripen bananas. However, Gunasekara et al.
(2015) recorded low sensory scores in ‘Embul’ bananas treated with either CaC2
or Ethrel. In tests in Thailand (Jiraporn and Thompson 2019, unpublished results)
with Food Technologist Academics mainly from Thailand, but also from India and
the UK, bananas from the same bunch of ‘Namwa’ were initiated to ripen with
CaC2 or left to ripen without any chemical at room temperature of about 30 °C. Their
response to the flavour and texture varied with some people preferring one or the
other or others said that they could not see any differences. From this it was con-
cluded that the CaC2 only affected the speed and evenness of ripening but not
sensory characteristics.
92 5 Initiation of Ripening

Toxicity of Calcium Carbide

Industrial grade CaC2 contains about 80% CaC2, 15% calcium oxide and 5% other
impurities (Kirk-Othmer 2004). Great care must be exercised and the operator must
wear protective clothing, including a protective face mask and leave the area imme-
diately after application. Hazard warnings should be put up and flames, cigarettes or
electrical fittings that could cause a spark must be eliminated from the area.
Industrial grade calcium carbide may contain arsenic and phosphorous hydride
(Siddiqui and Dhua 2010). When calcium carbide is applied as a ripening agent,
sulfur, arsenic and phosphorous from calcium carbide may diffuse into the peel and
flesh of fruits and may pose serious health risk (Islam et al. 2016), which appear to
be the main reason for concern about its effect on human health (Chow 1979).
Calcium carbide is not “Generally Recognized As Safe” (Pokhrel 2013) and prohib-
ited in several countries including Sri Lanka, India, Bangladesh and Nepal. Sri
Lanka under the Section 26 of the Food (Labelling and Miscellaneous) Regulation
of 1993. In India use of calcium carbide is strictly banned as per the Prevention of
Food Adulteration Act [Section 44AA]. Prevention of Food Adulteration Rules
1955. This prohibits the use of carbide gas for fruit ripening. Food Safety and
Standards Regulations 2011 prohibit the selling of artificially ripened fruits using
carbide gas. Bangladesh by the Pure Food Ordinance (Amendment) Act 2005,
Formation of National Food Safety Advisory Council (NFSAC); prohibit using cal-
cium carbide, formalin, and pesticides in foods. The penal code of Bangladesh must
penalize any individual selling illegally ripened fruits. The Nepal Food Regulation
2027, prohibits the use of carbide gas for fruit ripening. In India Chandel et al.
(2018) reported that arsine (AsH3) and phosphine (PH3) present in CaC2 as impuri-
ties might find entry in the calcium carbide ripened fruits. CaC2 ripened mangoes
contained harmful arsenic levels between 34.73 and 73.43 ppb as compared to high
residue levels in open market mangoes (106.27 ppb), while fruits ripened without
CaC2 did not contain detectable residues. Further, dipping fruit in 2% Na2CO3 or 2%
Agri-biosoft solution for 12 hr. was effective in reducing the arsenic residue from
71.02 ppb to 6.74–9.05 ppb from fruit surfaces and also removed arsenic from peel
and pulp although the authors do not report the chemical analysis of Agri-biosoft.
However, no information could be found on the residues of arsine and phosphine in
the pulp of bananas after being ripened with calcium carbide. Lakade et al. (2018)
developed a gold nanoparticle colorimetric method to detect residues of arsenic on
fruit surfaces in order to ascertain whether calcium carbide had been used in ripen-
ing the fruit.
Harvey (1928) reported “Ethylene oxide, methylene chloride, ethyl chloride, eth-
ylene chloride, propylene chloride, and amylene are desirable on account of their
physical property of being liquid at ordinary temperatures at comparatively low pres-
sure, but they were found not as effective as ethylene in ripening fruits, and they all
show greater toxicity, shown by the blackening. Amylene, as it is available commer-
cially, has an objectionable odour and produces objectionable flavours in the fruit.”
Alcohol 93

Propylene

Propylene is also called propene or methyl ethylene. Sfakiotakis et al. (1999)

showed that endogenous ethylene production in fruit could be induced by exposing
them to exogenous propylene. McMurchie et al. (1972) applied propylene to
bananas and also found that it increased both their respiration rate and endogenous
ethylene production. Golding et al. (1998) successfully initiated pre-climacteric
‘Williams’ bananas to ripen by exposure them to water saturated air containing
500 mL L−1 propylene for 24 h. Harvey (1928) reported “Tasting squads, including
horticulturists trained in testing fruits for flavour, have agreed that the flavor of
ethylene-treated fruit is superior to that of untreated fruit. Bananas treated with
propylene have a little better flavour than those treated with ethylene”. Stavroulakis
and Sfakiotakis (1997) treated kiwifruit, with 130 μL L−1 propylene in a continuous
flow-through system at 20 °C in combination with different O2 concentration and
found that ethylene biosynthesis and ripening were “stimulated” in the combination
of propylene with 21 kPa O2.

Esters

“The esters-amyl acetate, ethyl acetate, and methyl acetate produce browning and
blackening of the peel without ripening the fruit. Amyl acetate seems more toxic
than methyl or ethyl acetate. The vapor of amyl alcohol was not particularly toxic.
Acetaldehyde produced a brown colour similar to that of fruits ripened without gas”
(Harvey 1928).

Alcohol

Goonatilake (2008) tested ethylene glycol on bananas and several other fruits and
found it could initiate ripening. Their application method was to make up 20% eth-
ylene glycol in water then dip each fruit followed by brushing the solution over the
surface. The main impurities found in commercially available ethylene glycol were
diethylene glycol (3-oxapentane-1,5-diol), triethylene glycol (3,6-dioxaoctane-­1,8-
diol), methanol and aldehydic oxidation products (Marcus 1990). Stahler (1962)
took out a patent that stated “It has been found that the ripening time of green
bananas and citrus fruit can be significantly reduced by maintaining the fruit, after
picking and during storage, in contact with a higher alcohol. The alcohols used
according to this invention can be any of the alkyl alcohols containing between 6-
and 14- carbon atoms, used alone or in combination. For the treatment of green
bananas, lauryl alcohol is preferred.”
94 5 Initiation of Ripening

