**Meet the editors**

Jaya R. Soneji, PhD, works in the areas of plant breeding, genomics, bioenergy, genetic engineering, population, and eco-evolutionary genetics. She is the author of peer-reviewed research articles, book chapters, and popular articles and has guest-edited special issues for journals, edited books, and newsletters. She was an adjunct faculty at Polk State College, USA. Her work has been

broadcasted on Fox News, USA. She was invited by CBC Radio, Canada, to speak on air. She has served in the "Executive Committee" of GII. She was recognized as "Young Scientists" by Bioclues in 'Member in Spotlight' of GII and featured in ASPB News. The University of Florida's International Programs appraised her contribution in "International Focus." She has peer-reviewed manuscripts for prominent international journals and grant proposals for international institutions. Dr. Soneji obtained her BSc degree (rank second) and MSc degree (rank second) from the University of Mumbai and her PhD degree from BARC, India. She has received many scholarships and awards for her academic achievements. She taught plant genetics, physiology, tissue culture, and genetic engineering at Ramnarain Ruia College, India.

Madhugiri Nageswara-Rao, PhD, works in the areas of plant breeding, genomics, bioenergy, genetic engineering, disease diagnostics, population, and eco-evolutionary genetics. He is the author of peer-reviewed research articles, book chapters, and popular articles, and has guest-edited special issues for journals, edited books, and newsletters. He was an adjunct faculty at Polk State

College, USA. His work has been broadcasted on Fox News, USA. He was invited by CBC Radio, Canada, to speak on air. He has served in the "Executive Committee" of GII. He was recognized as 'Young Scientists' by Bioclues, in "Member in Spotlight" of GII and featured in ASPB News. The University of Florida's International Programs appraised his contribution in "International Focus." He has peer-reviewed manuscripts for prominent international journals and grant proposals for international institutions. Dr. Rao obtained his BSc and MSc degrees from Bangalore University and his PhD degree from FRI, India. He was featured in "Tomorrow's Principal Investigators: Rising Young Investigators" by Genome Technology, USA. He secured a "silver award" as a team member from the American Museum of Natural History, USA. He was also selected for AAAS/Science Program for Excellence in Science. He has served on the editorial boards of six journals.

Contents

**Preface VII**

**Section 1 Breeding and Genetics 1**

**Breeding 63** Boris Krška

**Section 2 Health Benefits 83**

Chapter 1 **Gojiberry Breeding: Current Status and Future Prospects 3** Jianjun Chen, ChihCheng T. Chao and Xiangying Wei

Chapter 3 **Peach Breeding Studies in Turkey and the Evaluation of Peach**

Ayzin Baykam Kuden, Songul Comlekcioglu, Kadir Sarier,

Turgut Yesiloglu, Berken Cimen, Meral Incesu and Bilge Yilmaz

**Phytochemicals and Functional Genomic Characteristics 85** Juan C. Castro, J. Dylan Maddox, Marianela Cobos and Sixto A. Imán

Jian Ding, Chengjiang Ruan, Ying Guan and Susan Mopper

Chapter 2 **Genetic Diversity and Breeding of Persimmon 21**

**and Nectarine Hybrids 47**

Burhanettin Imrak and Ali Kuden

**Hazelnut by GC-TOF/MS 117**

Chapter 7 **Clinical Applications of Pomegranate 127**

Sally Elnawasany

Chapter 4 **Genetic Apricot Resources and their Utilisation in**

Chapter 5 **Myrciaria dubia "Camu Camu" Fruit: Health-Promoting**

Chapter 6 **Qualitative and Quantitative Assessment of Fatty Acids of**

## Contents

#### **Preface XI**



Preface

tus and future prospects of fruit and nut crops.

The fruit and nut crops are laden with health benefits. As people are becoming more con‐ scious about their health and nutritional uptake, the worldwide demand and consumption of fruit and nut crops are steadily increasing. This has made it hard to keep pace between the rate of fruit and nut production and its consumption. To meet this increasing demand, there is a need to produce improved, better yielding, and high-quality fruit and nut crops. This book intends to provide the reader with a comprehensive overview of the current sta‐

Over the past decade, the standard of living has greatly improved and has led to an in‐ creased demand for fruits and nuts. It has led to an interest in consuming fruits and nuts with health-promoting phytochemicals. A lot of work is being done in the field of fruit and nut breeding, biotechnology, and genomics to produce varieties with traits of interest. Breeding of fruits and nuts has been enhanced by advanced technologies such as marker-

The chapters of this book discuss details of the botanical characteristics, origin and domesti‐ cation, genetic diversity, genetic resources, breeding, genetic improvement, biotechnology, genomics, effects of health-promoting phytochemicals, and edible oils or clinical applications of the fruit and nut crops. Such comprehensive information covered in this book will directly enhance both basic and applied research in fruit and nut crops and will particularly be useful

Department of Entomology, Plant Pathology, and Weed Science

**Dr. Jaya R. Soneji**

Las Cruces, USA

Las Cruces, USA

Department of Biology New Mexico State University

New Mexico State University

**Dr. Madhugiri Nageswara-Rao**

assisted selection, genome-wide association studies, and genomic selection.

for students, scientists, researchers, teachers, breeders, policy-makers, and growers.

## Preface

The fruit and nut crops are laden with health benefits. As people are becoming more con‐ scious about their health and nutritional uptake, the worldwide demand and consumption of fruit and nut crops are steadily increasing. This has made it hard to keep pace between the rate of fruit and nut production and its consumption. To meet this increasing demand, there is a need to produce improved, better yielding, and high-quality fruit and nut crops. This book intends to provide the reader with a comprehensive overview of the current sta‐ tus and future prospects of fruit and nut crops.

Over the past decade, the standard of living has greatly improved and has led to an in‐ creased demand for fruits and nuts. It has led to an interest in consuming fruits and nuts with health-promoting phytochemicals. A lot of work is being done in the field of fruit and nut breeding, biotechnology, and genomics to produce varieties with traits of interest. Breeding of fruits and nuts has been enhanced by advanced technologies such as markerassisted selection, genome-wide association studies, and genomic selection.

The chapters of this book discuss details of the botanical characteristics, origin and domesti‐ cation, genetic diversity, genetic resources, breeding, genetic improvement, biotechnology, genomics, effects of health-promoting phytochemicals, and edible oils or clinical applications of the fruit and nut crops. Such comprehensive information covered in this book will directly enhance both basic and applied research in fruit and nut crops and will particularly be useful for students, scientists, researchers, teachers, breeders, policy-makers, and growers.

> **Dr. Jaya R. Soneji** Department of Entomology, Plant Pathology, and Weed Science New Mexico State University Las Cruces, USA

> > **Dr. Madhugiri Nageswara-Rao** Department of Biology New Mexico State University Las Cruces, USA

**Section 1**

**Breeding and Genetics**

## **Breeding and Genetics**

**Chapter 1**

**Provisional chapter**

**Gojiberry Breeding: Current Status and Future**

**Gojiberry Breeding: Current Status and Future** 

DOI: 10.5772/intechopen.76388

Goji, gojiberry, or wolfberry is the fruit of *Lycium barbarum* L., *L. chinense* Mill., or *L. ruthenicum* Murr. in the family Solanaceae Juss. The fruit is bright orange-red or black and is edible with a sweet and tangy flavor. Gojiberry is rich in polysaccharides, flavonoids, carotenoids, betaine, kukoamine A, sitosterol, and other compounds which have antioxidant, anti-inflammatory, and anti-neoplastic properties and have been used for the treatment of various blood circulation disorders and diabetes. Recently, there is an increased demand for high-quality gojiberry and its products because they are considered a superfruit. China is the main producer and supplier of gojiberry in the world. Thus far, limited information is available about genetic resources, breeding activities, and major cultivars of gojiberry. This chapter is intended to review the current knowledge on gojiberry germplasm resources and their relationships as well as to describe gojiberry breeding activities. Future prospects on gojiberry cultivar development are also

**Keywords:** gojiberry, *Lycium barbarum*, *Lycium barbarum* polysaccharides (LBPs), *Lycium* 

Gojiberry generally includes *Lycium barbarum* L., *L. chinense* Mill., and *L. ruthenicum* Murr. (**Figure 1**). They are deciduous woody shrubs, often thorny, spiny, and growing from 1 m to 4 m in height [1]. Stems are slender, erect or spreading, often scrambling. The leaves are gray green, fleshy, ovate, lanceolate, or subcylindric shaped and are alternately arranged,

> © 2016 The Author(s). Licensee InTech. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

© 2018 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use,

distribution, and reproduction in any medium, provided the original work is properly cited.

**Prospects**

**Prospects**

Xiangying Wei

Xiangying Wei

**Abstract**

discussed.

**1. Introduction**

Jianjun Chen, ChihCheng T. Chao and

Jianjun Chen, ChihCheng T. Chao and

http://dx.doi.org/10.5772/intechopen.76388

Additional information is available at the end of the chapter

Additional information is available at the end of the chapter

*chinense*, *Lycium ruthenicum*, Solanaceae, wolfberry

#### **Gojiberry Breeding: Current Status and Future Prospects Gojiberry Breeding: Current Status and Future Prospects**

DOI: 10.5772/intechopen.76388

Jianjun Chen, ChihCheng T. Chao and Xiangying Wei Jianjun Chen, ChihCheng T. Chao and Xiangying Wei

Additional information is available at the end of the chapter Additional information is available at the end of the chapter

http://dx.doi.org/10.5772/intechopen.76388

#### **Abstract**

Goji, gojiberry, or wolfberry is the fruit of *Lycium barbarum* L., *L. chinense* Mill., or *L. ruthenicum* Murr. in the family Solanaceae Juss. The fruit is bright orange-red or black and is edible with a sweet and tangy flavor. Gojiberry is rich in polysaccharides, flavonoids, carotenoids, betaine, kukoamine A, sitosterol, and other compounds which have antioxidant, anti-inflammatory, and anti-neoplastic properties and have been used for the treatment of various blood circulation disorders and diabetes. Recently, there is an increased demand for high-quality gojiberry and its products because they are considered a superfruit. China is the main producer and supplier of gojiberry in the world. Thus far, limited information is available about genetic resources, breeding activities, and major cultivars of gojiberry. This chapter is intended to review the current knowledge on gojiberry germplasm resources and their relationships as well as to describe gojiberry breeding activities. Future prospects on gojiberry cultivar development are also discussed.

**Keywords:** gojiberry, *Lycium barbarum*, *Lycium barbarum* polysaccharides (LBPs), *Lycium chinense*, *Lycium ruthenicum*, Solanaceae, wolfberry

#### **1. Introduction**

Gojiberry generally includes *Lycium barbarum* L., *L. chinense* Mill., and *L. ruthenicum* Murr. (**Figure 1**). They are deciduous woody shrubs, often thorny, spiny, and growing from 1 m to 4 m in height [1]. Stems are slender, erect or spreading, often scrambling. The leaves are gray green, fleshy, ovate, lanceolate, or subcylindric shaped and are alternately arranged,

Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. © 2018 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

© 2016 The Author(s). Licensee InTech. This chapter is distributed under the terms of the Creative Commons

renowned Yin strengthening agent, and the root bark, known as "Di Gu Pi," is a cooling agent [3]. Traditional English vernacular names include boxthorn, fructus lycii, wolfberry, Chinese wolfberry, or matrimony vine [4]. Since the beginning of the century, the plant has been com-

Gojiberry Breeding: Current Status and Future Prospects http://dx.doi.org/10.5772/intechopen.76388 5

The berries harvested from August to October are eaten as fresh fruit, dehydrated to make dried fruit or soaked in liquor to produce *Lycium* juice. As a medicinal food, it is used as a condiment with steamed rice. Young soft leaves can also be used as a vegetable. Gojiberry has been used for its anti-aging activities, tranquilizing and thirst-quenching effects, and its ability to increase stamina [5]. Consuming gojiberry has been shown to improve health-related problems, such as diabetes, hyperlipidemia, cancer, hepatitis, immune disorders, thrombosis,

Scientific analysis of gojiberry constituents started in the 1970s. Qian et al. [9] reviewed 142 publications from 1975 to 2016 and summarized that at least 355 compounds occur in different species of *Lycium*, which were categorized as alkaloids at 20%, sterols, steroids, and their derivatives 16%, terpenoids 11%, amides 10%, flavonoids 9%, organic acid 9%, phenylpropanoids 9%, glycerogalactolipids 6%, ligans 4%, coumarins 3%, antraquinones 1%, peptides

The most intensively studied components are a group of water-soluble glycoconjugates, which are identified as arabinogalactan-proteins, commonly known as *L. barbarum* polysaccharides or LBPs. It is estimated that dried fruits comprise 5–8% of LBPs [10, 11]. Chinese Pharmacopeia [12] recommended using LBPs as an indicator for evaluation of gojiberry quality. Molecular weights of the LBPs range from 24 to 241 kDa, and they are mainly composed of six types of monosaccharides: arabinose, glucose, galactose, mannose, xylose, and rhamnose [13]. LBPs also contain galacturonic acid and 18 amino acids and share a glycan-o-ser glycopeptide structure [14]. The main chains of the glycan backbones of LBPs have been reported

A second major group of metabolites in fruit is carotenoids [3], the content of which increases during the fruit ripening process. A total of 11 free carotenoids and 7 carotenoid esters were detected from unsaponified and saponified *L. barbarum* extracts [16]. Zeaxanthin dipalmitate was found to account for 80% of the total carotenoids [17]. β-Cryptoxanthin palmitate, zeaxanthin monopalmitate, small amounts of free zeaxanthin and *β*-carotene are also present [3]. Seeds contain zeaxanthin (83%), *β*-cryptoxanthin (7%), *β*-carotene (0.9%), and mutatoxanthin

The fruits also consist of vitamins including ascorbic acid (vitamin C), riboflavin, and thiamin. The content of vitamin C (42 mg/100 g) is comparable to that of fresh lemon fruits [3].

to be either alpha-(1–N6)-D-glucans or alpha-(1–N4)-D polygalacturonans [3, 15].

monly called Goji, an appellation derived from the Chinese name "Gou Qi."

and male infertility and also benefit vision, kidney, and liver function [6–8].

