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Fruit Ripening: Physiology, Signalling and Genomics

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Fruit ripening is an important aspect of fruit production. The timing of it affects supply chains and buying behaviour, and for consumers ripeness not only affects perceptions of health but has nutritional effects too. Ripeness is closely related to spoilage which has a major financial impact on agricultural industries. Currently there are fast moving developments in knowledge of the factors affecting fruit ripeness, and this up-to-date monograph seeks to draw together the disparate research in this area. The aim of the book is to produce a comprehensive account covering almost every area related to fruit ripening including the latest molecular mechanisms regulating fruit ripening, its impact on human nutrition and emerging research and technologies.

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18 Chapters

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1. Climacteric and Non-climacteric Ripening



Climacteric and Non-climacteric Ripening

Kyoko Hiwasa-Tanase and Hiroshi Ezura*

University of Tsukuba, Tsukuba, Japan

1.1 Introduction: The Role of Fruit

Maturation and Ripening in Plants

Plants have developed a fruit architecture with the characteristics necessary to protect their seeds from the natural environment and disseminate those seeds. Seeds are frequently dispersed by animals as well as by wind and rain. Plants have made their fruit more attractive for surrounding animals by providing sources of energy and nutrition, thereby leading to successful seed spreading and propagation. Thus, fruit formation in plants is a reproductive strategy acquired during evolution, which has led to countless varieties of fruit types throughout the world.

Biologically, fruit is the framework that contains the seed in an angiosperm. Therefore, a fruit mainly consists of an ovary, including partial or whole carpel tissues.

However, the edible part of a fleshy fruit also develops from various floral components, such as the receptacle, sepal and inflorescence. The fruit tissues usually ripen in concert with the maturation of the seed, regardless of their derivation, and attract animals by their colour and aroma.


2. Fruit Ripening: Primary Metabolism



Fruit Ripening: Primary Metabolism

Sonia Osorio1,2 and Alisdair R. Fernie1* für Molekulare Pflanzenphysiologie, Potsdam2

Golm, Germany; IHSM-UMA-CSIC, Universidad de Málaga,

Málaga, Spain


2.1 Introduction

Fruit ripening is a complex and highly coordinated developmental process that yields succulent and flavourful tissues for organisms that consume and disperse the associated seed (Giovannoni, 2001). Ripening involves softening of the fruit tissues to facilitate seed dispersal. In addition to softening, fruits normally exhibit increased accumulation of sugars, acids, pigments and volatile compounds that increase interest and palatability to animals.

Additionally, fruits are an important source of supplementary diet, providing minerals, vitamins, fibre and antioxidants for humans. From an agronomical point of view, nutritional value, flavour, processing qualities and shelf-life determine the quality of fruit. Additional fruit attributes, including early maturity, enhanced colour and increased size, constitute the selection of so-called domestication traits.


3. Cellular, Metabolic and Molecular Aspects of Chromoplast Differentiation in Ripening Fruit



Cellular, Metabolic and Molecular

Aspects of Chromoplast Differentiation in

Ripening Fruit

Jean Claude Pech,1,2* Mondher Bouzayen1,2 and Alain Latché1,2 de Toulouse, Castanet-Tolosan, France; 2INRA, CastanetTolosan, France


3.1 Introduction

Chromoplasts are non-green plastids that are responsible for the yellow, orange and red colours of many fruit. They evolve during fruit ripening by differentiation of other forms of plastids. In a number of fruit, such as tomatoes and peppers, coloured chromoplasts are derived from green chloroplasts with the disintegration of the thylakoid membranes and the formation of new carotenoid-bearing structures (Frey-Wyssling and Kreutzer,

1958; Rosso, 1968). In other fruit, such as the flesh of developing papayas, chromoplasts evolve from leuco- or proplastids, as no intermediate amyloplast or chloroplast structures are encountered

(Schweiggert et al., 2011). A very complex origin of chromoplasts has been found in mango where a dynamic interconversion of plastids occurs, although it was not possible to establish a sequential pattern