Carbon Monoxide

CO is a colourless, tasteless, odourless and flammable gas with explosive limits in

air of between 12.5% and 74.2% by volume. It is extremely toxic to humans and
adequate safety precautions must be followed if and when it is used. Plant tissues
can oxidize CO to CO2. CO has been shown to initiate ripening of bananas when
they were exposed to concentrations of 0.1% for 24 h at 20°C (Thompson 1996) and
to “speed up ripening of tomatoes” (Beckles 2012). CO has other uses in posthar-
vest of fruit and vegetables and can have both beneficial and harmful effects. With
levels of O2 between 2 and 5%, CO can inhibit discoloration of lettuce on the cut
butts or from mechanical damage on the leaves. The respiration rate of lettuce was
reduced during a 10-day storage period at 2.5 °C when CO was added to the store,
but low concentrations of CO in the store atmosphere can increase the levels and
intensity of the physiological disorder russet spotting (Table 5.1). Zhang Shaoying
et al. (2015) showed that CO (10 μL L−1) delayed the internal browning of peaches.
The control fruit showed internal browning on the 10th day and thereafter the index
increased rapidly. CO effectively reduced the development and severity of internal
browning by 50%, after 30 days of storage at 8 °C. PAL activities in the CO-treated
fruit, showed a trend of first increased and then decreased, the peak appeared on
15th day. Compared with the control, the treatments with CO, inhibited the increase
of PPO activities. Firmness of the fruit treated with CO were significantly higher
(p ≤ 0.05) compared to the control during 25 days storage. The treatment with CO,
could inhibit the change of PG activity. A combination of 4% O2, 2% CO2 and 5%
CO was shown to be optimum in delaying ripening and maintaining good quality of
mature green tomatoes stored at 12.8 °C after being subsequently ripened at 20 °C
(Morris et al. 1981). In peppers and tomatoes the level of chilling injury symptoms
could be reduced, but not eliminated when CO was added to the store. CO has fun-
gistatic properties especially when combined with low O2. Botrytis cinerea on
strawberries was reduced where the carbon monoxide level was maintained at 5%
or higher in the presence of 5% O2 or lower. Decay in stored mature green tomatoes
stored at 12.8 °C was reduced when 5% CO was included in the storage atmosphere
(Morris et al. 1981). Tomatoes kept in 5 or 10% (v/v) CO with 4% (v/v) O2, had
superior TSS and total acidity profiles compared to those kept in air with less inci-
dence of B. cinerea (Kader 1983).
Figures followed by the same letter were not significantly different (p = 0.05).

Table 5.1 The effect of carbon monoxide on the development of russet spotting on harvested
lettuce. Modified from Kader 1987
Carbon monoxide concentration μL L−1 Russet spotting score
0 0a
80 16c
400 22d
2000 24d
10,000 20d
50,000 6b
Smoking 95

Smoking

Smoking can produce various gases, depending of the combustion material used to
provide the smoke, including acetylene, ethylene and carbon monoxide, all which
can initiate fruit ripening. Smoking is used in many developing countries and a study
in the Sudan showed that it appeared to be effective, but there was considerable varia-
tion in the speed of ripening of the bananas in different parts of the room (Thompson,
unpublished results) Kulkarni et al. (2011) reported that in India smoking was used
but was unpredictable and could result in uneven ripening and poor colour develop-
ment. In Sri Lanka a traditional method was described where bananas are laid in a pit
covered with banana leaves or a sheet and smoke is generated externally, from burn-
ings semi dried leaves, which is then directed into the pit. Narasimham et al. (1971)
compared the traditional practice in India of smoking bunches of bananas for 24–36 h
compared with non-smoking on the speed of ripening. They found that smoking
reduced the time taken for the peel to change from green to fully yellow from 7 days
in non-smoked fruit to 5 days for fruit that had been smoked. The smoking practice
they described was “What we do is this: We dig a pit in earth that is enough to put the
whole banana cluster in. Then, after safely laying the bananas in the pit, we cover up
the pit with a sheet such that only a small hole from a side remains: visualize a small
3–4 inch door to the pit. After that, we light a fire with semi-dry leaves just outside
the pit’s door. (Semi-dry leaves are used to get as much smoke as possible. Dry
leaves do not give that much smoke, because they completely oxidize quickly). And
the smoke is sent through the door by blowing it with the aid of a bamboo. This sends
a good amount of smoke and warms the inside of the pit considerably. And by experi-
ence I can tell you that this makes the bananas to ripen really quickly. I have done a
controlled experiment where half of the cluster was not put into the pit. Bananas in
the pit ripen overnight and the control sample took days to ripen.” Venkata Subbaiah
et al. (2013) tested smoking by burning straw, leaves and cow dung in a closed cham-
ber for 24 h and compared this with Ethrel dipping (1000 ppm). Fruit that had been
exposed to smoking had a higher proportion of firm fruits after ripening than those
that had been initiated to ripen with Ethrel. Mebratie et al. (2015) also compared
ripening initiation with Ethrel and smoking. Smoking resulted in faster ripening but
led to the least marketability, 29% at 10th day of storage, compared to the other treat-
ments that had more than 83% marketability. The reduction of marketability of
smoked bananas was due to blackening and over softening. Gautam and Dhakal
(1993) commented that Nepalese farmers have used their indigenous knowledge in
ripening of banana since time immemorial. In many places in Nepal banana bunches
(‘Malbhog’) were placed in jute sacks and hung over a fire.

Kerosene

Denny (1927) found that the smoke from kerosene lamps contained ethylene and
used the smoke from kerosene lamps to de-green citrus. In some countries, kerosene
burners are used to generate smoke in commercial scale banana ripening rooms and
96 5 Initiation of Ripening

Fig. 5.12 (a) Bananas stacked in heaps or wooded boxed in a ripening room in the Souk in
Khartoum in Sudan and (b) kerosene burners placed in the rooms to initiate ripening in Asmara in
Eritrea. Photographs taken (a) in 1973 and (b) in 2008

a study in the Sudan (Fig. 5.12) showed that it appeared to be effective, but there
was considerable variation in the speed of ripening throughout the room, which was
assumed to be due to inadequate air circulation. Banana bunches were exposed to
smoke generated by burning kerosene stoves inside the chambers for about 24 h. As
a result, the temperature inside the chamber also increases besides evolving ethyl-
ene gas with traces of other gases like acetylene and carbon monoxide (Ram et al.
1979). Other workers have reported that kerosene fumes also contain other gases
including sulphur, aromatic compounds and hydrocarbons (Khan et al. 2013) SO2,
NO, NO3 and CO (Islam et al. 2016).

Incense

In India Venkata Subbaiah et al. (2013) compared smoking with burning incense
sticks and Ethrel dipping in confined rooms on ripening of ‘Grand Naine’ bananas
and found that the incense was ineffective.

Heat

Clearly where fire is used in ripening rooms, along with gases, the temperature will
be elevated, often to high levels in tropical countries where it is commonly applied.
This high temperature could have an additional effect to the gases. However, Giri
et al. (2016) exposed bunches of ‘Cavendish’ at 40 or 45 °C for 30, 45 or 60 mins
Damage and Stress 97

and found that none of the heat treatments hastened the speed of ripening compared
to the control. The higher the temperature and the longer duration of heat exposure,
the slower was the rate of ripening. Kamdee et al. (2018) found that subsequent
ripening of ‘Sucrier’, previously ripened to colour index 3–4, was normal when
exposed 42 °C, even for 24 h, but they found their taste and odour was reduced to
unacceptable levels. Paull and Chen (2000) studied the extent of the alternation of
fruit ripening following heat treatment. They reported that sensitivity or tolerance to
heat stress of a fruit or vegetable is related to the level of heat protective proteins at
harvest and the postharvest production of heat shock proteins. This sensitivity can
be a normal cellular response (less than 42 °C) that can lead to reduced chilling
sensitivity, delayed or slowed ripening and a modification of quality or, at greater
than 45 °C, the pre-stress environmental conditions can modify the cellular response
to stress and cellular recovery.