**2. Nutraceutical and pharmaceutical values**

**2.1. Nutraceutical value**

1%, and others 1%.

(1.4%) [13].

**Figure 1.** Flower, fruit, and dried fruit appearance of *Lycium barbarum* L., *L. chinense* Mill., and *L. ruthenicum* Murr. *L. barbarum* bears royal purple flower (A) with lanceolate leaves and produces bright red berries (B) which resemble red raisins after they have been dried (C). The flower of *L. chinense* is purple (D), leaves are ovate, fresh (E) and dried (F) berries are orange-red. *L. ruthenicum* bears purple flowers that are lighter than *L. chinense* (G), leaves are subsessile and succulent in a linear shape (G). This species produces dark colored berries (H) that are harvested with peduncle and sepal; thus individual dried black berries often have peduncle and sepal (I).

sometimes in fascicles. Petioles are short. Flowers are solitary or clustered in leaf axils. The corolla is funnel or bell shaped in white, green, or purple. Five stamens are structured with filaments longer than the anthers. Anthers dehisce longitudinally. The fruit is a two-chambered, usually fleshy and juicy berry and typically orange-red or black (*L. ruthenicum*). Seeds are few or many, most with over 10. In the Northern Hemisphere, flowering occurs from June to September, and berry maturation starts from August to October.

Gojiberry has been consumed as food and used as medicine for more than 4000 years in China [2]. The first mentioned "Gou Qi" was in the ancient classic "Shen Nong Ben Cao Jing" (The Classic of Herbal Medicine), a Chinese book on agriculture and medicinal plants written between 200 and 250 AD. The fruit of *L. barbarum*, *L. chinense*, and *L. ruthenicum* are commonly called "Ningxia Gou Qi," "Gou Qi," and "Black Gou Qi," respectively. The fruit is a renowned Yin strengthening agent, and the root bark, known as "Di Gu Pi," is a cooling agent [3]. Traditional English vernacular names include boxthorn, fructus lycii, wolfberry, Chinese wolfberry, or matrimony vine [4]. Since the beginning of the century, the plant has been commonly called Goji, an appellation derived from the Chinese name "Gou Qi."

#### **2. Nutraceutical and pharmaceutical values**

The berries harvested from August to October are eaten as fresh fruit, dehydrated to make dried fruit or soaked in liquor to produce *Lycium* juice. As a medicinal food, it is used as a condiment with steamed rice. Young soft leaves can also be used as a vegetable. Gojiberry has been used for its anti-aging activities, tranquilizing and thirst-quenching effects, and its ability to increase stamina [5]. Consuming gojiberry has been shown to improve health-related problems, such as diabetes, hyperlipidemia, cancer, hepatitis, immune disorders, thrombosis, and male infertility and also benefit vision, kidney, and liver function [6–8].

#### **2.1. Nutraceutical value**

sometimes in fascicles. Petioles are short. Flowers are solitary or clustered in leaf axils. The corolla is funnel or bell shaped in white, green, or purple. Five stamens are structured with filaments longer than the anthers. Anthers dehisce longitudinally. The fruit is a two-chambered, usually fleshy and juicy berry and typically orange-red or black (*L. ruthenicum*). Seeds are few or many, most with over 10. In the Northern Hemisphere, flowering occurs from

**Figure 1.** Flower, fruit, and dried fruit appearance of *Lycium barbarum* L., *L. chinense* Mill., and *L. ruthenicum* Murr. *L. barbarum* bears royal purple flower (A) with lanceolate leaves and produces bright red berries (B) which resemble red raisins after they have been dried (C). The flower of *L. chinense* is purple (D), leaves are ovate, fresh (E) and dried (F) berries are orange-red. *L. ruthenicum* bears purple flowers that are lighter than *L. chinense* (G), leaves are subsessile and succulent in a linear shape (G). This species produces dark colored berries (H) that are harvested with peduncle and

Gojiberry has been consumed as food and used as medicine for more than 4000 years in China [2]. The first mentioned "Gou Qi" was in the ancient classic "Shen Nong Ben Cao Jing" (The Classic of Herbal Medicine), a Chinese book on agriculture and medicinal plants written between 200 and 250 AD. The fruit of *L. barbarum*, *L. chinense*, and *L. ruthenicum* are commonly called "Ningxia Gou Qi," "Gou Qi," and "Black Gou Qi," respectively. The fruit is a

June to September, and berry maturation starts from August to October.

sepal; thus individual dried black berries often have peduncle and sepal (I).

4 Breeding and Health Benefits of Fruit and Nut Crops

Scientific analysis of gojiberry constituents started in the 1970s. Qian et al. [9] reviewed 142 publications from 1975 to 2016 and summarized that at least 355 compounds occur in different species of *Lycium*, which were categorized as alkaloids at 20%, sterols, steroids, and their derivatives 16%, terpenoids 11%, amides 10%, flavonoids 9%, organic acid 9%, phenylpropanoids 9%, glycerogalactolipids 6%, ligans 4%, coumarins 3%, antraquinones 1%, peptides 1%, and others 1%.

The most intensively studied components are a group of water-soluble glycoconjugates, which are identified as arabinogalactan-proteins, commonly known as *L. barbarum* polysaccharides or LBPs. It is estimated that dried fruits comprise 5–8% of LBPs [10, 11]. Chinese Pharmacopeia [12] recommended using LBPs as an indicator for evaluation of gojiberry quality. Molecular weights of the LBPs range from 24 to 241 kDa, and they are mainly composed of six types of monosaccharides: arabinose, glucose, galactose, mannose, xylose, and rhamnose [13]. LBPs also contain galacturonic acid and 18 amino acids and share a glycan-o-ser glycopeptide structure [14]. The main chains of the glycan backbones of LBPs have been reported to be either alpha-(1–N6)-D-glucans or alpha-(1–N4)-D polygalacturonans [3, 15].

A second major group of metabolites in fruit is carotenoids [3], the content of which increases during the fruit ripening process. A total of 11 free carotenoids and 7 carotenoid esters were detected from unsaponified and saponified *L. barbarum* extracts [16]. Zeaxanthin dipalmitate was found to account for 80% of the total carotenoids [17]. β-Cryptoxanthin palmitate, zeaxanthin monopalmitate, small amounts of free zeaxanthin and *β*-carotene are also present [3]. Seeds contain zeaxanthin (83%), *β*-cryptoxanthin (7%), *β*-carotene (0.9%), and mutatoxanthin (1.4%) [13].

The fruits also consist of vitamins including ascorbic acid (vitamin C), riboflavin, and thiamin. The content of vitamin C (42 mg/100 g) is comparable to that of fresh lemon fruits [3]. Flavonoids are another important group of compounds. Total flavonoids of *L. barbarum* var. aurauticarpum, a yellow fruit variety, reach up to 2.0 mg/g which is four times higher than red-fruited *L. barbarum*. Fruits of *L. ruthenicum* contain oligomeric proanthocyanidins (OPC) at 3690 mg/100 g, which surpasses blueberries at 3380 mg/100 g [18]. Furthermore, aglycones myricetin, quercetin, and kaempferol were identified after hydrolysis [19].

zeaxanthin and antioxidant levels significantly, which protected eyes from hypopigmentation

Gojiberry Breeding: Current Status and Future Prospects http://dx.doi.org/10.5772/intechopen.76388 7

There are several reviewed articles regarding chemical constituents and nutraceutical value of gojiberry as well as pharmaceutical effects of gojiberry consumption. The reader is referred

Gojiberry has been commercialized with different names depending on the origin of plants and product types. "Ningxia Goji berry," produced from Ningxia (Ningxia Hui Autonomous Region, which is equivalent to a Province, located in Northwestern China) and harvested primarily from *L. barbarum*, is considered to be the authentic origin. "Black Wolfberry" is the berry harvested from *L. ruthenicum*. "Himalayan Goji berry" or "Tibetan Goji berry" are also on the global functional food market; these two names are used by health food promoters for a nomenclatural marketing advantage, though commercial cultivation of the crop does not occur in those regions [42]. In addition to being an important traditional Chinese medicinal herb, commercialized products vary from dried berries in various sized bags to juices, beers, and wines. Gojiberry is also found in cookies, chocolate candies, muesli, sausages, and snack bars. Gojiberry products have been marketed online since 2002 and are termed as a "super-

Gojiberry and its products are sold as food or food supplements in the US and in Europe [16]. These products, however, are not allowed to be promoted as drugs, and therapeutic claims are prohibited thus far. In the US, the Food and Drug Administration (FDA) warned about some Gojiberry juice distributors using marketing claims which violated the Food Drug and Cosmetic Act [3]. In the United Kingdom (UK), the Food Standards Agency in 2007 concluded that there were sufficient records of alimentary use of Gojiberry in the UK before 1997 and thus the fruit does not fall under the Novel Food legislation. In the US, however, Gojiberry is

China is the main producer and supplier of gojiberry in the world. The majority of commercially produced gojiberry comes from Ningxia with gojiberry plantations typically ranging from 40 and 400 ha. Gojiberry, primarily *L. barbarum,* has been produced along the fertile floodplains of the Yellow River for more than 700 years. Fresh fruits are uniformly orange-red, and dried fruit has a sweet-and-tangy flavor. Ningxia has earned a reputation for premium quality gojiberries. As of 2015, more than 66 million hectares were planted with gojiberry in Ningxia. The region is recognized as the largest annual harvest in China, accounting for 45% of the nation's total yield of gojiberries. Fresh red or black berries are harvested starting in August. The harvested fruits are immediately washed and then preserved by drying them directly under full sun or through dry machines. Gojiberries are celebrated each August in Ningxia with an annual festival coinciding with the berry harvest. The celebration was originally held in Ningxia's capital, Yinchuan City;

not listed on the generally regarded as safe (GRAS) list by the FDA [3].

and accumulation of oxidative stress compounds that can damage the macula [41].

to these reviews [3, 9, 11, 16, 31] for more detailed information.

**2.3. Uses**

food" [43].

**3. Gojiberry production**

Gojiberry fruits contain 1–2.7% free amino acids, of which proline is the major constituent [3]. Non-proteinogenic amino acids include taurine, *γ*-aminobutyric acid, and betaine (trimethylglycine) [20, 21]. Some miscellaneous compounds, such as β-sitosterol and its glucoside daucosterol, scopoletin, p-coumaric acid, the dopamine derivative lyciumide A, and L-monomethyl succinate occur in fruits [22–24].

#### **2.2. Pharmaceutical value**

Consuming gojiberry has been shown to improve general well-being, anti-myelosuppression, and sleep quality. Five randomized clinical studies conducted in the US [25–28] and China [29, 30] indicated that daily consumption of standardized *L. barbarum* fruit juice [GoChi juice (Goji juice) 120 ml = equivalent to 150 g of fresh fruit] for 14 or 30 days improved general well-being including neurological and psychological status, cardiovascular, joint, and muscle functions as well as gastrointestinal regularity without any adverse effects.

LBPs from *L. barbarum* have anti-aging and neuroprotective activities [31]. An arabinogalactan-protein (LBP-III) isolated from LBPs exhibited cytoprotective effects against stress. The protective role was mediated by reducing the phosphorylation of double-stranded RNAdependent protein kinase (PKR) and also by decreasing the dithiothreitol (DTT)-induced LDH release and caspase-3 activity [32]. The reduction of PKR was caused by beta-amyloid peptide. It is well known that the phosphorylation state of PKR increases with age, the reduction of phosphorylation triggered by beta-amyloid peptide suggests that LBP-III from gojiberry could be a potential neuroprotective agent [33, 34]. *L. barbarum* intake was effective to control waist circumference in humans and may reduce the risks of metabolic syndrome [28].