4. Cell-wall Metabolism and Softening during Ripening



Cell-wall Metabolism and Softening during Ripening

Mark L. Tucker*

US Department of Agriculture, Beltsville, MD, USA

4.1 Introduction

The final stage of fruit development is ripening, which typically includes transformation from an inedible hard organ into a palatable softer version. Fruit softening is a combination of changes in firmness and texture. Firmness can be defined as compressibility or the force required to deform the surface of the fruit (Brookfield et al.,

2011). Texture is defined as a sensory attribute and is more difficult to measure with instrumentation (Mohamed et al.,

1982; Garcia-Ramos et al., 2005; Brookfield et al., 2011). Texture includes crispness, viscosity and juiciness (Brookfield et al.,

2011). Texture is often best measured by human tasters (Brookfield et al., 2011).

Before moving on to the specific details of what happens during ripening to contribute to softening, let us identify a few basic principles to help visualize what softening really is. The edible parts of fruit are not woody (lignified), not even before they ripen. In other words, the cell walls in the edible parts of fruit are not rigid. They can flex. The flexibility of the cell wall is more obvious in thinner structures like leaves. Leaves are not as ‘stiff as a board’.


5. Aroma Volatiles



Aroma Volatiles

Bo Zhang and Kun-Song Chen*

Laboratory of Fruit Quality Biology, Zhejiang University, Hangzhou,

PR China

5.1 Introduction

Plants are multifaceted chemical factories that produce at least 1000 volatile compounds, and the molecules involved in the biosynthesis and release of these volatiles comprise more than 1% of plant secondary metabolites (Qualley and Dudareva, 2009;

Dicke and Loreto, 2010). Some plants allocate up to 10% of their carbon to the production of volatile secondary metabolites (Firn and Jones, 2006). Several years ago, an argument was raised focusing on the role of volatile compounds released by plants. This argument was presented in an article entitled, ‘Plant volatiles: a lack of function or a lack of knowledge?’

(Pichersky et al., 2006). In recent years, thanks to the development of integrative biological approaches, more and more papers regarding volatiles have been published, giving rise to a molecular understanding of the function of volatiles. In

March 2010, a special issue of Trends in


6. Making the Surface of Fleshy Fruit: Biosynthesis, Assembly and Role of the Cuticular Layer



Making the Surface of Fleshy Fruit:

Biosynthesis, Assembly and Role of the

Cuticular Layer

Justin Lashbrooke,1,2,3 Fabrizio Costa2 and Asaph Aharoni1*

1Department of Plant Sciences, Weizmann Institute of Science,

Rehovot, Israel; 2Research and Innovation Centre, Fondazione Edmund

Mach, TN, Italy; 3Institute for Wine Biotechnology, Stellenbosch

University, Stellenbosch, South Africa

6.1 Introduction to the Plant Cuticle

The primary barrier between the atmosphere and the aerial parts of higher plants is the cuticular membrane or the cuticle.

The constituents of this hydrophobic extracellular membrane, typically comprised of soluble waxes and polymerized lipids, are produced and secreted by the plant’s epidermal cells (Kunst and

Samuels, 2003; Pollard et al., 2008).

Lipids consisting mostly of C16 to C20 fatty acids are polymerized to form a matrix known as cutin (Schreiber, 2010).

The cutin matrix is both embedded with waxes (intracuticular) and covered with a thin layer of surface (epicuticular) waxes


7. Antioxidants and Bioactive Compounds in Fruits



Antioxidants and Bioactive Compounds in Fruits

Angelos K. Kanellis1* and George A. Manganaris2 of Pharmaceutical Sciences, Aristotle University of

Thessaloniki, Thessaloniki, Greece; 2Department of Agricultural

Sciences, Biotechnology and Food Science, Cyprus University of

Technology, Lemesos, Cyprus


7.1 Introduction

Quality is determined based mainly on external appearance, background colour and fruit size, whilst flavour, taste and aroma perception occur during or following consumption (Struik et al., 2005).