Damage and Stress

Wounding the banana bunch stalks or fruit may produce ethylene in response to the
wound. Generally, any damage to the fruit tissues induces ethylene biosynthesis.
Bautista (1990) described a simple technique used in Southeast Asia where a stick
is inserted into the stalk of Jackfruit (Fig. 5.13), which not only makes a convenient
handle for carrying but damage to the fruit also initiates it to ripen. Bautista also
mentions that other methods including cutting, scraping or “pinching” papaya,
chico or avocado can hasten ripening.
Ferris et al. (1993) assessed the effects of different types of injury on the speed
of ripening of three plantain genotypes in Nigeria and found some hastening of
ripening in injured plantains (Table 5.2), but results were not consistent. When a
similar experiment was carried out under controlled temperature there were
­consistent increases in the ripening rate due to damage, but only damage caused by
abrasion was consistently significantly (p = 0.05) higher than the controls (Fig. 5.14).

Fig. 5.13 Jackfruit in

Thailand being initiated to
ripen by pushing a stick
into the fruit, before
sending it to market.
Photograph taken in
February 2019
98 5 Initiation of Ripening

Table 5.2 Effects of type of injury on number of days for green plantains, harvested at mature or
fully mature, to be ripe. Modified from Ferris et al. (1993)
Type of damage
Harvest maturity None Impact Abrasion Incision
Fully mature 11.0 9.4 9.9 9.8
Mature 16.2 16.9 13.0 15.1

Yellow LSD(P = 0.05)

6
Peel colour score

Green
1
0 2 4 6 8 10 12
Time after harvest (days)

Fig. 5.14 Effects of different types of damage to ‘False Horn’ plantains on ripening at 20 °C.
□ = control, ∆ = impact, o = abrasion, x = firmness (static pressure) (Modified from Ferris et al.
(1993)

In Brazil Maia et al. (2014) working with ‘Dwarf-Prata’ bananas showed that all
fruits subjected to mechanical injury had increased weight loss, electrolyte leakage,
PPO activity, accelerated peel colour changes and an earlier climacteric peak, com-
pared to controls. The damage caused by abrasion caused higher weight loss. The
starch conversion to sugars in the pulp was affected by impact damage. The impact
and compression damages hastened climacteric ethylene peak and, consequently,
fruit ripening. The impact damage greatly increased PPO and POD activities. Stress-­
induced ethylene biosynthesis was reported by Abeles and Abeles (1972) to be pro-
duced by the same biochemical pathway as that produced during the ripening of
climacteric fruit. Exposing fruit postharvest to low humidity may also result in suf-
ficient stress to the fruit to initiate ripening. This was shown by Thompson et al.
(1974) for plantains. Sfakiotakis et al. (1999) showed that ethylene production was
Fruit Generation 99

induced autocatalytically by exposure to chilling temperatures, wounding and

Botrytis infection in Kiwifruit.
In a comparison of transporting ‘Dwarf Cavendish’ bananas from the field to
ripening rooms in the Sudan, Thompson and Silvis (1975) found that dehanding and
packing the hands in wooden boxes for transport compared with transporting them
as bunches in the traditional way resulted in the ones that had been transported as
hands ripening in 22 days compared to those transported as bunches ripening in
18 days, both at 19–23 °C. This difference was significant (p = 0.05). It was thought
that this shortening of the pre-climacteric period could have been due to the greater
damage inflicted on bunches during transport compared to hands packed carefully
in boxes. Also, because of the lack of transport in rural areas in those days people
would often travel sitting on top of the pile of bunches of bananas, causing more
damage.

Fruit Generation

Fruit that are ripening, and thus giving out ethylene, can be placed in an airtight
room with green fruit. A continuous system could be worked out for commercial
application of this method. However, the room would need to be frequently venti-
lated to ensure there was no build-up of CO2, which is known to inhibit the effect of
ethylene. Kitinoja and Kader (2002) commented that ‘Small-scale wholesalers and
retailers can ripen fruits in bins or large cartons by placing a small quantity of
ethylene-­generating produce such as ripe bananas in with the produce to be ripened.
Cover the bin or carton with a plastic sheet for 24 h, then remove the plastic cover’.
Small trials were carried out by A K Thompson (unpublished) in the 1970s in the
Sudan with bananas for the local market, but the persons operating the ripening
rooms found that using bananas that had begun to ripen to initiate those that had not
to be insufficiently predictable and preferred other methods. A study of fruit being
used to initiate ripening in bananas was carried out in India by Gandhi et al. (2016).
They ripened bananas at ambient temperature in either paper bags or plastic bags in
which were included calcium carbide, an apple, a pear or a tomato. There were 3
bananas in each bag. All the fruit in paper bags took 10 days to ripen, as did the
control in a plastic bag. However, those with calcium carbide ripened in 5 days (for
both 1 and 2 g) while those with an apple ripened in 4 days. Those with a pear rip-
ened in 7 days and those with a tomato in 9 days. Tigist Nardos Tadesse (2014)
stored ‘Dwarf Cavendish’ bananas with avocado fruits and found that the ripening
period of the bananas was reduced without any undesirable effect on their quality.
Pokhrel (2013) also recommended this method and suggested that ripe fruits can be
placed with unripe mature fruits in a ratio from 1:20 in an open environment and in
a ratio of 1:100 in a closed environment to initiate ripening. Ram et al. (2009) mixed
ripe bananas with green bananas and found that the green bananas began to ripen in
3 days and were ready for use in just over 4 days (with an excellent flavour) com-
pared to control fruit that did not begin to ripen until 9 days. Burg (2004) described
100 5 Initiation of Ripening

an experiment where bananas and tomatoes were stored at 14.4 °C, either together
or separately, either under 760 or 80 mm Hg hypobaric pressure. The bananas stored
with tomatoes under 760 mm Hg initiated to ripen because of the ethylene given out
by the tomatoes while the bananas under 80 mm Hg did not initiate to ripen.

Leaves

A one-day treatment of bananas with rain tree leaves (Gliricidia sp.) at 5% of fruit
weight enhanced ripening of ‘Latundan’ (Musa ABB) and ‘Saba’ (Musa BBB)
bananas but was less effective than calcium carbide treatment (Acedo and Bautista
1993). Sogo-Temi et al. (2014) tested exposing fruit to freshly harvested leaves
from two trees, Irvingia gabonesis and Jathropha curcas, and compared their effect
with calcium carbide and potash (there was no definition of exactly the form of
potash used). The calcium carbide and potash were wrapped in polyethene film
before being dropped into a black polythene bag containing the bananas. The I.
gabonesis and the J. curcas leaves were washed in cold water and also placed in
polythene bags containing bananas, while others were placed in the polyethene bags
without any ripening agent and used as the control. The order of ripening of the
bananas was: calcium carbide (3 days) then potash, I. gabonesis and J. curcas and
finally the fruit with no ripening agent ripening in 6 days. Pokhrel (2013) reported
that in some parts of Nepal, Asuro and Dhurseli (Colebrookea oppositifolia) leaves
and fresh rice straw, are also used to ripen bananas using about ten times the amount
of bananas than the amount of leaves and leaving them covered for a week to ripen.
Ram et al. (2009) included dried leaves of Asuro (Adhatoda vesical or Justicia
adhatada) and Koiralo (Bahunia veriagata) in sealed PE bags with bananas (100 g
each per kg of fruits) and found that the bananas began to soften in 4 days, or just
over 4 days, compared to 9 days for controls, and were suitable for consumption
after 6 days. They were considered to have a good flavour. da Costa Nascimento
et al. (2019) ripened ‘Pacovan’ bananas using Bowdichia virgilioides leaves, com-
pared to calcium carbide, following the empirical method used by Borborema farm-
ers in Brazil. They found that bananas exposed to B. virgilioides leaves had a higher
respiration rate and ascorbic acid production and reduced acidity and chlorophyll
compared to those not treated and there were no significant differences from the
bananas that had been initiated to ripen with calcium carbide.
Chapter 6
Ripening Technology