Several experimental and clinical studies showed that *L. barbarum* exhibited anti-diabetic effects*. L. barbarum* reduced oxidation in patients with retinopathy [35]. In a randomized diabetic retinopathy study, the intake of fruit of *L. barbarum* for 3 months was shown to increase the contents of vitamin C by 61% and the activities of SOD by 87% and also to reduce serum content of lipid peroxide by about 20% compared to a control group [36]. *L. barbarum* was also effective when used for improving immunologic function of red blood cells in patients with diabetic retinopathy [36]. In a study with 44 patients with diabetic retinopathy, RBC C<sup>3</sup> b receptor rosette (RBC-C<sup>3</sup> bRR) in the patients treated with *L. barbarum* for 3 months decreased [37]. LBPs were shown to have immunomodulatory effects on patients with type-2 diabetes by reducing T8 and interleukin 6 (IL-6) by 23% and increasing T4/T8 and IL-2 by 30 and 62% compared to the normal level, respectively [38].

Gojiberry has been considered to be the richest source for zeaxanthin [39, 40] varying from 1.18 to 2.41 mg/g dried fruit [11]. Gojiberry is, thus, a natural pill for eye health. Drinking goji berry juice daily as a dietary supplementation for 90 days was reported to increase plasma zeaxanthin and antioxidant levels significantly, which protected eyes from hypopigmentation and accumulation of oxidative stress compounds that can damage the macula [41].

There are several reviewed articles regarding chemical constituents and nutraceutical value of gojiberry as well as pharmaceutical effects of gojiberry consumption. The reader is referred to these reviews [3, 9, 11, 16, 31] for more detailed information.

#### **2.3. Uses**

Flavonoids are another important group of compounds. Total flavonoids of *L. barbarum* var. aurauticarpum, a yellow fruit variety, reach up to 2.0 mg/g which is four times higher than red-fruited *L. barbarum*. Fruits of *L. ruthenicum* contain oligomeric proanthocyanidins (OPC) at 3690 mg/100 g, which surpasses blueberries at 3380 mg/100 g [18]. Furthermore, aglycones

Gojiberry fruits contain 1–2.7% free amino acids, of which proline is the major constituent [3]. Non-proteinogenic amino acids include taurine, *γ*-aminobutyric acid, and betaine (trimethylglycine) [20, 21]. Some miscellaneous compounds, such as β-sitosterol and its glucoside daucosterol, scopoletin, p-coumaric acid, the dopamine derivative lyciumide A, and

Consuming gojiberry has been shown to improve general well-being, anti-myelosuppression, and sleep quality. Five randomized clinical studies conducted in the US [25–28] and China [29, 30] indicated that daily consumption of standardized *L. barbarum* fruit juice [GoChi juice (Goji juice) 120 ml = equivalent to 150 g of fresh fruit] for 14 or 30 days improved general well-being including neurological and psychological status, cardiovascular, joint, and muscle

LBPs from *L. barbarum* have anti-aging and neuroprotective activities [31]. An arabinogalactan-protein (LBP-III) isolated from LBPs exhibited cytoprotective effects against stress. The protective role was mediated by reducing the phosphorylation of double-stranded RNAdependent protein kinase (PKR) and also by decreasing the dithiothreitol (DTT)-induced LDH release and caspase-3 activity [32]. The reduction of PKR was caused by beta-amyloid peptide. It is well known that the phosphorylation state of PKR increases with age, the reduction of phosphorylation triggered by beta-amyloid peptide suggests that LBP-III from gojiberry could be a potential neuroprotective agent [33, 34]. *L. barbarum* intake was effective to control

waist circumference in humans and may reduce the risks of metabolic syndrome [28].

Several experimental and clinical studies showed that *L. barbarum* exhibited anti-diabetic effects*. L. barbarum* reduced oxidation in patients with retinopathy [35]. In a randomized diabetic retinopathy study, the intake of fruit of *L. barbarum* for 3 months was shown to increase the contents of vitamin C by 61% and the activities of SOD by 87% and also to reduce serum content of lipid peroxide by about 20% compared to a control group [36]. *L. barbarum* was also effective when used for improving immunologic function of red blood cells in patients with diabetic retinopathy [36]. In a study with 44 patients with diabetic retinopathy, RBC C<sup>3</sup>

[37]. LBPs were shown to have immunomodulatory effects on patients with type-2 diabetes by reducing T8 and interleukin 6 (IL-6) by 23% and increasing T4/T8 and IL-2 by 30 and 62%

Gojiberry has been considered to be the richest source for zeaxanthin [39, 40] varying from 1.18 to 2.41 mg/g dried fruit [11]. Gojiberry is, thus, a natural pill for eye health. Drinking goji berry juice daily as a dietary supplementation for 90 days was reported to increase plasma

bRR) in the patients treated with *L. barbarum* for 3 months decreased

myricetin, quercetin, and kaempferol were identified after hydrolysis [19].

functions as well as gastrointestinal regularity without any adverse effects.

L-monomethyl succinate occur in fruits [22–24].

6 Breeding and Health Benefits of Fruit and Nut Crops

**2.2. Pharmaceutical value**

receptor rosette (RBC-C<sup>3</sup>

compared to the normal level, respectively [38].

Gojiberry has been commercialized with different names depending on the origin of plants and product types. "Ningxia Goji berry," produced from Ningxia (Ningxia Hui Autonomous Region, which is equivalent to a Province, located in Northwestern China) and harvested primarily from *L. barbarum*, is considered to be the authentic origin. "Black Wolfberry" is the berry harvested from *L. ruthenicum*. "Himalayan Goji berry" or "Tibetan Goji berry" are also on the global functional food market; these two names are used by health food promoters for a nomenclatural marketing advantage, though commercial cultivation of the crop does not occur in those regions [42]. In addition to being an important traditional Chinese medicinal herb, commercialized products vary from dried berries in various sized bags to juices, beers, and wines. Gojiberry is also found in cookies, chocolate candies, muesli, sausages, and snack bars. Gojiberry products have been marketed online since 2002 and are termed as a "superfood" [43].

Gojiberry and its products are sold as food or food supplements in the US and in Europe [16]. These products, however, are not allowed to be promoted as drugs, and therapeutic claims are prohibited thus far. In the US, the Food and Drug Administration (FDA) warned about some Gojiberry juice distributors using marketing claims which violated the Food Drug and Cosmetic Act [3]. In the United Kingdom (UK), the Food Standards Agency in 2007 concluded that there were sufficient records of alimentary use of Gojiberry in the UK before 1997 and thus the fruit does not fall under the Novel Food legislation. In the US, however, Gojiberry is not listed on the generally regarded as safe (GRAS) list by the FDA [3].

#### **3. Gojiberry production**

b

China is the main producer and supplier of gojiberry in the world. The majority of commercially produced gojiberry comes from Ningxia with gojiberry plantations typically ranging from 40 and 400 ha. Gojiberry, primarily *L. barbarum,* has been produced along the fertile floodplains of the Yellow River for more than 700 years. Fresh fruits are uniformly orange-red, and dried fruit has a sweet-and-tangy flavor. Ningxia has earned a reputation for premium quality gojiberries. As of 2015, more than 66 million hectares were planted with gojiberry in Ningxia. The region is recognized as the largest annual harvest in China, accounting for 45% of the nation's total yield of gojiberries. Fresh red or black berries are harvested starting in August. The harvested fruits are immediately washed and then preserved by drying them directly under full sun or through dry machines. Gojiberries are celebrated each August in Ningxia with an annual festival coinciding with the berry harvest. The celebration was originally held in Ningxia's capital, Yinchuan City; the festival has been moved since 2000 to Zhongning County. Furthermore, gojiberries are drought-tolerant plants. As Ningxia's borders merge with three deserts, gojiberries are also planted to control erosion and reclaim irrigable soils from desertification. Commercial volumes of gojiberry also grow in other provinces of China including Xingjiang, Hebei, Inner Mongolia, Qinghai, Gansu, and Shaanxi. Jinghe County in Xingjiang, Julu County in Hebei, and Huangjinhou in Inner Mongolia also produce high quality gojiberries

America, primarily in Argentina and Chile, with more than 30 species. There are about 21 species in Southwestern and North America, approximately 17 species in southern Africa [45], and about 10 species in Eurasia [46]. Based on the analysis of chloroplast DNA sequences, Fukuda et al. [46] proposed that *Lycium* originated from the New World. All species in southern Africa, Australia, and Eurasia have a common progenitor from the New World. Australian and Eurasian species originated once from a southern African progenitor, and *L. sandwicense* differentiated from the New World species. As a result, phylogenetic analysis showed that gojiberries (*L. barbarum*, *L. chinense*, and *L. ruthenicum*) are clustered with *L. europaceum* L. as they belong to Eurasian species. The Eurasian species are closely related to species from Australia such as *L. australe* F. Muell. and also those from southern Africa, such as *L. afrum* L., *L. cinereum* Thumb., *L. ferocissimum* Miers, *L. pilifolium* C. H. Wright, *L. prunus-spinosa* Dunal, *L. schizocalyx* C. H. Wright, and *L. villosum* Schinz. Species from North or South America as

Gojiberry Breeding: Current Status and Future Prospects http://dx.doi.org/10.5772/intechopen.76388 9

Most *Lycium* species have perfect flowers and are bisexual or hermaphrodites. However, like some others in the family Solanaceae, *Lycium* species are generally considered to be outcrossed due to gametophytic self-incompatibility [47]. For example, allelic diversity at the self-incompatibility (*S*) gene in *L. andersonii* was estimated to have more than 35 alleles, and coalescence analysis showed that the *S*-allele lineages in this species are older than the genus as a whole, indicating that self-incompatibility is the basal condition for *Lycium* [48]. Most species are diploid with chromosome number of 2n = 2x = 24. However, Miller and Venable [49] reported that three North American species, *L. californicum* Nutt. Ex Gray, *L. exsertum* A. Gray, and *L. fremontii* A. Gray are polyploids and display functional dioecy. In addition, seven species in Africa have separate male and female plants [50, 51]. Levin and Miller [52] believed that gender dimorphism (the presence of two sexual morphs in a population) evolved twice among North American *Lycium* and probably three times in Africa. Thus, gender dimorphism is more common among African *Lycium*, occurring in 7 of the 27 African species (26%) compared to only 3 of 50 American species (6%). Furthermore, gender dimorphism has been shown to be uniformly associated with polyploidy, which resulted in a proposition that polyploidy disrupts self-incompatibility of North America diploid *Lycium* species and resultant self-compatible polyploids are then subject to invasion by male sterile

It is unknown when and how *Lycium* species were dispersed to Eurasian regions. Among the 10 Eurasian species, there are 7 species that have been naturalized in China including *L. barbarum*, *L. chinense*, *L. cylindricum* Kuang, *L. dasystemum* Pojark., *L. ruthenicum*, *L. truncatum* Y.C. Wang, and *L. yunnanense* Kuang [53], of which *L. barbarum*, *L. chinense*, and *L. ruthenicum* are the most popular species. *L. barbarum*, *L. dasystemum*, and *L. ruthenicum* are primarily distributed in northwest China including Ningxia, Gansu, Inner Mongolia, Qinghai, Xinjiang, Shaanxi, and Shanxi. *L. chinense* is largely dispensed in central and east China. *L. cylindricum* is spread in Gansu, Inner Mongolia, Ningxia, Qinghai, North Shaanxi, Xinjiang, Tibet as well as Afghanistan, Kazakhstan, Kyrgyzstan, Mongolia, Pakistan, Russia, Tajikistan, Turkmenistan, and Uzbekistan. *L. truncatum* is distributed in dry regions with altitude ranging from 800 to 1500 m including Gansu, Inner Mongolia, Ningxia, Shanxi, and Xinjiang. *L. yunnanense* is

well as Pacific Island were clustered together [46].

mainly situated in Yunnan, southwest China.

plants [47].

Gojiberry has also been commercially produced in the other countries including India, Korea, Japan, and other Asian countries. During the first decade of the twenty-first century, farmers in Canada and the US began cultivating gojiberry on a commercial scale to meet potential markets for fresh berries, juice, and processed products. Gojiberry has been propagated by tissue culture by Agri-starts, Inc. in Apopka, Florida, US and tissue-cultured liners are sold nationwide for production in the US and Canada. Goji farm USA celebrated the harvest of gojiberry from 8000 plants grown in Sonoma Valley of the moon region in California, US (GojiFarmUSA.com)

#### **4. Current concerns on gojiberry production**

The demand for quality gojiberry fruit has become much greater than the supply. A variety of factors are implicated in the shortage of supply, which includes the availability of reliable cultivars, the lack of corresponding production protocols, disease and pest control information, appropriate methods for processing fruits, quality control, and food safety issues. Among these factors, the availability of new and reliable cultivars is a key factor. China is the only country with gojiberry breeding programs. Current breeding efforts have been largely limited to *L. barbarum*, *L. chinense*, and *L. ruthenicum*, and only a few cultivars have been released and utilized. Some of the cultivars are unstable in fruit setting, fruit size, and final yield due to seed propagation or inappropriate planting of comparable cultivars adjacently for pollination. Additionally, the evaluation of gojiberry germplasm has remained in the descriptive stage; their potential has not been fully exploited. Thus, a better understanding of the current status of gojiberry breeding is important for future development of new cultivars and for increasing its production.