Fruits are also considered to be beneficial sources of antioxidant potency due to their phytochemical properties. Numerous bioactive compounds are theorized to have a role in preventing or ameliorating various chronic human diseases such as cancer, coronary vascular disease, Alzheimer’s disease and diabetes. Metabolic pathways are not completely understood and as-yetundefined non-antioxidant mechanisms may be responsible. Nevertheless, the consumer cannot perceive such attributes.


8. Vitamins in Fleshy Fruits



Vitamins in Fleshy Fruits

Pierre Baldet, Carine Ferrand and Christophe Rothan*

INRA, Villenave d’Ornon, France; and University of Bordeaux,

Villenave d’Ornon, France

8.1 Introduction

Grains (rice, wheat, maize) and tubers

(potato, cassava) constitute the staple foods in most human diets. They provide major nutrients required for sustaining a healthy and productive life in humans, such as polysaccharides, lipids and proteins.

Cereal and tuber-based diets, however, need to be diversified by the addition of a variety of foodstuffs, such as legumes, vegetables and fruits, which will add micronutrients to the staple food. Micronutrients are essential dietary elements required in very small quantities and not synthesized by humans. They comprise minerals (e.g. selenium, zinc) and vitamins, deficiencies of which are responsible for numerous human diseases.

The crucial role in the human diet of fresh fruits and vegetables for preventing various diseases has been known for centuries. The well-known example of the discovery of the role of L-ascorbic acid, or vitamin C, illustrates how the link between a diet including fleshy fruits, the prevention of scurvy and the intake of vitamin C was made. Scurvy is a fatal disease that frequently affected sailors on long voyages, or soldiers, whose diet consisted mostly of bread, meat or beans.


9. Polyphenols




Agnès Ageorges, Véronique Cheynier* and Nancy Terrier

INRA, Montpellier cedex, France

9.1 Introduction

Polyphenols are a large class of plant secondary metabolites, ubiquitous in plants and structurally diverse. The earlier definition of polyphenols, proposed by

Bate-Smith and Swain (1962), implied the ability to precipitate alkaloids and proteins from solution, while many recent papers refer to all phenolic compounds as polyphenols. In fact, the term polyphenols should be restricted to plant phenolic compounds ‘derived exclusively from the shikimate derived phenylpropanoid and/or the polyketide pathway(s), featuring more than one phenolic ring and being devoid of any nitrogen-based functional group in their most basic structural expression’, as stated recently by Quideau et al. (2011).

This definition covers several groups, including flavonoids, hydroxystilbenes, lignans and benzoic acid derivatives such as gallotannins and ellagitannins. Wide structural diversity is encountered within each group, and especially the flavonoid family, comprising over 8000 molecules


10. Ethylene Biosynthesis



Ethylene Biosynthesis

Donald Grierson*

Laboratory of Fruit Quality Biology/The State Agriculture Ministry

Laboratory of Horticultural Plant Growth, Development and Quality

Improvement, Zhejiang University, Hangzhou, China; Plant and Crop

Sciences Division, School of Biosciences, University of Nottingham,

Loughborough, UK

10.1 Introduction

Research during the first few decades of the 20th century showed that hydrocarbon gases in the environment influence plant growth, development and fruit ripening.

Once it was realized that ethylene was the key molecule in this process, and that plants produce it themselves, it was recognized as a bona fide hormone. This stimulated interest in determining the pathway of ethylene biosynthesis and led, ultimately, to the discovery of the enzymes, genes and regulatory factors that control ethylene production and action at different stages in the life cycle. All plants produce ethylene, but increased ethylene production occurs at many stages of development, particularly in response to developmental signals (e.g. flower development and sex determination, abscission, fruit ripening, leaf senescence), hormones


11. Ethylene Perception and Signalling in Ripening Fruit



Ethylene Perception and Signalling in

Ripening Fruit

Rahul Kumar and Arun K. Sharma*

Department of Plant Molecular Biology, University of Delhi South

Campus, New Delhi, India

11.1 Introduction

Fruits are an indispensable part of the human diet as they are a rich source of vitamins, minerals, antioxidants, sugars,

fibres and flavour compounds. Accumulation of the majority of these chemicals/ products starts once ripening commences in fruits. The triggering of the ripening initiates a phase change in fruit development, which is accompanied by a dramatic shift in primary and secondary metabolism. These coordinated changes result in the accumulation of carotenoids, flavour and volatile components, antioxidants and sugars in the ripening fruits and add to the nutritional quality of the fruits. In plants, the bright colours and complex aromas act as attractants and help in seed dispersal, while in humans these pigments with antioxidative properties are involved in prevention of certain diseases, such as cancer and heart attack (Fraser and Bramley,