Ripening Rooms

Ripening rooms are designed for climacteric fruit that are harvested before they are
ripe and placed under controlled conditions in order to initiate and control ripening.
Since the 1990s there has been an increasing demand, in many countries, for all the
bananas being offered for sale in a supermarket to be at the same stage of ripeness
so that it has an acceptable and predictable shelf-life. This led to the development of
a system called “pressure ripening”. The system involves the direction of the circu-
lating air in the ripening room being channelled through boxes of fruit so that exog-
enous ethylene is in contact equally with all the fruit in the room. At the same time
the CO2, which can impede ripening initiation, is not allowed to concentrate around
the fruit (Fig. 6.1). In pressurized ripening rooms air is forced though each pallet or
series of pallets before returning to the evaporator. Therefore, any “air-stacking” or
“cross-stacking” of boxes is not necessary, and the result is less handling of the fruit
and improved product quality. In some designs, inflatable bags are placed between
pallets to help direct the air through the boxes of bananas. For non-pressurized
rooms, the boxes of bananas should be “air stacked”. That is, the boxes should be
offset to allow the air to circulate among all the boxes since a non-pressurized room
design may not facilitate the circulating air to pass through boxes but around them,
since the air will take the path of least resistance. In summary the primary require-
ments for ripening rooms are that they should have:
good temperature control
good and effective air circulation
a good system for introducing fresh air
and be gas tight
Before the introduction of pressure ripening, boxes of bananas were removed
from their pallets and stacked in the ripening room so that here was space around

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2019 101
A. K. Thompson et al., Banana Ripening, SpringerBriefs in Food, Health,
and Nutrition, https://doi.org/10.1007/978-3-030-27739-0_6
102 6 Ripening Technology

Fig. 6.1 Pressure ripening rooms (a) and pallets of bananas loaded into a pressure ripening room
(b). Reproduced with permission of Dave Rodden, Advanced ripening technologies Ltd. UK

each box in order to facilitate air circulation (Fig. 6.2). In international trade the
boxes of bananas are lined with polyethylene film (Fig. 6.3) and usually transported
to ripening rooms stacked on pallets. Several systems have been used. One is to
remove each box from the pallet, pull back the plastic film and re-stack them on
pallets so that there is a space between boxes (Fig. 6.2). In other cases, the pallets
are stacked so that one hand hole in each box is facing outwards. As the fruit are
loaded into the rooms the plastic just inside each hand-hold is torn to facilitate air
exchange. This is especially important where fruit have been vacuum-packed.
Air circulation systems are usually largely convectional. Air is blown through the
cooler and then across the top of the store just below the ceiling. The cooled air falls
by convection through the boxes of fruit and is taken up at floor level for recircula-
tion. Many modern ripening rooms have air channels in the floor through which air
is circulated at high pressure. This forces it up through the pallets of boxes and
should give better air circulation. Special devices such as inflatable air bags placed
between pallets are now used to ensure better air circulation and, therefore, more
even fruit ripening.
Good ventilation to enable fresh air to be introduced is very important for suc-
cessful fruit ripening. During the period of initiation of ripening, which is usually
24 h, no fresh air is introduced to the rooms. This is the period when ethylene is
introduced. Directly after this period the rooms must be thoroughly ventilated.
Setting up a ripening system for bananas in Ecuador, Thompson and Seymour
(1984) found that the CO2 level of ripening rooms had gone up from about zero to
7 kPa during this initial 24 h initiation period. Even with a good fan extraction sys-
tem it took 40 min of ventilation to bring the CO2 levels to below 1 kPa. This venti-
lation with fresh air must be repeated every 24 h during subsequent ripening to
prevent levels of CO2 becoming too high. If rooms are not frequently ventilated
ripening can be delayed, or abnormal ripening can occur.
R ipening Rooms 103

Fig. 6.2 Commercial

banana ripening room in
the 1970s

The rooms need to be gas tight in order to ensure that threshold levels of ethylene
are maintained around the fruit during the initiation period. The most common place
where leakage occurs is around the doors. It is, therefore, crucial that special gas
tight doors are fitted and that they have suitable rubber gaskets. These must be
inspected regularly to ensure that they have not been damaged. Gas can also be lost
through the walls of ripening rooms. Commonly these are metal-lined on the inside
with mastic between joints to ensure no gas can pass through them. Gas tight paint
can be used on walls. All holes through the walls for plumbing and electrical fittings
must be blocked with mastic. Safety aspects of rooms have already been mentioned.
All electric fittings and thermostats must be of a special ‘spark-free’ type.
It is advisable to have high humidity of 90–95% RH in ripening rooms. To this end
many rooms are fitted with some humidification device such as a spinning disc humid-
ifier. However, if the rooms are full of fruit and the refrigerant in the cooling coils used
to maintain the room temperature is regulated to within a few degrees of the required
room temperature then this should be sufficient to keep the humidity high.
104 6 Ripening Technology

Fig. 6.3 ‘Cavendish’ bananas, on arrval in Europe, in polyethylene film bags that will give a
modified atmosphere around the fruit

Catalytic Generators (Fig. 5.4) recommended, for ripening rooms, the following
“Banana ripening rooms are very important, not just any room will suffice. A proper
ripening room must have the following:
The room must be as air tight as possible to prevent too much of the ethylene from
leaking out.
The room must be properly insulated to be able to control the temperature within a
few degrees.
The room must have adequate refrigeration. Bananas produce large quantities of
heat when they are ripening. The refrigeration equipment must have the capacity
to accurately control the pulp temperature.
The room may need heating equipment in order to maintain proper room tempera-
ture in cold weather. Electric heating elements have proven the most satisfactory
and are often a part of the cooling system. Open flame type heating should never
be used.
The room must have adequate air circulation. Because uniform pulp temperatures
throughout the load are essential for even ripening, the refrigerated air in the
room must circulate at all times and uniformly throughout the load. The room
should be constructed so that the air flow path from the refrigeration system,
through the load and back to the refrigeration system is unobstructed. Proper air
flow patterns are of the utmost importance.”
Modelling 105