#### **5. Genetic improvement of gojiberry**

It has been well documented that naturally occurring variation among wild relatives of cultivated crops is an underexploited resource in plant breeding [44]. The genus *Lycium*, a member of the family Solanaceae, comprises about 80 species [2]. Most of them remain in the wild, and their reproduction systems have been studied, but their agronomic traits and nutraceutical and pharmaceutical value have barely been exploited*.*

#### **5.1. Genetic resources**

*Lycium* species are distributed in temperate and subtropical regions worldwide, but are absent in both the Old and New World tropics. Areas of greatest species richness are in South America, primarily in Argentina and Chile, with more than 30 species. There are about 21 species in Southwestern and North America, approximately 17 species in southern Africa [45], and about 10 species in Eurasia [46]. Based on the analysis of chloroplast DNA sequences, Fukuda et al. [46] proposed that *Lycium* originated from the New World. All species in southern Africa, Australia, and Eurasia have a common progenitor from the New World. Australian and Eurasian species originated once from a southern African progenitor, and *L. sandwicense* differentiated from the New World species. As a result, phylogenetic analysis showed that gojiberries (*L. barbarum*, *L. chinense*, and *L. ruthenicum*) are clustered with *L. europaceum* L. as they belong to Eurasian species. The Eurasian species are closely related to species from Australia such as *L. australe* F. Muell. and also those from southern Africa, such as *L. afrum* L., *L. cinereum* Thumb., *L. ferocissimum* Miers, *L. pilifolium* C. H. Wright, *L. prunus-spinosa* Dunal, *L. schizocalyx* C. H. Wright, and *L. villosum* Schinz. Species from North or South America as well as Pacific Island were clustered together [46].

the festival has been moved since 2000 to Zhongning County. Furthermore, gojiberries are drought-tolerant plants. As Ningxia's borders merge with three deserts, gojiberries are also planted to control erosion and reclaim irrigable soils from desertification. Commercial volumes of gojiberry also grow in other provinces of China including Xingjiang, Hebei, Inner Mongolia, Qinghai, Gansu, and Shaanxi. Jinghe County in Xingjiang, Julu County in Hebei, and

Gojiberry has also been commercially produced in the other countries including India, Korea, Japan, and other Asian countries. During the first decade of the twenty-first century, farmers in Canada and the US began cultivating gojiberry on a commercial scale to meet potential markets for fresh berries, juice, and processed products. Gojiberry has been propagated by tissue culture by Agri-starts, Inc. in Apopka, Florida, US and tissue-cultured liners are sold nationwide for production in the US and Canada. Goji farm USA celebrated the harvest of gojiberry from 8000 plants grown in Sonoma Valley of the moon region in California, US (GojiFarmUSA.com)

The demand for quality gojiberry fruit has become much greater than the supply. A variety of factors are implicated in the shortage of supply, which includes the availability of reliable cultivars, the lack of corresponding production protocols, disease and pest control information, appropriate methods for processing fruits, quality control, and food safety issues. Among these factors, the availability of new and reliable cultivars is a key factor. China is the only country with gojiberry breeding programs. Current breeding efforts have been largely limited to *L. barbarum*, *L. chinense*, and *L. ruthenicum*, and only a few cultivars have been released and utilized. Some of the cultivars are unstable in fruit setting, fruit size, and final yield due to seed propagation or inappropriate planting of comparable cultivars adjacently for pollination. Additionally, the evaluation of gojiberry germplasm has remained in the descriptive stage; their potential has not been fully exploited. Thus, a better understanding of the current status of gojiberry breeding is important for future development of new cultivars and for increasing its production.

It has been well documented that naturally occurring variation among wild relatives of cultivated crops is an underexploited resource in plant breeding [44]. The genus *Lycium*, a member of the family Solanaceae, comprises about 80 species [2]. Most of them remain in the wild, and their reproduction systems have been studied, but their agronomic traits and nutraceutical

*Lycium* species are distributed in temperate and subtropical regions worldwide, but are absent in both the Old and New World tropics. Areas of greatest species richness are in South

Huangjinhou in Inner Mongolia also produce high quality gojiberries

8 Breeding and Health Benefits of Fruit and Nut Crops

**4. Current concerns on gojiberry production**

**5. Genetic improvement of gojiberry**

**5.1. Genetic resources**

and pharmaceutical value have barely been exploited*.*

Most *Lycium* species have perfect flowers and are bisexual or hermaphrodites. However, like some others in the family Solanaceae, *Lycium* species are generally considered to be outcrossed due to gametophytic self-incompatibility [47]. For example, allelic diversity at the self-incompatibility (*S*) gene in *L. andersonii* was estimated to have more than 35 alleles, and coalescence analysis showed that the *S*-allele lineages in this species are older than the genus as a whole, indicating that self-incompatibility is the basal condition for *Lycium* [48]. Most species are diploid with chromosome number of 2n = 2x = 24. However, Miller and Venable [49] reported that three North American species, *L. californicum* Nutt. Ex Gray, *L. exsertum* A. Gray, and *L. fremontii* A. Gray are polyploids and display functional dioecy. In addition, seven species in Africa have separate male and female plants [50, 51]. Levin and Miller [52] believed that gender dimorphism (the presence of two sexual morphs in a population) evolved twice among North American *Lycium* and probably three times in Africa. Thus, gender dimorphism is more common among African *Lycium*, occurring in 7 of the 27 African species (26%) compared to only 3 of 50 American species (6%). Furthermore, gender dimorphism has been shown to be uniformly associated with polyploidy, which resulted in a proposition that polyploidy disrupts self-incompatibility of North America diploid *Lycium* species and resultant self-compatible polyploids are then subject to invasion by male sterile plants [47].

It is unknown when and how *Lycium* species were dispersed to Eurasian regions. Among the 10 Eurasian species, there are 7 species that have been naturalized in China including *L. barbarum*, *L. chinense*, *L. cylindricum* Kuang, *L. dasystemum* Pojark., *L. ruthenicum*, *L. truncatum* Y.C. Wang, and *L. yunnanense* Kuang [53], of which *L. barbarum*, *L. chinense*, and *L. ruthenicum* are the most popular species. *L. barbarum*, *L. dasystemum*, and *L. ruthenicum* are primarily distributed in northwest China including Ningxia, Gansu, Inner Mongolia, Qinghai, Xinjiang, Shaanxi, and Shanxi. *L. chinense* is largely dispensed in central and east China. *L. cylindricum* is spread in Gansu, Inner Mongolia, Ningxia, Qinghai, North Shaanxi, Xinjiang, Tibet as well as Afghanistan, Kazakhstan, Kyrgyzstan, Mongolia, Pakistan, Russia, Tajikistan, Turkmenistan, and Uzbekistan. *L. truncatum* is distributed in dry regions with altitude ranging from 800 to 1500 m including Gansu, Inner Mongolia, Ningxia, Shanxi, and Xinjiang. *L. yunnanense* is mainly situated in Yunnan, southwest China.

#### **5.2. Domestication of** *L. barbarum*

How *L. barbarum* became naturalized in northwest China and centralized in Ningxia, particularly Zhongning County, is unclear. One possible reason could be due to the geographical location of Zhongning, as berries of *L. barbarum* produced in this county had the highest quality. An important irrigation project during Qin (221–206 BC), Han (206 BC–220 AD), and Tang (618–907 AD) dynasties channeled water from the Yellow River to irrigate farmland on the Yinchuan Plain in Ningxia, where Zhongning County was the gateway of the irrigation project. This project created favorable conditions for *L. barbarum* production. During Ming (1368–1644) and Qing (1644–1912) dynasties as well as the China Republic (1912–1949) periods, *L. barbarum* production was largely concentrated in Zhongning County with production acreage of about 200 hectares. Growers might have consciously harvested large fruits from individual plants. The harvested seeds might have been combined or kept separately by trees and planted the next year. Such conscious practices or domestication over thousands of years might have resulted in plants differing from their wild progenitors in several morphophysiological traits. Improved traits could be associated with seed retention and germination, growth habit, fruit size and coloration, and fruit edibility and taste. The domestication resulted in the establishment of landraces. No attention was given to the number of landraces in Zhongning County until the 1960s when Mr. Guofeng Qin conducted a survey in the County and identified 10 landraces, which included "Damaye," "Xiaomaye," "Heyemaye," and "Baitiaogouqi" [54]. "Damaye" was found in Mr. Zhuohan Zhang's garden who was a gojiberry grower. He probably never realized what important role this landrace has played in gojiberry cultivar development. "Damaye" grew to a height of 1.5 m with a canopy diameter of 1.7 m in 6 years. The fruit is spheroid-shaped, and 1000 fresh fruits weigh from 450 to 510 g. A single plant produces 7–8 kg fresh fruit. "Damaye" has been considered to be the most reliable landrace as it produces large, uniform fruits with little variation over the years.

associates released "Ningqi 3" in 2005. Another scientist, Mr. Zhongqin Hu at the Zhongning Wolfberry Industry Bureau made selections from "Damaye" and released "Ningqi 4" in 2005 [55]. Using "Damaye" and "Ningqi 1" as primary resources, subsequent selections resulted in the release of "Ningqi 5," "Ningqi 6," "Ningqi 7," "Ningqi 8," and "Ningqi 9." "Ningqi 5" is a male sterile cultivar, "Ningqi 6" and "Ningqi 8" require outcrossing, while "Ningqi 7" is self-compatible. "Ningqi 9" is a tetraploid. Both "Ningqi 5" and "Ningqi 9"

Gojiberry Breeding: Current Status and Future Prospects http://dx.doi.org/10.5772/intechopen.76388 11

Among the Ningqi series, "Ningqi 1" has been highly successful. It has a similar growth rate and canopy shape to "Damaye," but it produces ellipsoid-shaped fruits with 1000 fresh fruits weighing more than 586 g, which is 15–30% greater than "Damaye." "Ningqi 1" is particularly stable in production and is able to adapt to a wide range of environments. Its production was quickly expanded in the entire Ningxia region, subsequently Xinjiang, Gansu, Inner Mongolia, Hubei, and Shanxi, moving from northwest China to Central China in over 20 provinces. Its production acreage totaled 88,000 ha. Mr. Shenyuan Zhong, the inventor of "Ningqi 1," "Ningqi 2," and "Ningqi 3" was considered the father of Ningxia gojiberry. He received the National Science and Technology Progress Second Prize in China in 2005. Mr. Zhong passed away in 2012, but production of his cultivars, particularly "Ningqi 1," has rap-

The success of "Ningqi 1" is closely related to the reproduction mode. As mentioned earlier, *Lycium* species are self-incompatible. Recent studies of "Damaye" and "Ningqi 1" showed that the two are self-compatible [56], which ensures the stability of agronomic traits when propagated through seed. The selection scheme used by Mr. Zhong was similar to the application of the pure-line theory [57] with a modification by vegetative propagation to fix a phenotype if needed. The finding of self-compatibility in "Damaye" and "Ningqi 1" was somewhat unexpected because *L. barbarum* has been considered to be an outcrossing species. The finding provides explanation as to why "Damaye" was the most valuable landrace and also why "Ningqi

The performance of selected Ningqi series in the field suggested that several reproduction modes occurred in *L. barbarum.* Some are self-compatible, such as "Ningqi 1," and "Ningqi 7." Some were self-incompatible, such as "Ningqi 3," "Ningqi 4," "Ningqi 6," and "Ningqi 8." Whereas "Ningqi 5" showed high male sterility, and "Ningqi 9" was a tetraploid. As a result, intraspecific hybridizations have been carried out for breeding of new gojiberry cultivars. The hybridizations within *L. barbarum* included the use of self-incompatibility and male sterility and the crosses between Ningqi series cultivars with cultivars from the other regions [58]. For the use of the self-incompatible cultivars, one cultivar was planted in a row with another in a 1:1 ratio. The percentage of fruit set, 1000 fresh fruit weight, and total fruit yield per tree were evaluated for identifying the best combinations. For example, "Ningqi 6" should be planted with "Ningqi 8" in a 1:1 ratio for high fruit set. "Ningqi 5," the male sterility line, should be planted with "Ningqi 1" or "Ningqi 4" in a 1:1 or 1:2 ratio for maximizing fruit set. Ningqi cultivars were also hybridized with cultivars from Xinjiang and other provinces. An et al. [59]

were selected from "Ningqi 1."

idly increased in China.

1" was the most popular cultivar.