12. Other Hormonal Signals during Ripening



Other Hormonal Signals during


Christopher Davies* and Christine Böttcher

CSIRO Plant Industry, Glen Osmond, SA, Australia

12.1 Introduction

Ask any plant biologist which hormone is involved in fruit ripening and the answer will almost inevitably be ‘ethylene’. The role of ethylene during fruit development has been much discussed, and the case for it being pivotal in climacteric ripening is well established (see Grierson, Chapter 10, and Kumar and Sharma, Chapter 11, this volume). This simple molecule has dominated the research effort into the control of fruit ripening. This is partly because of its rather obvious effects on the ripening of some fruit and partly because it coordinates the ripening of many commercially important fruits that can also serve as model species for study, such as tomato. However, ethylene is far from being the only hormonal influence on fruit ripening. There is increasing interest in other hormones that deserve our attention with regard to the control of ripening in both climacteric and non-climacteric fruits.


13. Genetic Diversity of Tropical Fruit



Genetic Diversity of Tropical Fruit

Surendra Kumar Malik, Susheel Kumar and Kailash C. Bansal*

National Bureau of Plant Genetic Resources, Pusa Campus,

New Delhi, India

13.1 Introduction

Botanically, a true fruit is a mature ovary; however, other flower and inflorescence parts also form a part of the fruit in some taxa. There is a vast diversity of fruits in angiosperms. Fruits are classified based on their morphology and development: simple

(fruit from a single ovary); accessory (fruit from inferior ovary); aggregate (fruit from several separate ovaries); and multiple

(fruit from several independent flowers).

Simple fruits can be dry or fleshy and result from the ripening of a simple or compound ovary with only one pistil. Dry fruits may be dehiscent (opening to discharge seeds) or indehiscent (not opening to discharge seeds). Ecological parameters and the habitat of a plant play an important part in the fate of a fruit, facilitating the reproductive mechanism, dissemination of seeds and eventually propagation of the species.


14. Natural Diversity and Genetic Control of Fruit Sensory Quality



Natural Diversity and Genetic Control of

Fruit Sensory Quality

Bénédicte Quilot-Turion and Mathilde Causse*

INRA, Unité de Génétique et Amélioration des Fruits et Légumes,

Domaine Saint-Maurice, Montfavet Cedex, France

14.1 Introduction

Fruit sensory quality has only recently become a target for breeders. Due to consumer dissatisfaction relating especially to fruit flavour, genetic improvement of this quality is now required (Ulrich and

Olbricht, 2011). Fruit sensory quality is a complex trait that contributes a combination of flavour and texture components, together with general fruit appearance attributes. Most sensory traits are difficult to measure by methods other than sensory analysis. However, some of the major components of flavour and texture such as sweetness, sourness or fruit

firmness can be assessed by physical or chemical measurements (Baldwin et al.,

1998). The complexity of fruit quality (due to the number of parameters to take into account, their polygenic inheritance and their multiple interactions) and generation length for fruit trees has limited genetic progress. Today, molecular markers enable dissection of the genetic basis of complex traits, and our increasing knowledge about the genomes offer new and efficient tools to breeders.


15. Ripening Mutants



Ripening Mutants

Cornelius S. Barry*

Department of Horticulture, Michigan State University, MI, USA

15.1 Introduction

Fleshy fruits are botanically and chemically diverse, yet ripening processes are surprisingly conserved and often include changes in colour and cell-wall dissolution, together with subsequent fruit softening, the synthesis of aroma compounds and conversion of starch to sugars.