Catalytic Generators also recommend to apply ethylene for a minimum of 24 h dur-

ing the initial phase of the ripening cycle at 100–150 ppm. To achieve this, the
generator setting will depend on the size of the ripening room for example:
Setting 1 for rooms 1600–2500 cubic feet.
Setting 2 for rooms 2500–5000 cubic feet.
Setting 3 for rooms 5000–7500 cubic feet.
Setting 4 for rooms 7500–10,000 + cubic feet.
CO2 concentrations above 1 kPa can retard ripening, delay the effects of ethylene
and cause quality problems. Therefore, it is recommended to ventilate rooms, after
the first 24 h of ripening, by opening the doors for 20 min every 12 h. Other venting
methods are by automatic fan (either timed or sensor-based) or “flow-though” (con-
stant) ventilation.
An example of the current ripening practices of one major company in UK who
ripen 600 to 800 pallets daily was given by Bateman (2019). Each pallet is given a
bar code on arrival. The number from the farm that produced the fruit is written on
each box enabling feedback to growers to ensure quality control. It takes about
6 days from arrival in the ripening facility to despatch to the retailer but it may take
less time depending on growing conditions. On arrival, pallets are tightly packed
together and air is directed through each banana box via the handholds and ventila-
tion holes. On first day, the fruit is allowed to come to the correct temperature for
ripening (16 °C) as opposed to the transport temperature of 13–14 °C. Then on the
second day, ethylene is introduced using catalytic generators. There is a heater and
refrigeration unit at the top of each ripening room and the air is forced down. On day
three the bananas are heated to about 18–19 °C and on day four or five they are
cooled to 13.8–14 °C.

Modelling

Toemmers et al. (2010) described a dynamic model of banana ripening using the
conversion of starch to sugars and based on CO2 emissions. The model was tested
on ripening bananas and was shown to describe the characteristic changes in the
CO2 concentration in the air of the ripening facility and the starch concentration in
the pulp and the peel colour of the bananas. Based on this information, it was pos-
sible, using the model, to estimate the stage of ripeness of the bananas. The model
was also used for automated predictive and adaptive process control in banana rip-
ening facilities using an “Open-Loop-Feedback-Optimal” controller. The starch
level, the CO2 reaction rate and the peel colour change rate were fitted to changing
process courses, which enabled the control function of the banana ripening. The
set-point temperature profile was optimised to automatically control the process to
the required ripening stage and the optimal storage temperature at the end of the
ripening procedure.
106 6 Ripening Technology

Seo and Hosokawa (1983a) and Seo and Hosokawa (1983b), evaluating automa-
tion of banana ripening, confirmed, what had been previously shown, that sugar
content of bananas increased during ripening initiation and this was closely corre-
lated with the cumulative quantity of evolved CO2 by respiration. Thus, a sugar
content of bananas at any moment during ripening could be predicted by knowing
the cumulative quantity of CO2 evolved from bananas and that the cumulative quan-
tity of evolved CO2 at any moment of ripening process could be calculated by CO2
evolution rate curves at the particular temperature schedule. Conversely, it would be
possible to construct a ripening temperature schedule so as to achieve a given value
of cumulative quantity of evolved CO2 or a sugar content of bananas at the termina-
tion of the ripening process. Amano et al. (1993) tried to manage cumulative quan-
tity of evolved CO2 by a conventional control algorithm to control the banana
ripening process, but the stable control of cumulative quantity of evolved CO2 was
not always carried out. With fuzzy control, the experience and the knowledge of
skilled operators, usually expressed qualitatively in words, are applied to control
processes by putting them into fuzzy control rules. Therefore, Seo et al. (1995) con-
sidered fuzzy control to be appropriate to apply for automation of banana ripening.
Fuzzy logic is widely used in machine control and is based on a mathematical sys-
tem that analyses analogue input values in terms of logical variables that take on
continuous values between 0 and 1. Fuzzy control was applied by Seo et al. (1995)
to bananas, which had been initiated to ripen with ethylene, over a 7 day period with
the fuzzy rules of three inputs and one output. The operating conditions for ventila-
tion was based on CO2 concentrations and the deviation of cumulative quantity of
evolved CO2 from the target value. These were considered to reflect the effects of
CO2 and O2 concentrations in the ripening chamber on the CO2 evolution rate of
bananas. Changes in the measured value of cumulative quantity of evolved CO2 in
ripening by fuzzy control, closely followed changes in the target value. Ventilating
operations were effectively conducted to improve the control. Results appeared
encouraging with 6050 mg kg−1 of cumulative quantity of evolved CO2 and 10.1%
increase in TSS content at the end of ripening against the target values of
6030 mg kg−1 CO2 and 11.8 TSS.

Transport

International trade in bananas was first reported to be 500 stems to from Jamaica to
Boston USA in 1866, which took 14 days and the fruit were said to have been sold
at a profit (Sealy et al 1984). There was no indication of temperature control during
this or subsequent shipments in the latter part of the nineteenth century. In 1901 the
vessel Port Morant carried 23,000 stems of bananas from Jamaica to Britain using
mechanical refrigerated holds for the company Elder and Fyffes (Sinclair 1988).
Green banana transport is usually by sea freight at 13-14 °C, 85–95% RH and
25–60 cm of water (cmH) ventilation for about 18–22 days. Using controlled atmo-
spheres for transporting bananas internationally, in order to prevent ripening initia-
Reducing Ripening Initiation in Transit 107

tion, has been used for many years in reefer containers and reefer ships. See section
on controlled atmosphere storage (Chap. 4). Janssen (2014) and Jedermann et al.
(2015) tested the construction of a container prototype that was capable of remote
supervision of the physiological stage of fruit. It had a sensor system comprising of
a set of wireless temperature and humidity sensors to detect local maximum tem-
peratures and hot spots as well as sensors for CO2 and ethylene. Tests were carried
out using a system for automated transport supervision on commercial shipments of
bananas that focused on the detection of “ship ripes” by their higher respiration rate
and ethylene production as well as by the detection of potential hot spots caused by
higher respiration resulting in heat production combined with insufficient air flow.
These tests were carried out during transport of bananas from Costa Rica to Europe
using a simple heat transfer model that allowed estimation of index values for local
cooling effects and respiration rate in each pallet from the measured temperature
curves. They found that the model could predict the risk of hot spots, but the major
obstacle was the large amount of heat generated that was addressed by using differ-
ent modifications in the packing scheme to improve the air flow, which showed clear
benefits with regard to the amount of heat removed by cooling.
Using controlled atmospheres for transporting bananas internationally has been
used for many years. For example, Dr. Errol Reid carried out experiments in 1997
on CA transport of bananas in reefer containers from the Windward Islands to
Britain and subsequently supervised the use of CA reefer ships in 1998. CA reefer
ships were being used for transport of bananas from the Windward Islands to the
UK with some 50,000–60,000 tonnes per year in 2010 often combined with bananas
from the Dominican Republic to reduce the risk of dead-freight. Maersk market CA
containers (called Starcare™) that they claim will extend the green life of bananas
for to up to 50 days (Maersk 2019).