**5.4. Hybridization**

#### **5.3. Individual plant selection**

An organized breeding effort on gojiberry was initiated in the 1970s at the Ningxia Research Center of Wolfberry Engineering Technology, which was renamed as National Wolfberry Engineering Research Center in 2011. The breeding center is located in Yinchuan City, Ningxia, China. This is the only federally sponsored institute devoted to gojiberry research. Initial breeding efforts were primarily focused on the selection of individual plants with vigorous growth rates, larger fruit, and higher fruit yield. Mr. Shenyuan Zhong volunteered to work in Ningxia after graduation from the Northwest Agricultural College and stayed in Zhongning County from 1965 to 1985. Mr. Zhong started individual plant selection from "Damaye" populations. A large number of individual plants were selected from "Damaye." Seeds were harvested from each selected individual plant, and individual plant's progenies were evaluated separately. If progenies from selected plants were variable, cloning propagation, such as rooting of cuttings, was used to stabilize the phenotypes. Using this selection method, Mr. Zhong selected 12 lines. After large-scale progeny testing, he released cultivars: "Ningqi 1" and "Ningqi 2" [54, 55]. Through mass selection, Mr. Zhong and his associates released "Ningqi 3" in 2005. Another scientist, Mr. Zhongqin Hu at the Zhongning Wolfberry Industry Bureau made selections from "Damaye" and released "Ningqi 4" in 2005 [55]. Using "Damaye" and "Ningqi 1" as primary resources, subsequent selections resulted in the release of "Ningqi 5," "Ningqi 6," "Ningqi 7," "Ningqi 8," and "Ningqi 9." "Ningqi 5" is a male sterile cultivar, "Ningqi 6" and "Ningqi 8" require outcrossing, while "Ningqi 7" is self-compatible. "Ningqi 9" is a tetraploid. Both "Ningqi 5" and "Ningqi 9" were selected from "Ningqi 1."

Among the Ningqi series, "Ningqi 1" has been highly successful. It has a similar growth rate and canopy shape to "Damaye," but it produces ellipsoid-shaped fruits with 1000 fresh fruits weighing more than 586 g, which is 15–30% greater than "Damaye." "Ningqi 1" is particularly stable in production and is able to adapt to a wide range of environments. Its production was quickly expanded in the entire Ningxia region, subsequently Xinjiang, Gansu, Inner Mongolia, Hubei, and Shanxi, moving from northwest China to Central China in over 20 provinces. Its production acreage totaled 88,000 ha. Mr. Shenyuan Zhong, the inventor of "Ningqi 1," "Ningqi 2," and "Ningqi 3" was considered the father of Ningxia gojiberry. He received the National Science and Technology Progress Second Prize in China in 2005. Mr. Zhong passed away in 2012, but production of his cultivars, particularly "Ningqi 1," has rapidly increased in China.

The success of "Ningqi 1" is closely related to the reproduction mode. As mentioned earlier, *Lycium* species are self-incompatible. Recent studies of "Damaye" and "Ningqi 1" showed that the two are self-compatible [56], which ensures the stability of agronomic traits when propagated through seed. The selection scheme used by Mr. Zhong was similar to the application of the pure-line theory [57] with a modification by vegetative propagation to fix a phenotype if needed. The finding of self-compatibility in "Damaye" and "Ningqi 1" was somewhat unexpected because *L. barbarum* has been considered to be an outcrossing species. The finding provides explanation as to why "Damaye" was the most valuable landrace and also why "Ningqi 1" was the most popular cultivar.

#### **5.4. Hybridization**

**5.2. Domestication of** *L. barbarum*

10 Breeding and Health Benefits of Fruit and Nut Crops

with little variation over the years.

**5.3. Individual plant selection**

How *L. barbarum* became naturalized in northwest China and centralized in Ningxia, particularly Zhongning County, is unclear. One possible reason could be due to the geographical location of Zhongning, as berries of *L. barbarum* produced in this county had the highest quality. An important irrigation project during Qin (221–206 BC), Han (206 BC–220 AD), and Tang (618–907 AD) dynasties channeled water from the Yellow River to irrigate farmland on the Yinchuan Plain in Ningxia, where Zhongning County was the gateway of the irrigation project. This project created favorable conditions for *L. barbarum* production. During Ming (1368–1644) and Qing (1644–1912) dynasties as well as the China Republic (1912–1949) periods, *L. barbarum* production was largely concentrated in Zhongning County with production acreage of about 200 hectares. Growers might have consciously harvested large fruits from individual plants. The harvested seeds might have been combined or kept separately by trees and planted the next year. Such conscious practices or domestication over thousands of years might have resulted in plants differing from their wild progenitors in several morphophysiological traits. Improved traits could be associated with seed retention and germination, growth habit, fruit size and coloration, and fruit edibility and taste. The domestication resulted in the establishment of landraces. No attention was given to the number of landraces in Zhongning County until the 1960s when Mr. Guofeng Qin conducted a survey in the County and identified 10 landraces, which included "Damaye," "Xiaomaye," "Heyemaye," and "Baitiaogouqi" [54]. "Damaye" was found in Mr. Zhuohan Zhang's garden who was a gojiberry grower. He probably never realized what important role this landrace has played in gojiberry cultivar development. "Damaye" grew to a height of 1.5 m with a canopy diameter of 1.7 m in 6 years. The fruit is spheroid-shaped, and 1000 fresh fruits weigh from 450 to 510 g. A single plant produces 7–8 kg fresh fruit. "Damaye" has been considered to be the most reliable landrace as it produces large, uniform fruits

An organized breeding effort on gojiberry was initiated in the 1970s at the Ningxia Research Center of Wolfberry Engineering Technology, which was renamed as National Wolfberry Engineering Research Center in 2011. The breeding center is located in Yinchuan City, Ningxia, China. This is the only federally sponsored institute devoted to gojiberry research. Initial breeding efforts were primarily focused on the selection of individual plants with vigorous growth rates, larger fruit, and higher fruit yield. Mr. Shenyuan Zhong volunteered to work in Ningxia after graduation from the Northwest Agricultural College and stayed in Zhongning County from 1965 to 1985. Mr. Zhong started individual plant selection from "Damaye" populations. A large number of individual plants were selected from "Damaye." Seeds were harvested from each selected individual plant, and individual plant's progenies were evaluated separately. If progenies from selected plants were variable, cloning propagation, such as rooting of cuttings, was used to stabilize the phenotypes. Using this selection method, Mr. Zhong selected 12 lines. After large-scale progeny testing, he released cultivars: "Ningqi 1" and "Ningqi 2" [54, 55]. Through mass selection, Mr. Zhong and his

The performance of selected Ningqi series in the field suggested that several reproduction modes occurred in *L. barbarum.* Some are self-compatible, such as "Ningqi 1," and "Ningqi 7." Some were self-incompatible, such as "Ningqi 3," "Ningqi 4," "Ningqi 6," and "Ningqi 8." Whereas "Ningqi 5" showed high male sterility, and "Ningqi 9" was a tetraploid. As a result, intraspecific hybridizations have been carried out for breeding of new gojiberry cultivars. The hybridizations within *L. barbarum* included the use of self-incompatibility and male sterility and the crosses between Ningqi series cultivars with cultivars from the other regions [58]. For the use of the self-incompatible cultivars, one cultivar was planted in a row with another in a 1:1 ratio. The percentage of fruit set, 1000 fresh fruit weight, and total fruit yield per tree were evaluated for identifying the best combinations. For example, "Ningqi 6" should be planted with "Ningqi 8" in a 1:1 ratio for high fruit set. "Ningqi 5," the male sterility line, should be planted with "Ningqi 1" or "Ningqi 4" in a 1:1 or 1:2 ratio for maximizing fruit set. Ningqi cultivars were also hybridized with cultivars from Xinjiang and other provinces. An et al. [59] made crosses between "Ningqi 1" and "Ningqi 9" (a tetraploid) and selected a triploid cultivar, which produced fruit that had no or little seeds, contained more sugar and amino acids with better taste compared to "Ningqi 1," and was resistant to aphid infestation.

shoots, not in the fruit. Additionally, field test of the transgenic plants showed that rhizo-

Gojiberry Breeding: Current Status and Future Prospects http://dx.doi.org/10.5772/intechopen.76388 13

In a study of genetic engineering of targeted genes, five carotenogenic genes from *L. barbarum*: geranylgeranyl diphosphate synthase, phytoene synthase and delta-carotene desaturase gene, lycopene beta-cyclase, and lycopene epsilon-cyclase were functionally analyzed in transgenic tobacco (*Nicotiana tabacum* L.) plants. Results showed that all transgenic tobacco plants constitutively expressing these genes and beta-carotene contents in their leaves and flowers increased [72]. These results imply that such genes could be used for improving *L.* 

There is a growing interest in gojiberry around the world, but some questions about *Lycium* species are still unanswered. *Lycium* species originated in South and North America, but how and when they were dispersed to Africa and Eurasian regions remain unclear. How has *L. sandwicense* been distributed across different island archipelagos? Why has *L. barbarum* been domesticated mainly in northwest China, and how has it become an important medicinal plant known as Ningxia gojiberry? A total of 355 compounds have been identified thus far [9], do any other compounds remain to be discovered? Nutraceutical and pharmaceutical value

A fundamental question from a breeding point of view, however, is the reproduction modes of *Lycium* species. Almost all reports in the literature described that diploid *Lycium* species are outcrossed due to their gametophytic self-incompatibility. Miller and Venable [47] used North American species to show that gender dimorphism has evolved in polyploid, selfcompatible taxa from co-sexual, self-incompatible diploids. They proposed that polyploidy is a trigger of unrecognized importance for the evolution of gender dimorphism, which operates by disrupting self-incompatibility and leading to inbreeding depression. Subsequently, male sterile mutants invade and increase because they are unable to inbreed. Research results from China, however, showed that *L. barbarum* is diploid, no occurrence of dimorphism, and landraces range from self-compatible to self-incompatible. The primary reason for the dominance of "Damaye" is its self-compatibility. The most popular cultivar selected from "Damaye," "Ningqi 1," is also widely produced and reproduced due to its self-compatibility. Was *L. barbarum* originally a self-compatible species being dispensed to Eurasian regions? Baker's Law [73–75] stated that self-compatible species are more likely to be successful island colonizers than obligate out-crossers that require pollen transfer between plants (i.e., self-incompatible species). In support of this law, a higher frequency of self-compatibility, as opposed to selfincompatibility, has been documented in island flora. Although the region where *L. barbarum* naturalized is not an island, its surroundings may be similar to an isolated region. Another possibility could be that *L. barbarum* is a self-incompatible species, and adaptation to northwest China caused switching to self-compatibility. The transition from self-incompatibility to self-compatibility has occurred often in the history of Solanaceae [76]. Thus, the reproduction

sphere microorganisms were not affected by the expression of the transgene [71].

*barbarum* in beta-carotene biosynthesis.

of important compounds require further investigation.

**6. Retrospect and prospect**

Interspecific hybridization has also been used for developing new cultivars. Crosses between "Ningqi 1" with a wild species by Li et al. [60] resulted in the release of "Ningqi Cai 1." It produces green, tender shoots and leaves, which is used as a vegetable, not for fruit production. This unique cultivar contains 352 g/kg protein, 18 amino acids with vitamin C at 135 mg/kg, and has been widely grown in China as a vegetable crop. Reciprocal crosses were made between either "Ningqi 1" or "Ningqi 4" with cultivars of *L. ruthenicum*, and results showed that they were highly compatible, resulting in the fruit set rate ranging from 52.38 to 91.43% [58]. Furthermore, Wang et al. [61] crossed *L. barbarum* with tomato (*Solanum lycopersicum* L.). Seven lines were selected from 21 cross combinations, and two of them flowered and produced fruit, suggesting *L. barbarum* has a wide range of crossability within the family Solanaceae.

#### **5.5. Breeding through chromosome manipulation**

Chromosome manipulation has been another approach for improving *L. barbarum* cultivars. In vitro culture of *L. barbarum* anthers produced haploid plants (2n = 12), and subsequent culture of hypocotyls of the haploids caused simultaneous doubling, resulting in homozygous diploid plants [62]. In vitro culture of endosperm of *L. barbarum* by Wang et al. [63] and Gu et al. [64] resulted in the isolation of triploid plants from mixoploid populations. Colchicine treatment of in vitro cultured meristems [65] or in vitro culture of ovary [66] produced tetraploid *L. barbarum*. In general, fruits produced from triploid and tetraploid plants were larger than diploid plants. Additionally, polyploid plants had larger flowers and thicker fruit pulp than the diploid plants.

#### **5.6. Biotechnological approaches**

Biotechnological approaches for improving gojiberry have been limited thus far. Methods for in vitro culture of existing meristems, anthers, embryos, and endosperm have been reported, and some of the methods have become well established. In vitro cultured materials were also used for inducing mutation, and some progress has been made. For example, a breeding line for resistance to *Fusarium graminearum* was produced from in vitro cultured embryonic calluses treated by 60Co-γ ray under the selection pressure of crude toxin of *Fusarium graminearum* [67]. Selection of EMS-treated embryonic calluses in the presence of 1.5% NaCl resulted in the development of salt tolerant plants [68].

*Agrobacterium tumefaciens*-mediated genetic transformation was established in *L. barbarum* [69]. In attempts to improve *L. barbarum* resistance to aphids, transgenic plants containing the gene encoding snowdrop lectin (*Galanthus nivalis* agglutinin, GNA) were generated [70]. The transgenic plants showed aphid resistance and also increased fruit weight and total sugar content, but seed count decreased. Because GNA was under the control of constitutive and phloem-specific promoters, the transgene product was only detected in leaves and young shoots, not in the fruit. Additionally, field test of the transgenic plants showed that rhizosphere microorganisms were not affected by the expression of the transgene [71].