These changes increase palatability and help to signal seed maturity, facilitating dispersal by frugivores. Due to the importance of fleshy fruits in providing sources of sugars, vitamins, minerals, antioxidants and fibre to the human diet, considerable research effort has focused on identifying the processes, enzymes and regulatory proteins that contribute to the development and ripening of fleshy fruits yet limit their postharvest deterioration. In recent years, the development of genomics resources for fleshy fruit-bearing species, including available genome sequences, large expressed sequence tag collections and publically available gene expression data, have greatly increased our understanding of the genes correlated with events that occur during the ripening process. However, despite the development of these resources, functional analyses of putative ripening-related genes is the main factor limiting understanding of the ripening process. While gene-silencing approaches are useful in some fruit crop species, some of the most significant


16. Biotechnology of Fruit Quality



Biotechnology of Fruit Quality

Avtar K. Handa,1* Raheel Anwar1,2 and Autar K. Mattoo3

1Department of Horticulture and Landscape Architecture, Purdue

University, West Lafayette, IN, USA; 2Institute of Horticultural Sciences,

University of Agriculture, Faisalabad, Punjab, Pakistan; 3Sustainable

Agricultural Systems Laboratory, USDA-ARS,Beltsville Agricultural

Research Center, Beltsville, MD, USA

16.1 Introduction

Fruit and vegetable crops are the major dietary source of vitamins, antioxidants and minerals and have the potential not only to ameliorate physiological disorders but also to decrease the incidence of human diseases such as cancer. Consequently, consumption of fruits and vegetables has increased in recent years, further increasing their global demand. Consumers expect good-quality fruit to be flavourful, succulent, juicy and nutritional, in addition to being attractive in size and appearance.

Other consumer-desirable characteristics of fruits include crispness, chewiness and oiliness. However, for the fruit handler, shipper and retailer, the desirable fruit quality attributes include being less prone to handling and shipping damages, slow softening during storage and longer shelflife, without affecting consumer appeal.


17. Insights into Plant Epigenome Dynamics



Insights into Plant Epigenome Dynamics

James Giovannoni*

US Department of Agriculture and Boyce Thompson Institute for Plant

Research, Cornell University, Ithaca, NY, USA

17.1 Introduction

Genetic information is housed and passed to subsequent generations via the DNA code. The epigenome provides additional information, context and regulatory constraint in addition to both transient and long-term genetic memory. The epigenome consists of information carried in the nature of chromatin packaging and organization, histone modifications (e.g. acetylation, methylation) and DNA (specifically cytosine) methylation. Specific genes involved in DNA or protein methylation, acetylation, small RNA (sRNA) processing and sRNA transcription contribute to epigenome architecture, and their mutations have provided opportunities to develop insights into the intricacies of the epigenome. Advanced sequencing and informatics capabilities permit genomescale analyses at modest cost, resulting in a wealth of data on epigenomes, and their variation and dynamics in response to development and external stimuli.


18. Functional Genomics for the Study of Fruit Ripening and Quality: Towards an Integrative Approach



Functional Genomics for the Study of

Fruit Ripening and Quality: Towards an

Integrative Approach

Federico Martinelli1,2 and Abhaya Dandekar3*

1Dipartimento di Scienze Agrarie e Forestali, University of Palermo,

Palermo, Italy; 2Istituto Euro Mediterraneo di Scienza e Tecnologia,

Palermo, Italy; 3Department of Plant Sciences, University of

CaliforniaDavis, CA, USA

18.1 Introduction

Fruit development is controlled by genetically programmed processes influenced by environmental factors. Different ‘omics’ approaches (deep sequencing, microarray analysis, suppression subtractive hybridization) have identified and characterized genes involved in this process in several fruit species. The mass of knowledge concerning transcriptional regulatory networks affecting important physiological and developmental processes has expanded in the last two decades.

Expressed sequence tag (EST) sequencing uses microarray technology and realtime PCR to generate comprehensive data for functional genomics studies. Following the pioneering work of Aharoni and coworkers (2000) on strawberry, microarrays have been used in many different fruit species. In tomato, large-scale EST sequencing projects have clarified molecular mechanisms of fruit ripening and identified important transcription factors



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