Reducing Ripening Initiation in Transit

For international transport of bananas storage conditions are commonly set at

13.3 °C and 85–95% RH and ventilation at 25 m3 hr.−1, which should give a pre-­
climacteric life of about 4 weeks. An equation describing the relationship between
temperature, ethylene concentration and green life of ‘Cavendish’ bananas was
developed and applied to a shipment protocol of 19 days for bananas exported from
Central America to southern Europe by Wills et al. (2014). The equation predicted
that fruit could be transported without refrigeration if ethylene levels were main-
tained at 0.04 μL L−1 during the winter temperature of 17 °C and at 0.002 μL L−1 at
the summer transport temperature of 24 °C.
Hyperbaric conditions, even at variable room temperatures of up to 37 °C, have
been shown to preserve foods and thus achieve significant energy savings (Fernandes
et al. 2015). Designing shipping containers with the facility of increasing the atmo-
spheric pressure inside each container would be an alternative way of maintaining
bananas in a preclimacteric condition during transport. Hyperbaric storage, at room
108 6 Ripening Technology

temperature, could be more energy efficient that refrigeration since the only energy
costs are during compression and no additional energy is required to subsequently
maintain the product under pressure. However, the capital costs of high-pressure
equipment are high (Saraiva 2014), but expansion of high-pressure food processing
should help to reduce equipment costs (Balasubramaniam et al. 2008). Bartlett
pears in storage at 20 °C had a higher ethylene production rate in 100 kPa O2 than
in air (Frenkel 1975). Similar results were reported by Morris (1981) for mature-­
green and breaker tomatoes stored at 20 °C when they were exposed to 30 or 50 kPa
O2 but exposure to 80 or 100 kPa O2 reduced ethylene production rates and musk-
melons stored in 100 kPa O2 at 20 °C had similar ethylene production level as those
stored in air (Altman and Corey 1987). Zheng et al. (2008) found that zucchini
squash exposed to 60 kPa O2 had a lower ethylene production compared to those
stored in 100 kPa O2 or in air. Hypobaric containers have been tested for interna-
tional transport of fresh food. While hypobaric conditions have been shown to
increase the postharvest life of fresh bananas it has so far not been considered prac-
tical and economic for commercial use in banana transport (Burg 2004; Thompson
2016; see also Chap. 4 Hypobaric Conditions).

Ripening in Transit

Ripening rooms are expensive to build and operate. Since bananas cannot be suc-
cessfully allowed to ripen before harvest and they are often transported over large
distances for protracted periods, it seems logical that at least ripening during trans-
port should be considered as an option. Jedermann et al. (2014, 2015) commented
that ripening of bananas can be carried out directly in the reefer container, used for
transport, after it arrives at its destination. Several companies have developed ripen-
ing systems for bananas in containers during actual transit from the field to the retail
outlet. University of Queensland (2017) commented, referring particularly to man-
goes, that “in-transit ripening is of significant interest to the industry with the oppor-
tunity to substantially reduce time to market and minimize the pressure on large and
expensive ripening room infrastructure”. For in-transit ripening of mangoes using
their IC powder (ethylene-apha-cyclodextrin), Ho et al. (2016) found ethylene at
between 4.9 and 10.5 μL L−1 in the headspace of the containers over 48 h where the
IC powder had been used. Mangoes from the treated containers had a shorter ripen-
ing time by 3–6 days compared to the non-treated control fruit.
Dole Food adopted a technology that ripens bananas in shipping containers. A
briefcase-like metal box containing ethylene canisters is used to release ethylene
gas at a controlled rate for shipping times from 1 to 7 days, so that fruit can be at the
required stage of ripeness on arrival. In this way fruit can be distributed directly to
the retail market without first being taken to a ripening facility.
The American company SmartAir Technology use modified container and refrig-
eration units for in-transit ripening. Two pods are installed at each side of the front
bulkhead of the container with four blowers in two fan pods controlled by a SmartAir
R ipening in Transit 109

control box mounted at the front of the trailer, which also operates the ethylene gas
generator and the fresh air exchange system the unit runs continuously when the
SmartAir system is in use. Discharge air from the refrigeration unit evaporator emp-
ties into the mixing chamber behind the front bulkhead. The discharge mixes with
air in the trailer for precise temperature control, typically within ±1 °F at any point
in the trailer. Most of the mixed air is captured by the fan pods and directed down
the trailer sidewalls instead of flowing above the load as it would in a conventional
refrigerated trailer (Macklin 2001).
A patent was taken out for ripening bananas inside a shipping container in 2013
by Axel Moehrke. The patent was summarized as “To ripen bananas during ship-
ping, unripened bananas are first placed in ethylene-permeable containers within a
shipping box. The boxes of bananas are arranged onto shipping pallets which are
then placed into a shipping container for shipping. The arrangement of the bananas
within the reefer container is such that there is adequate airflow around the fruits to
ensure stable temperature throughout the shipping and ripening process. While
enclosed in the reefer container, during shipping, the bananas are exposed to ethyl-
ene gas at a slow rate over a number of days, to ensure slow, even ripening. The
bananas may be fully or partially ripe upon removal from the shipping container and
are then shipped directly to retail outlets or distribution centers, without the need for
additional ripening in traditional ripening rooms.” (Moehrke 2014).
Chapter 7
Conclusions

Refrigeration technology is continuously improving including improved energy

efficiency. The desire for energy efficiency is a top concern with higher energy sav-
ings with further advances in the mechanical equipment of ripening rooms includ-
ing less heat given off by the mechanical equipment. Technology control of the
atmosphere remotely is progressing with the provision of software that allows
remote access to control ripening rooms, including adjusting alerts, gas parameters
and temperatures through Internet connections.
The maturity of individual banana fingers varies with the climate, environment
and weather during growth, but also within a single bunch. So, if bananas are
allowed to ripen without artificial initiation there will be variation as to when they
reach optimum eating maturity. This may be acceptable or even advantages for a
household, but it not acceptable in most commercial production.
There seems no clear consensus on whether bananas taste better if not initiated
to ripen artificially. The enormous changes in organoleptic properties as fruit ripen
makes it difficult to differentially determine differences and no clear conclusions
can be made from chemical analyses. If the function of ethylene, or any of the other
ways of initiating ripening, have any other function that initiating, again is not clear
and seems unlikely.

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2019 111
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and Nutrition, https://doi.org/10.1007/978-3-030-27739-0_7
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Index

A Auxins, 68
Abscisic acid (ABA), 25, 45, 68, 69 Avrami’s equation, 87
Acetylene
advantage, 88
biological activity, 89 B
calcium carbide, 89–91 Banana Bunchy Top Virus (BBTV), 4
initiation of ripening, 88 Banana Pouyat, 4
proportional to temperature and exposure Bananas (Musa)
time, 88, 89 area of production, 1
sensory analysis, 91 breeding programmes, 4, 5
temperature, 88–90 genotypes (see Genotypes)
toxicity of calcium carbide, 92 international trade, 1
Acyl steryl glycosides, 54 overseas transport and primary production, 1
Agri-biosoft, 92 quantity, 1
Air circulation systems, 102 ripening index, 1, 2
Alcohol, 93 sun damage, 14, 15
Ambul bananas, 20 taxonomy, 2–4
1-Aminocyclopropane-1-carboxylic acid types, 6
(ACC), 28–30 Valery, 2
Aminoethoxy-vinylglycine (AVG), 69, 70 Banana Streak Virus (BSV), 4
Amritsagar, 7 Bananitos, 7
Amylene, 92 Banavac®, 60
Angularity, 20–22 Bautista, 97
Anthracnose, 16 Biotic and abiotic stresses, 67
Antihypertensive principle, 54 Blackened skin, 75
Apem, 6 Black Sigatoka, 4, 15
Apple banana, 6 Bluggoe, 7
Aroma, 46, 47 Bodles Altafort, 7
Arsine (AsH3), 92 Botryodiplodia theobromae, 17
Ascorbic acid, 51 Bowdichia virgilioides leaves, 100
Australimusa, 2 Breeding programmes, 4, 5
Automation, banana ripening, 106 Bunch covers, 19, 20