In a study of genetic engineering of targeted genes, five carotenogenic genes from *L. barbarum*: geranylgeranyl diphosphate synthase, phytoene synthase and delta-carotene desaturase gene, lycopene beta-cyclase, and lycopene epsilon-cyclase were functionally analyzed in transgenic tobacco (*Nicotiana tabacum* L.) plants. Results showed that all transgenic tobacco plants constitutively expressing these genes and beta-carotene contents in their leaves and flowers increased [72]. These results imply that such genes could be used for improving *L. barbarum* in beta-carotene biosynthesis.

#### **6. Retrospect and prospect**

made crosses between "Ningqi 1" and "Ningqi 9" (a tetraploid) and selected a triploid cultivar, which produced fruit that had no or little seeds, contained more sugar and amino acids

Interspecific hybridization has also been used for developing new cultivars. Crosses between "Ningqi 1" with a wild species by Li et al. [60] resulted in the release of "Ningqi Cai 1." It produces green, tender shoots and leaves, which is used as a vegetable, not for fruit production. This unique cultivar contains 352 g/kg protein, 18 amino acids with vitamin C at 135 mg/kg, and has been widely grown in China as a vegetable crop. Reciprocal crosses were made between either "Ningqi 1" or "Ningqi 4" with cultivars of *L. ruthenicum*, and results showed that they were highly compatible, resulting in the fruit set rate ranging from 52.38 to 91.43% [58]. Furthermore, Wang et al. [61] crossed *L. barbarum* with tomato (*Solanum lycopersicum* L.). Seven lines were selected from 21 cross combinations, and two of them flowered and produced fruit, suggesting *L. barbarum* has a wide range of crossability within the family

Chromosome manipulation has been another approach for improving *L. barbarum* cultivars. In vitro culture of *L. barbarum* anthers produced haploid plants (2n = 12), and subsequent culture of hypocotyls of the haploids caused simultaneous doubling, resulting in homozygous diploid plants [62]. In vitro culture of endosperm of *L. barbarum* by Wang et al. [63] and Gu et al. [64] resulted in the isolation of triploid plants from mixoploid populations. Colchicine treatment of in vitro cultured meristems [65] or in vitro culture of ovary [66] produced tetraploid *L. barbarum*. In general, fruits produced from triploid and tetraploid plants were larger than diploid plants. Additionally, polyploid plants had larger flowers and thicker fruit pulp

Biotechnological approaches for improving gojiberry have been limited thus far. Methods for in vitro culture of existing meristems, anthers, embryos, and endosperm have been reported, and some of the methods have become well established. In vitro cultured materials were also used for inducing mutation, and some progress has been made. For example, a breeding line for resistance to *Fusarium graminearum* was produced from in vitro cultured embryonic calluses treated by 60Co-γ ray under the selection pressure of crude toxin of *Fusarium graminearum* [67]. Selection of EMS-treated embryonic calluses in the presence of 1.5% NaCl

*Agrobacterium tumefaciens*-mediated genetic transformation was established in *L. barbarum* [69]. In attempts to improve *L. barbarum* resistance to aphids, transgenic plants containing the gene encoding snowdrop lectin (*Galanthus nivalis* agglutinin, GNA) were generated [70]. The transgenic plants showed aphid resistance and also increased fruit weight and total sugar content, but seed count decreased. Because GNA was under the control of constitutive and phloem-specific promoters, the transgene product was only detected in leaves and young

with better taste compared to "Ningqi 1," and was resistant to aphid infestation.

Solanaceae.

than the diploid plants.

**5.6. Biotechnological approaches**

**5.5. Breeding through chromosome manipulation**

12 Breeding and Health Benefits of Fruit and Nut Crops

resulted in the development of salt tolerant plants [68].

There is a growing interest in gojiberry around the world, but some questions about *Lycium* species are still unanswered. *Lycium* species originated in South and North America, but how and when they were dispersed to Africa and Eurasian regions remain unclear. How has *L. sandwicense* been distributed across different island archipelagos? Why has *L. barbarum* been domesticated mainly in northwest China, and how has it become an important medicinal plant known as Ningxia gojiberry? A total of 355 compounds have been identified thus far [9], do any other compounds remain to be discovered? Nutraceutical and pharmaceutical value of important compounds require further investigation.

A fundamental question from a breeding point of view, however, is the reproduction modes of *Lycium* species. Almost all reports in the literature described that diploid *Lycium* species are outcrossed due to their gametophytic self-incompatibility. Miller and Venable [47] used North American species to show that gender dimorphism has evolved in polyploid, selfcompatible taxa from co-sexual, self-incompatible diploids. They proposed that polyploidy is a trigger of unrecognized importance for the evolution of gender dimorphism, which operates by disrupting self-incompatibility and leading to inbreeding depression. Subsequently, male sterile mutants invade and increase because they are unable to inbreed. Research results from China, however, showed that *L. barbarum* is diploid, no occurrence of dimorphism, and landraces range from self-compatible to self-incompatible. The primary reason for the dominance of "Damaye" is its self-compatibility. The most popular cultivar selected from "Damaye," "Ningqi 1," is also widely produced and reproduced due to its self-compatibility. Was *L. barbarum* originally a self-compatible species being dispensed to Eurasian regions? Baker's Law [73–75] stated that self-compatible species are more likely to be successful island colonizers than obligate out-crossers that require pollen transfer between plants (i.e., self-incompatible species). In support of this law, a higher frequency of self-compatibility, as opposed to selfincompatibility, has been documented in island flora. Although the region where *L. barbarum* naturalized is not an island, its surroundings may be similar to an isolated region. Another possibility could be that *L. barbarum* is a self-incompatible species, and adaptation to northwest China caused switching to self-compatibility. The transition from self-incompatibility to self-compatibility has occurred often in the history of Solanaceae [76]. Thus, the reproduction modes in *L. barbarum* should be further investigated as the modes are fundamentally important for new cultivar development as documented in this article.

**Author details**

\*, ChihCheng T. Chao<sup>2</sup>

2 USDA-ARS, Plant Genetic Resources Unit, Geneva, NY, USA

\*Address all correspondence to: jjchen@ufl.edu

University of Florida, Apopka, FL, USA

Macmillan Press; 1992

Springer; 2015. pp. 1-26

2010;**76**:7-19

s13065-017-0287-z

and Xiangying Wei1,3

Gojiberry Breeding: Current Status and Future Prospects http://dx.doi.org/10.5772/intechopen.76388 15

1 Institute of Food and Agricultural Sciences, Mid-Florida Research and Education Center,

[1] Huxley A. The New Royal Horticultural Society Dictionary of Gardening. London, UK:

[2] Wang Y, Chen H, Wu M, Zeng S, Liu Y, Dong J. Chemical and genetic diversity of wolfberry. In: Chang RC-C, So K-F, editors. *Lycium barbarum* And Human Health. Dordrecht:

[3] Potterat O. Goji (*Lycium barbarum* and *L. chinense*): Phytochemistry, pharmacology and safety in the perspective of traditional uses and recent popularity. Planta Medica.

[4] Hänsel R, Keller K, Rimpler H, Schneider G. Hagers. Handbuch der Pharmazeutischen Praxis. In: Drogen E, editor. Berlin, Heidelberg, New York: Springer Verlag 1993

[5] Islam T, Yu X, Badwal TS, Xu B. Comparative studies on phenolic profiles, antioxidant capacities and carotenoid contents of red goji berry (*Lycium barbarum*) and black goji berry (*Lycium ruthenicum*). Chemistry Central Journal. 2017;**11**:59. DOI: 10.1186/

[6] Jung K, Chin YW, Kim YC, Kim J. Potentially hepatoprotective glycolipid constituents of

[7] Li X, Ma Y, Liu X. Effect of the *Lycium barbarum* polysaccharides on age-related oxidative

[8] Kocyigit E, Sanlier N. A review of composition and health effects of *Lycium barbarum*.

[9] Qian D, Zhao Y, Yang G, Huang L. Systematic review of chemical constituents in the

[10] Wang Q, Chen S, Zhang Z. Determination of polysaccharide contents in Fructus Lycii.

*Lycium chinense* fruits. Archives of Pharmacal Research. 2005;**28**:1381-1385

stress in aged mice. Journal of Ethnopharmacology. 2007;**111**:504-511

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genus *Lycium* (Solanaceae). Molecules. 2017;**22**:911

Chinese Traditional Herbal Drugs. 1991;**22**:67-68

3 College of Life Sciences, Fujian Agriculture and Forestry University, Fuzhou, China

Jianjun Chen1

**References**

Significant progress has been made in *L. barbarum* breeding compared to other emerging fruit crops, such as pawpaw (*Asimina triloba* Dunal.), quince (*Cydonia oblonga* Mill.), or blue honeysuckle (*Lonicera caerulea* L.) [42]. The *Lycium* story is that a South and North America species was naturalized in northwest China and the domestication of this species produced more than 10 landraces. Selection from the landraces resulted in the release of a series of cultivars. Its reproduction modes, cross-ability, self-incompatibility, male sterility, and phylogenetic relationships with other species have been revealed. Regeneration and transformation have been developed. From a local, traditional medicinal plant, it has now received increasing attention as an important nutraceutical and pharmaceutical crop. However, *L. barbarum* and its relatives in the genus require further attention:


With the increased recognition of their roles as functional foods, plants in the genus *Lycium* will draw more attention for systematic research. It is anticipated that the potential of this small fruit crop will be fully exploited and valuable products that benefit human health and well-being will be developed and utilized.

#### **Author details**

modes in *L. barbarum* should be further investigated as the modes are fundamentally impor-

Significant progress has been made in *L. barbarum* breeding compared to other emerging fruit crops, such as pawpaw (*Asimina triloba* Dunal.), quince (*Cydonia oblonga* Mill.), or blue honeysuckle (*Lonicera caerulea* L.) [42]. The *Lycium* story is that a South and North America species was naturalized in northwest China and the domestication of this species produced more than 10 landraces. Selection from the landraces resulted in the release of a series of cultivars. Its reproduction modes, cross-ability, self-incompatibility, male sterility, and phylogenetic relationships with other species have been revealed. Regeneration and transformation have been developed. From a local, traditional medicinal plant, it has now received increasing attention as an important nutraceutical and pharmaceutical crop. However, *L. barbarum* and

**1.** Genetic potential of other species should be exploited. Current research has been largely focused on *L. barbarum*, and to a certain extent on *L. chinense* and *L. ruthenicum*. Attention should be expanded to species from South and North America as well as Africa. Fruit constitutes of those species should be analyzed and those producing valuable compounds should be used for breeding purposes. Their reproduction modes, cross-ability, and corresponding breeding schemes should be developed. Since *L. barbarum* has been shown to be able to cross with tomato, it is believed that *L. barbarum* could be easily crossed with other *Lycium* species. With the use of other species, it is anticipated that new cultivars with more

**2.** Breeding objectives should include not only fruit size and yield but also disease and pest resistance, valuable compound content, and adaptability. Early maturity should also be important. Constitutes of fruits or leaves should be systematically analyzed, and compounds with beneficial functions or negative effects should be clearly identified. Breeding schemes should be designed to maximize production of beneficial compounds and mini-

**3.** Effective methods for breeding of *Lycium* species should be developed. Current methods are based on individual plant selection and to some extent the use of self-incompatibility or male sterility. Individual plant selection continues to be useful but it should be accompanied with *in vitro* shoot culture to produce a large number of clones for commercial production. Pure lines derived from simultaneous doubling of anther cultured haploids should be tested to determine if homozygosity is an option for improving gojiberry productivity. If not, methods for maximizing heterozygosity should be evaluated to increase fruit production. Furthermore, molecular marker technologies should be incorporated into

With the increased recognition of their roles as functional foods, plants in the genus *Lycium* will draw more attention for systematic research. It is anticipated that the potential of this small fruit crop will be fully exploited and valuable products that benefit human health and

tant for new cultivar development as documented in this article.

desirable traits could be developed for commercial production.

its relatives in the genus require further attention:

14 Breeding and Health Benefits of Fruit and Nut Crops

mize those with negative effects.

breeding schemes to increase breeding efficiency.

well-being will be developed and utilized.

Jianjun Chen1 \*, ChihCheng T. Chao<sup>2</sup> and Xiangying Wei1,3

\*Address all correspondence to: jjchen@ufl.edu

1 Institute of Food and Agricultural Sciences, Mid-Florida Research and Education Center, University of Florida, Apopka, FL, USA

2 USDA-ARS, Plant Genetic Resources Unit, Geneva, NY, USA

3 College of Life Sciences, Fujian Agriculture and Forestry University, Fuzhou, China

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**Chapter 2**

**Provisional chapter**

**Genetic Diversity and Breeding of Persimmon**

**Genetic Diversity and Breeding of Persimmon**

DOI: 10.5772/intechopen.74977

The genus *Diospyros,* which is distributed in tropical and subtropical regions of the world, contains hundreds of plant species. However, four species of them have commercial importance. *D. kaki* Thunb. is the most widely cultivated species of the *Diospyros* genus. Persimmon (*D. kaki* Thunb.) is grown in many parts of the world that display subtropical climate conditions. In recent years, the cultivation of persimmon has found renewed interest in various countries of the Mediterranean basin. In China (which is the origin of persimmon) and in Japan and Korea (where it is grown widely), persimmons were selected from some well-known old varieties. Recently in countries such as Italy, Spain, USA, Brazil, Turkey and Israel, persimmons were selected from new cultivars. Currently China, Japan and Korea have the big persimmon germplasm collections with a large number of varieties and other *Diospyros* species. Also, Italy, Spain, USA, Brazil, Turkey, Israel, Azerbaijan, Uzbekistan and Pakistan have constituted the germplasms by high commercial value cultivars and/or local varieties. In this chapter, we tried to provide an overview of the genetic

diversity and breeding of persimmon by combining literature with our studies.