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2019 137
A. K. Thompson et al., Banana Ripening, SpringerBriefs in Food, Health,
and Nutrition, https://doi.org/10.1007/978-3-030-27739-0
138 Index

C Cucumbers, 14
Calcium carbide, 85, 89–92 Cyanide, 29
Callimusa, 2 Cycloartane triterpenes, 54
Calliper grade, 20–22 Cylinders, 82, 83
Carbohydrates, 48, 49
Carbon monoxide (CO), 94
Caribbean Islands grades, 21 D
Carotenoids, 52, 53 Deliquescent salts, 88
Cascade application, 17 Diazocyciopentadiene (DACP), 68
Catalytic generators, 84, 104, 105 Dielectric constant, 27
Catecholamines, 54 1,1-Diphenyl-2-picrylhydrazyl, 20
Cavendish, 1, 5, 7, 16 Diseases of bananas
Cellulose, 55 anthracnose, 16
Cell walls, 44 Botryodiplodia theobromae, 17
Ceratocystis paradoxa, 17 cascade application, 17
Chemicals Cavendish, 16
ABA, 68 Ceratocystis paradoxa, 17
AVG, 69, 70 field-ripe, 16
DACP, 68 fungicides, 15
gibberellic acid, 67, 68 infection, 15
IAA, 68 international trade, 17
LPE, 69 leaf damage, 15
maleic acid, 70 local marketing, 16
1-MCP, 64–67 packing station environment, 18
metal ions, 64 pitting disease, 17
NO, 70 plastic covering, 17, 18
salicylic acid, 67 Pulpa Crema, 16
Cheni Champa, 7 shipment, 16
Chilling injury, 74–76 Dominico, 7
Chromatographic analytical techniques, 81 Dominico Harton, 7
Cis-butenedioic acid, 70 Dwarf Brazilian, 7
Climacteric fruit Dwarf Cavendish, 3, 7, 19
ABA, 26, 68 Dwarf-Prata bananas, 98
Asian pears and peppers, 25
autocatalytic, 28
in avocadoes, 37 E
biphasic respiratory, 28 Effectively enhanced ripening, 90
development, 25 EIN3 binding F-box protein (EBF), 33
ethylene (see Ethylene) Encapsulation of ethylene, 87, 88
low P, 14 Esters, 93
1-MCP treatment, 65 Etacelasil, 84
pattern of ripening, 25 Ethanol
physiology, 28 concentrations, 58
in respiration rate and starch, 68 Ethephon, 84
Coatings, 71, 72 Ethrel, 84–87
CO2 evolution, 27 Ethyl chloride, 92
Colour index, 1, 2 Ethylene
Commercial banana ripening room, 102, 103 biosynthesis, 25–28
Commercial coatings, 71 catalytic generators, 84
Controlled atmosphere (CA) storage, 32 commercial rooms, 81
effects after ripening initiation, 58, 59 cylinders, 82, 83
pre-climacteric bananas, 57, 58 dependent and independent pathways, 25
Cooked/boiled green, 74 encapsulation, 87, 88
Crown tissue, 16 endogenous, 25, 81
Index 139

ethrel, 84–87 folates, 54

exogenous, 25, 26 genetic analysis, 26
flammable limits, 80 minerals, 47, 48
fruit maturity, 81 moisture, 43, 44
fruit ripening rooms, 80 non-climacteric fruit, 25
generation, 26 peel chemicals, 40, 41
identification, 81 peel colour, 37, 39
internal, 28–30 peel spotting, 40
maturity of fruit and response, 34–37 phenolics, 49, 50
physiological levels, 81 phytochemicals, 54, 55
post ripening initiation, 31, 32 pre-climacteric phase, 26, 27
in pre-climacteric bananas, 57 proteins, 49
production, 25, 27 texture, 44–46
responsive factor, 33 vitamin A, 52, 53
signalling interactions, 26 weight loss, 41, 42
Ethylene absorbents, 62–64 Fungicides, 15, 17
Ethylene chloride, 92 Fuzzy logic, 106
Ethylene glycol, 87
Ethylene oxide, 92
Ethysorb, 62 G
Eumusa, 2 Gamma irradiation, 72
Gamma radiation, 73
Gas parameters, 111
F Gas tight, 103
False Horn plantains, 98 Gene expression, 32
Fertilizers, 13, 14 Genetic analysis, 26
FHIA-01, 7 Genotypes
FHIA-02, 8 cultivar, 6
FHIA-03, 8 in international trade, 5, 6
FHIA-17, 8 Giant Cavendish, 7, 27, 31
FHIA-18, 8 Gibberellic acid, 67, 68
FHIA-21, 8 Goldfinger, 4
FHIA-23, 8 Grand Naine, 8, 85
Field-ripe, 16 Green bananas, 75, 76
Film thickness, 62 Green banana transport, 106
Finger drop, 41 Green Keeper (GK), 62, 63
Flavonoids, 54 Gros Michel, 8
Flavour, 46, 47 Guineo, 8
Folates, 54 Guineo Negro, 8
Fruit generation, 99, 100
Fruit ripening
ABA, 25 H
acidity, 50 Hari chhal, 8
aroma, 46, 47 Harton, 8
ascorbic acid, 51 Harvest maturity
carbohydrates, 48, 49 Ambul bananas, 20
carotenoids, 52, 53 angularity, 20–22
classification, 25 bunches, 20–22
climacteric and non-climacteric, 25 calliper grade, 20–22
composition, 37, 38 fruit and response to ethylene
ethylene (see Ethylene) Cavendish, 35
exposure to exogenous ethylene, 25, 26 colour of the fingers, 35, 36
finger drop, 41 factors, 37
flavour, 46, 47 inhibitors, 34
140 Index

Harvest maturity (cont.) binding site, 30

low concentrations, 34 biosynthesis, 29, 30
position on the bunch, 35, 36 cyanide, 29
taste panel scores, 36 MA-ACS genes, 29
texture, 35, 36 MA-ACS1–9, 29
TSS, 35, 36 metabolic pathway, 28
in international trade, 20 methionine to SAM, 28
irrigation/rainfall, 23 post-climacteric peak stage, 30
and microorganism infection, 23 pulp, 30
plastic bunch cover, 20 respiration rate, 28
quality and ripening, 20 sucrose phosphate synthase activity, 30
recommendations, 21 synthesis, 30
ribbon tagging, 20 International Institute of Tropical Agriculture
Robusta, 23 (IITA), 4
second hand form, 21, 22 International trade, 1, 5, 6, 17, 20, 102, 106
Hazard warnings, 92 Internet, 111
Heat, 96, 97 In-transit ripening, 108–109
Hemiterpenoid glucoside, 55 Irradiation, 72–74
Highgate, 8 Irvingia gabonesis, 100
Hom Khiew, 8, 9
Honey, 8
Horn, 8 J
Humidity, 76–78 Jackfruit, 97
Hyperbaric conditions, 107 Jathropha curcas, 100
Hypobaric storage, 31, 59, 60

K
I Karpuravalli, 8
Incense, 96 Kerosene, 95, 96
Indole-3-acetic acid (IAA), 68 Khai, 8, 9
Ingentimusa, 2 Khuai Hom Khiew Korm, 8
Initiation of ripening Kinetics analysis, 87
acetylene, 88–92 Kluai Hom, 8
alcohol, 93 Kluai Hom Khiew, 8
CO, 94 Kluai Hom Taiwan, 8
comparative effectiveness, 79 Kluai Hom Thong, 8, 86, 87
damage and stress, 97–99 Kluai Hug Mook, 8
disadvantage, 79 Kluai Khai, 9
esters, 93 Kluai Leb Mua Nang, 9, 10
ethylene (see Ethylene) Kluai Nam Wah, 9
Food Technologists in Thailand, 79 Kluai Nark, 9
fruit generation, 99, 100 Kluai Teparod, 9
leaves, 100
local markets, 79
Namwa, 79, 80 L
non-using ethephon, 80 Lady Finger, 9, 10
pre-requisite, 79 Latundan, 9
propylene, 93 Leaves, 100
quality, 80 Lecture tubes, 83
smoking, 95–97 Light and day length, 14, 15
Internal ethylene Local markets, 79
ACC, 28–30 Low Temperature Research Station, 74
ACO activity, 29 Lysophosphatidylethanolamine (LPE), 69
Index 141

M Phenolics, 49, 50
MA-ACS1, 27 Phenylananine ammonia lyase (PAL)
MA-ACS1–9 genes, 29 activity, 67
MADS-box, 32 Phosphine (PH3), 92
Magnapothe grisea, 17 Phytochemicals, 54, 55
Malbhog, 9, 95 Phytotoxicity, 73
Maleic acid, 70 Pisang Ambon Lumut, 11
Mangoes, 31 Pisang Awak, 11
1-MCP treatment, 31 Pisang Lilin, 11
Mechanical damage, 18 Pisang Mas, 11, 58, 60
Metal ions, 64 Pisang Ustrali, 11
Methionine to SAM, 28 PITA 3, 11
1-Methylcyclopropene (MCP), 64–67 PITA 24, 11
Methylene chloride, 92 PITA 314, 11
Methyl ethylene, 93 Pitting disease, 17
Microorganism infection, 23 Plastic bunch cover, 20
Minerals, 47, 48 Polyethylene film, 78
Mitogen-activated protein kinase, 32 Postharvest treatments
Modified atmosphere packing (MAP), 60–63 CA (see Controlled atmosphere (CA)
Moisture, 43, 44 storage)
Musa acuminata, 2 chemicals (see Chemicals)
Musa balbisiana, 2 coating, 71, 72
Musa cavendishii, 6 ethylene absorbents, 62–64
humidity, 76–78
hypobaric storage, 59–60
N irradiation, 72–74
Namwa, 10, 79, 80 MAP, 60–62
Nanica, 10 temperature, 74–76
Nepal banana bunches, 95 Pouyat, 11
Ney Poovan, 10, 13 Poyo, 11
Nitrous oxide (NO), 70 Prata, 11
Non-fungicidal methods, 17 Pre-climacteric bananas, 57, 58
Non-treated fruit, 85 Pre-climacteric phase, 26, 27
Preharvest effects
bunch covers, 19, 20
O disease, 15–18
Open-Loop-Feedback-Optimal controller, 105 fertilizers, 13, 14
Organic production, 14 light and day length, 14, 15
Organoleptic properties, 111 maturity, 20–23
Orishele, 10, 42 mechanical damage, 18
organic production, 14
water stress, 18
P Pressure ripening, 101, 102
Pacha Naadan, 10 Pre-storage treatments, 34
Packing station environment, 18 Prevention of Food Adulteration
Pacovan, 10, 100 Act, 92
Panama disease, 74 Propene, 93
Peel chemicals, 40, 41 Propylene, 93
Peel colour, 37, 39 Propylene chloride, 92
Peel spotting, 40 Proteins, 49
Pei Chiao, 10 Provitamin A carotenoids (pVACs), 52
Pentose phosphate pathway, 73 Pulpa Crema, 16, 38
Petite Naine, 10 Pyricularia grisea, 17
142 Index

R and ethrel, 95
Radiation-induced injury, 73 heat, 96, 97
Rasabale, 11 incense, 96
Red Banana, 11 kerosene, 95, 96
Red Decca, 11 vs. non-smoking, 95
Refrigeration technology, 111 semi-dry leaves, 95
Rhodochlamys, 2 in Sri Lanka, 95
Ribbon tagging, 20 Sodium hydroxide, 85
RipeLock™ technology, 67 Spark-free type, 103
Ripening Starcare™, 107
CA effects after initiation, 58, 59 Starch granules, 44
characterization, 28 Storage humidity, 76
ethylene biosynthesis, 28 Storage temperature, 34
genetic and biochemical pathways, 28 Strawberries, 32
genetic effects, 32–34 Stress, 97–99
initiation (see Initiation of ripening) Succinic dehydrogenase activity, 73
mechanisms, 28 Sucrier, 11, 31, 59
technology (see Technology of ripening) Sucrose-ester coating, 71
Ripening index, 1, 2 Sucrose phosphate synthase
Ripening rooms activity, 30
air circulation systems, 102 Sugar, 11
air-stacking, 101 Sun damage, 14, 15
catalytic generators, 104, 105
CO2 concentrations, 105
commercial banana, 102, 103 T
cross-stacking, 101 Tasting squads, 93
gas tight, 103 Taxonomy, 2–4
good ventilation, 102 Technology of ripening
international trade, 102 modelling, 105, 106
polyethylene film bags, 102, 104 reducing ripening initiation
pressure ripening, 101, 102 in transit, 107, 108
requirements, 101 rooms, 101–105
RH, 103 in transit, 108–109
sale in supermarket, 101 transport, 106, 107
spark-free type, 103 Temperature, 74–76, 111
Robusta, 11, 14, 23 Tetraploids, 4
Texture, 27, 44–46
Thylakoid membranes, 39
S Tomato rin mutants, 32
Saba, 11 Transcription factors, 33
S-adenosyl-methionine (SAM), 28 Transport, 106, 107
Salicylic acid, 67 Triploidy, 3
Santa Catarina Prata, 11 Triterpenes, 54
Sensory analysis, 91 Tryptophan, 54
Serotonin, 54
SmartAir Technology, 108
Smoking U
by burning straw, 95 Under peel discoloration (UPD), 75
developing countries, 95 Unique biphasic respiratory climacteric, 30
Index 143

V W
Vacuum packing, 60 Washing water, 18
Valery, 2, 11 Water stress, 18, 42, 77
Varieties, 6 Weight loss, 41, 42
Ventilating operations, 106
Ventilation, 102
Vitamin A, 52, 53 Y
Volatile compounds, 46 Yellow pulp, 38