Persimmon is fleshy fibrous subtropical and tropical, deciduous fruit belonging to *Ebenaceae* family. The Oriental persimmon (*Diospyros kaki* Thunb.) is an exotic fruit rich in vitamins, nutrients and antioxidants vital for optimum health with various medicinal and chemical uses of fruits and leaves. Its fruit is usually consumed as a fresh or dried fruit. It is believed to have originated in the mountain area of southern China and has been cultivated as an important fruit crop in China, Korea as well as in Japan for centuries [1]. It is commonly

**Keywords:** *Diospyros*, genetic resources, selection, hybridization, breeding

© 2016 The Author(s). Licensee InTech. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

© 2018 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use,

distribution, and reproduction in any medium, provided the original work is properly cited.

Turgut Yesiloglu, Berken Cimen, Meral Incesu and

Turgut Yesiloglu, Berken Cimen, Meral Incesu and

Additional information is available at the end of the chapter

Additional information is available at the end of the chapter

http://dx.doi.org/10.5772/intechopen.74977

Bilge Yilmaz

Bilge Yilmaz

**Abstract**

**1. Introduction**


#### **Chapter 2 Provisional chapter**

#### **Genetic Diversity and Breeding of Persimmon Genetic Diversity and Breeding of Persimmon**

DOI: 10.5772/intechopen.74977

Turgut Yesiloglu, Berken Cimen, Meral Incesu and Bilge Yilmaz Turgut Yesiloglu, Berken Cimen, Meral Incesu and Bilge Yilmaz

Additional information is available at the end of the chapter Additional information is available at the end of the chapter

http://dx.doi.org/10.5772/intechopen.74977

#### **Abstract**

[67] Cao Y, Jia Y, Luo Q. Selection of Gojiberry resistance to *Fusarium graminearum* through in vitro tissue culture. In: Bei S, editor. Ningxia Gojiberry Research: Ningxia People's

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20 Breeding and Health Benefits of Fruit and Nut Crops

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1955;**9**:347-349

2006;**103**:1359-1363

The genus *Diospyros,* which is distributed in tropical and subtropical regions of the world, contains hundreds of plant species. However, four species of them have commercial importance. *D. kaki* Thunb. is the most widely cultivated species of the *Diospyros* genus. Persimmon (*D. kaki* Thunb.) is grown in many parts of the world that display subtropical climate conditions. In recent years, the cultivation of persimmon has found renewed interest in various countries of the Mediterranean basin. In China (which is the origin of persimmon) and in Japan and Korea (where it is grown widely), persimmons were selected from some well-known old varieties. Recently in countries such as Italy, Spain, USA, Brazil, Turkey and Israel, persimmons were selected from new cultivars. Currently China, Japan and Korea have the big persimmon germplasm collections with a large number of varieties and other *Diospyros* species. Also, Italy, Spain, USA, Brazil, Turkey, Israel, Azerbaijan, Uzbekistan and Pakistan have constituted the germplasms by high commercial value cultivars and/or local varieties. In this chapter, we tried to provide an overview of the genetic diversity and breeding of persimmon by combining literature with our studies.

**Keywords:** *Diospyros*, genetic resources, selection, hybridization, breeding

#### **1. Introduction**

Persimmon is fleshy fibrous subtropical and tropical, deciduous fruit belonging to *Ebenaceae* family. The Oriental persimmon (*Diospyros kaki* Thunb.) is an exotic fruit rich in vitamins, nutrients and antioxidants vital for optimum health with various medicinal and chemical uses of fruits and leaves. Its fruit is usually consumed as a fresh or dried fruit. It is believed to have originated in the mountain area of southern China and has been cultivated as an important fruit crop in China, Korea as well as in Japan for centuries [1]. It is commonly

© 2016 The Author(s). Licensee InTech. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. © 2018 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

cultivated in warm regions of the world including China, Korea, Japan, Brazil, Spain, Turkey, Italy and Israel. The world's total persimmon production was low between 1961 (990,079 tons) and 1993 (1,290,971 tons). However, persimmon production has increased since 1995. Total persimmon production of the world was 5,190,624 tons in 2014 (**Table 1**). Similarly, while persimmon acreage was low from 1961 to 1993, the acreage expanded rapidly increasing from 262,039 ha in 1993 to 1,028,051 ha in 2014 [2].

As represented in **Table 2**, China is the highest producer with a production of about 3,730,800 tonnes. This amount of production accounts for about 71.88% of the world's production. It is followed by Korea (428,363 tonnes), Spain (245,000 tonnes), Japan (240,600 tonnes), Brazil (182,290 tonnes), Azerbaijan (140,405 tonnes), Taiwan (72,674 tonnes), Uzbekistan (66,000 tonnes), Italy (39,149 tonnes), Israel (36,592 tonnes) and Turkey (33,470 tonnes) [2, 3]. The variation of production amounts of the countries differed by years. As is presented in **Table 2**, China's persimmon production has steadily increased from 495,000 tonnes in 1961 to 3,730,800 tonnes in 2014. Similarly, Korea's production increased by 32.28 times and has reached 428,363 tonnes. Spain has shown the maximum rate of increase with the production increasing from 591 tonnes in 1991 to 245,000 tonnes in 2014. In Spain, about 20 years ago, persimmon was grown for local consumption. The selection of cultivar 'Rojo Brillante' (PVA) having high fruit yield and quality and the application of the technique for removing astringency without losing fruit firmness led to the expansion of the persimmon culture in the 1990s [4]. Israel and Turkey that have lower production have significantly increased their production (respectively, 36,592 and 33,470 tonnes). On the other hands, while Japan's and Italy's production were 393,500 tonnes and 70,740 tonnes in 1961, they have dramatically decreased to 240,600 and 39,149 tonnes in 2014.

About 5% of the world's total persimmon production is exported as fresh fruit. The rest of persimmon is consumed in the internal market, and a good part of production is dried or processed. Although Azerbaijan is the sixth persimmon producer country, it is the first fresh fruit persimmon exporting country in the world. In 2013, Azerbaijan exported 95,118 tonnes of persimmon fruits. It is followed by Spain (40,121 tonnes), China mainland (35,799 tonnes), Israel (13,084 tonnes) and Poland (12,142 tonnes) (**Table 3**). Further, Russia (114,596 tonnes), Kazakhstan (58,464 tonnes) and Germany (30,233 tonnes) are the largest persimmon-importing countries in the World (**Table 4**).

Persimmons have proved to be highly adaptable to a wide range of climate conditions, ranging from subtropical coastal regions to mild coastal areas and warm inland temperate areas. Persimmons do best in areas that have moderate winters and relatively mild summers. Generally, astringent varieties prefer cooler climatic conditions than non-astringent varieties. Non-astringent cultivars require warmer growing conditions. If the cultivars of nonastringent types are grown in cooler regions, the fruit flesh may not lose all of its astringency and have lower sugar content at harvest. They can tolerate temperatures of −18°C when fully dormant. Persimmons need only a short chilling period (about 100–400 h below 7.2°C). The chilling requirement of non-astringent varieties is lower than that of astringent varieties. If the dormancy is broken or the chilling requirement of the variety has been supplied early by the warming climate, the new shoots can be damaged by a late spring frost. The leaves are killed by −3.3°C when growing. However, persimmons generally bloom late in the spring

**Year** 1961 1962 1963 1964 1965 1966 1967 1968 1969 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981

506,000

39,254

—

260,500

37,598

—

7313

—

62,800

5600

—

919,207

23

560,400

31,837

—

265,200

39,958

—

6238

—

61,100

3400

—

968,198

506,500

33,386

—

263,900

40,385

—

5267

—

58,500

2500

—

910,503

570,000

29,984

—

287,000

36,340

—

4463

—

57,800

1900

—

987,547

Genetic Diversity and Breeding of Persimmon http://dx.doi.org/10.5772/intechopen.74977

> 525,000

30,138

—

275,400

23,801

—

4422

—

62,100

2400

—

923,321

530,000

16,946

—

264,100

22,364

—

3826

—

60,690

1700

—

899,686

528,000

20,890

—

274,700

22,114

—

2843

—

60,000

—

—

908,607

538,000

41,928

—

283,600

20,446

—

2745

—

63,500

—

—

950,249

535,000

32,284

—

347,200

19,374

—

2515

—

59,000

—

—

995,403

480,000

31,115

—

306,900

25,188

—

2359

—

61,700

—

—

907,292

427,000

22,887

—

303,200

21,558

—

2411

—

59,200

—

—

836,286

455,000

30,310

—

342,700

21,659

—

2341

—

59,600

—

—

911,635

414,000

33,854

—

444,100

20,849

—

2152

—

74,000

—

—

988,980

470,000

34,579

—

450,100

20,427

—

2333

—

73,300

—

—

1,050,764

475,000

23,609

—

504,400

20,037

—

2203

—

73,600

—

—

1,098,869

495,000

22,075

—

419,300

19,586

—

2101

—

71,400

—

—

1,029,482

515,000

23,510

—

346,400

19,988

—

1997

—

72,000

—

—

978,915

465,000

23,602

—

464,000

17,198

—

2340

—

73,500

—

—

1,045,660

475,000

14,114

—

383,500

16,239

—

1804

—

75,200

—

—

965,877

440,000

16,594

—

322,500

16,005

—

2109

—

72,830

—

—

870,058

495,000

13,271

—

393,500

15,298

—

2250

—

70,740

—

—

990,079

**China**

**Korea**

**Spain**

**Japan**

**Brazil**

**Azerbaijan**

**Taiwan**

**Uzbekistan**

**Italy**

**Israel**

**Turkey\***

**World**


cultivated in warm regions of the world including China, Korea, Japan, Brazil, Spain, Turkey, Italy and Israel. The world's total persimmon production was low between 1961 (990,079 tons) and 1993 (1,290,971 tons). However, persimmon production has increased since 1995. Total persimmon production of the world was 5,190,624 tons in 2014 (**Table 1**). Similarly, while persimmon acreage was low from 1961 to 1993, the acreage expanded rap-

As represented in **Table 2**, China is the highest producer with a production of about 3,730,800 tonnes. This amount of production accounts for about 71.88% of the world's production. It is followed by Korea (428,363 tonnes), Spain (245,000 tonnes), Japan (240,600 tonnes), Brazil (182,290 tonnes), Azerbaijan (140,405 tonnes), Taiwan (72,674 tonnes), Uzbekistan (66,000 tonnes), Italy (39,149 tonnes), Israel (36,592 tonnes) and Turkey (33,470 tonnes) [2, 3]. The variation of production amounts of the countries differed by years. As is presented in **Table 2**, China's persimmon production has steadily increased from 495,000 tonnes in 1961 to 3,730,800 tonnes in 2014. Similarly, Korea's production increased by 32.28 times and has reached 428,363 tonnes. Spain has shown the maximum rate of increase with the production increasing from 591 tonnes in 1991 to 245,000 tonnes in 2014. In Spain, about 20 years ago, persimmon was grown for local consumption. The selection of cultivar 'Rojo Brillante' (PVA) having high fruit yield and quality and the application of the technique for removing astringency without losing fruit firmness led to the expansion of the persimmon culture in the 1990s [4]. Israel and Turkey that have lower production have significantly increased their production (respectively, 36,592 and 33,470 tonnes). On the other hands, while Japan's and Italy's production were 393,500 tonnes and 70,740 tonnes in 1961, they have dramatically decreased to 240,600 and 39,149 tonnes in 2014.

About 5% of the world's total persimmon production is exported as fresh fruit. The rest of persimmon is consumed in the internal market, and a good part of production is dried or processed. Although Azerbaijan is the sixth persimmon producer country, it is the first fresh fruit persimmon exporting country in the world. In 2013, Azerbaijan exported 95,118 tonnes of persimmon fruits. It is followed by Spain (40,121 tonnes), China mainland (35,799 tonnes), Israel (13,084 tonnes) and Poland (12,142 tonnes) (**Table 3**). Further, Russia (114,596 tonnes), Kazakhstan (58,464 tonnes) and Germany (30,233 tonnes) are the largest persimmon-importing

Persimmons have proved to be highly adaptable to a wide range of climate conditions, ranging from subtropical coastal regions to mild coastal areas and warm inland temperate areas. Persimmons do best in areas that have moderate winters and relatively mild summers. Generally, astringent varieties prefer cooler climatic conditions than non-astringent varieties. Non-astringent cultivars require warmer growing conditions. If the cultivars of nonastringent types are grown in cooler regions, the fruit flesh may not lose all of its astringency and have lower sugar content at harvest. They can tolerate temperatures of −18°C when fully dormant. Persimmons need only a short chilling period (about 100–400 h below 7.2°C). The chilling requirement of non-astringent varieties is lower than that of astringent varieties. If the dormancy is broken or the chilling requirement of the variety has been supplied early by the warming climate, the new shoots can be damaged by a late spring frost. The leaves are killed by −3.3°C when growing. However, persimmons generally bloom late in the spring

idly increasing from 262,039 ha in 1993 to 1,028,051 ha in 2014 [2].

22 Breeding and Health Benefits of Fruit and Nut Crops

countries in the World (**Table 4**).


**Year** 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 **2014**

**3,730,800** Source: FAO, 2017. \*

**Table 1.**

World persimmon production amount (tonnes).

**428,363**

**245,000**

**240,600** Represents data from Turkish Statistical Institute (TUIK), 2017.

**182,290**

**140,405**

**72,674**

**66,000**

**39,149**

**36,592**

**33,470**

5,190,624

Genetic Diversity and Breeding of Persimmon http://dx.doi.org/10.5772/intechopen.74977 25

3,538,823

351,990

242,800

214,700

173,169

143,106

63,694

75,000

41,858

35,692

33,232

4,890,000

3,417,586

401,049

212,300

253,800

158,241

140,082

81,894

56,000

51,165

31,292

32,392

4,813,100

3,187,239

390,820

159,400

207,500

154,625

146,084

90,100

53,400

50,347

29,271

28,295

4,476,946

2,875,600

390,630

125,280

189,400

167,215

142,188

58,401

38,000

48,165

28,201

26,277

4,070,375

2,834,165

416,705

100,200

258,000

171,555

135,549

37,032

40,000

51,593

32,291

25,281

4,084,050

2,710,988

430,521

95,400

266,600

173,297

132,179

33,899

31,000

51,600

45,350

24,302

3,977,744

2,574,143

395,614

67,000

244,800

159,851

128,407

32,962

28,000

52,500

37,347

23,713

3,727,432

2,320,346

352,822

63,000

232,700

168,274

124,485

26,395

27,213

53,100

24,606

19,297

3,399,970

2,185,041

363,822

51,950

285,900

164,849

108,965

27,111

21,000

51,332

48,000

18,000

3,314,731

1,998,214

299,046

45,800

232,500

162,288

48,089

36,177

19,000

57,110

38,700

17,000

2,943,525

1,795,110

249,207

45,200

265,000

158,131

114,899

38,247

17,000

47,000

40,100

15,000

2,776,510

**China**

**Korea**

**Spain**

**Japan**

**Brazil**

**Azerbaijan**

**Taiwan**

**Uzbekistan**

**Italy**

**Israel**

**Turkey\***

**World**


**Year** 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002

1,740,591

281,143

36,050

269,300

141,364

104,800

34,747

16,500

54,170

35,700

15,000

2,720,098

1,584,660

270,338

34,500

281,800

105,000

86,000

26,169

16,000

48,240

14,900

13,500

2,473,149

1,591,906

287,847

33,000

278,800

63,300

70,300

23,891

16,000

42,450

14,186

12,000

2,427,646

1,481,327

273,846

29,469

286,000

64,096

60,300

25,754

—

40,769

14,000

11,500

2,280,753

1,314,136

260,671

21,842

260,100

60,423

47,900

18,962

—

62,000

17,400

10,500

2,068,143

1,075,417

239,570

18,688

301,700

52,198

—

19,018

—

59,800

16,800

10,000

1,787,234

1,025,219

210,766

14,757

240,500

52,534

—

15,811

—

67,800

18,200

9400

1,649,285

969,363

194,585

10,826

254,100

51,685

—

16,440

—

61,300

11,000

9200

1,573,421

826,870

167,471

6895

302,200

55,406

—

15,100

—

48,999

13,800

9300

1,440,701

789,113

116,070

2963

241,900

48,086

—

16,380

—

56,753

15,780

10,000

1,290,971

724,329

155,111

591

307,700

46,611

—

14,636

—

66,546

17,390

10,000

1,336,383

641,576

109,722

591

248,800

47,662

—

14,935

—

61,810

16,200

10,000

1,144,666

624,773

95,758

0

285,700

46,712

—

15,457

—

68,770

17,200

—

1,157,241

650,283

113,403

—

268,100

46,836

—

16,404

—

72,920

16,000

—

1,185,538

732,921

98,337

—

287,600

45,745

—

15,694

—

69,170

7000

—

1,257,684

820,000

75,677

—

290,200

45,000

—

11,811

—

69,820

19,200

—

1,332,767

750,000

98,906

—

291,300

43,488

—

10,626

—

69,700

11,900

—

1,277,032

680,000

97,031

—

289,700

43,658

—

10,945

—

56,200

8100

—

1,186,165

24 Breeding and Health Benefits of Fruit and Nut Crops

608,000

68,812

—

297,400

41,915

—

9525

—

78,400

13,900

—

1,118,477

553,000

91,052

—

309,900

45,298

—

7850

—

76,700

12,700

—

1,096,858

482,000

57,807

—

333,700

38,396

—

6621

—

65,400

8100

—

992,416

**China**

**Korea**

**Spain**

**Japan**

**Brazil**

**Azerbaijan**

**Taiwan**

**Uzbekistan**

**Italy**

**Israel**

**Turkey\***

**World**

**Table 1.** World persimmon production amount (tonnes).


**Table 2.** Persimmon production amount (ton), planted area (ha) and yield (kg ha−1) in 2014.

(mid-April) to avoid spring frosts. On the other hand, persimmon has been damaged by earlyautumn frosts. Early-autumn frosts can lead to skin blemishes on fruit and early defoliation. Persimmon does not tolerate wind. It does not provide a good fruit yield and quality, if strong winds occur during the growing season. Fruit is also prone to wind rub from leaves and branches causing skin blemish on fruit. Windbreakers can be used to reduce the wind speed.

Full sun with some air movement is recommended for persimmon trees in inland areas, although they will tolerate some partial shade. But trees do not produce well in the high summer heat of desert regions, which sunburn the bark.

Kaki persimmons are drought tolerant. Persimmon trees can withstand drought, but fruit yield and quality (especially size) are reduced. Also, adequate moisture in the soil is required to produce sufficient shoot growth and formation of flower buds for next year's crop. The trees should be irrigated during dry periods.

to water-logging should be avoided. The preferred soil pH for optimum tree growth is in the range of 6.0–7.5. However, persimmon trees can tolerate a wider variety of conditions than

Pest and disease problems: protection of fruits from bats and birds are required. Fruit flies are the potential problem as are aphids and mealybugs. Persimmon trees are also susceptible to

collar rot, thus keeping mulch clear of the trunk is required.

**Table 4.** Top 10 countries with highest persimmon imports in 2013.

**Rank Country Amount (tonnes)**

Genetic Diversity and Breeding of Persimmon http://dx.doi.org/10.5772/intechopen.74977 27

**World total 232,247**

**Rank Country Amount (tonnes)**

**World total 323,858**

 Russian Federation 114,596 Kazakhstan 58,464 Germany 30,233 Belarus 14,788 France 12,929 Italy 11,427 Lithuania 10,177 Poland 9715 Thailand 6997 Canada 6407

**Table 3.** Top 10 countries with highest persimmon exports in 2013.

 Azerbaijan 95,118 Spain 40,121 China, mainland 35,799 Israel 13,084 Poland 12,142 Lithuania 9057 Republic of Korea 7379 Netherlands 6957 Georgia 6781 South Africa 5809

most fruit trees.

The persimmons trees can grow well on a wide range of soil types but do best in deep, welldrained loam soils with a good supply of organic matter. Heavy clay loam soils that are prone


**Table 3.** Top 10 countries with highest persimmon exports in 2013.


**Table 4.** Top 10 countries with highest persimmon imports in 2013.

(mid-April) to avoid spring frosts. On the other hand, persimmon has been damaged by earlyautumn frosts. Early-autumn frosts can lead to skin blemishes on fruit and early defoliation. Persimmon does not tolerate wind. It does not provide a good fruit yield and quality, if strong winds occur during the growing season. Fruit is also prone to wind rub from leaves and branches causing skin blemish on fruit. Windbreakers can be used to reduce the wind speed. Full sun with some air movement is recommended for persimmon trees in inland areas, although they will tolerate some partial shade. But trees do not produce well in the high sum-

TUİK [3]. Turkish Statistic Council records.

**World total production 5,190,624 1,028,051 5049.5**

**Table 2.** Persimmon production amount (ton), planted area (ha) and yield (kg ha−1) in 2014.

**Rank Country Production in tonnes Area (ha) Yield (kg/ha)** China, mainland 3,730,800 931,907 4003.4 Republic of Korea 428,363 27,988 15,305.2 Spain 245,000 13,370 18,324.6 Japan 240,600 21,300 11,295.8 Brazil 182,290 8323 21,902.0 Azerbaijan 140,405 8712 16,116.3 China, Taiwan Province of 72,764 5263 13,825.6 Uzbekistan 66,000 4218 15,647.0 Italy 39,149 2531 15,467.8 Israel 36,592 1374 26,631.7 Turkey\* 33,470 2062 16,232.6 New Zealand 2600 164 15,853.7 Iran (Islamic Republic of) 2452 275 8926.6 Nepal 1918 288 6667.4 Australia 715 86 8272.1 Slovenia 801 70 11,442.9 Mexico 175 18 9722.2 Chile — 102 —

Kaki persimmons are drought tolerant. Persimmon trees can withstand drought, but fruit yield and quality (especially size) are reduced. Also, adequate moisture in the soil is required to produce sufficient shoot growth and formation of flower buds for next year's crop. The

The persimmons trees can grow well on a wide range of soil types but do best in deep, welldrained loam soils with a good supply of organic matter. Heavy clay loam soils that are prone

mer heat of desert regions, which sunburn the bark.

trees should be irrigated during dry periods.

FAO stat [2]. http://www.faostat.com.\*

26 Breeding and Health Benefits of Fruit and Nut Crops

to water-logging should be avoided. The preferred soil pH for optimum tree growth is in the range of 6.0–7.5. However, persimmon trees can tolerate a wider variety of conditions than most fruit trees.

Pest and disease problems: protection of fruits from bats and birds are required. Fruit flies are the potential problem as are aphids and mealybugs. Persimmon trees are also susceptible to collar rot, thus keeping mulch clear of the trunk is required.

#### **2. Origin and history**

Zeven and Zhukovsky [5] suggested that persimmon (*D. kaki*) has a primary center of genetic origin in the mountains of central China and a secondary center in Japan. Persimmon cultivation in China began more than 2000 years ago, and it is also scientifically known as *D. chinensis*. In China, it is found wild at altitudes up to 6000–8000 ft. [6]. It spread from China to Korea and to Japan many years ago. Since from prehistoric times, permission is consumed as food source in these countries. There are some trees that are 400–500 years old. It was imported in Europe (South France) for the first time in 1760. Thereafter it spread to the Mediterranean coast (Italy, Spain, Greece, Turkey and Algeria). The persimmon plant was introduced in North America (California, Florida), South America (Brazil) and Australia in the mid-1800s. Early in the fourteenth century, the explorer Marco Polo recorded the Chinese trade in persimmons [7]. Its cultivation has recent traditions in western countries where it is present only since the second half of the nineteenth century. Currently, persimmon is one of the most important fruit crops in Asian countries and, there is also steady increase in its production in some European countries.

*(D. oleifera*, *D. glandulosa*, *D. confertiflora*, *D. discolorare*, *D. ehretioides*, *D. lycioides*, *D. mollis*, *D. rhodocalyx* and *D. sumatrana*) are *2*n = 30, except for *2*n = 60 for *D. rhombifolia* and *2*n = 90 for *D. ebenum* [13–15]. *D. kaki* L. is a hexaploid (2n = 6x = 90). However, octoploid (2n = 8x = 120) cultivars such as Hasshu and nonaploid (*2*n = 9x =135) cultivars such as 'Hiratanenashi' and 'Tonewase' have also been reported [14–16]. On the other hand, *D. virginiana* has two karyo-

Genetic Diversity and Breeding of Persimmon http://dx.doi.org/10.5772/intechopen.74977 29

Persimmon fruit is highly astringent due to soluble tannins in the vacuoles of the fruit flesh. However, some cultivars lose astringency naturally on the tree as fruit ripens, while others retain astringency until maturity. Therefore, persimmons are classified into two major groups (based on the presence or absence of astringency in the fruit at maturity) as astringent (A) and non-astringent or sweet (NA) cultivars. Water-soluble tannins which cause astringency in the flesh of astringent types decrease as the fruit softens and becomes edible. However, astringency can be removed by various chemical treatments. Carbon dioxide gas or alcohol can be used to remove astringency, while the fruit remains firm. If ethylene is used for removing astringency, the fruit softens very quickly. Fruit of the non-astringent types naturally loses astringency, while the fruit is still firm. Thus, the fruit of non-astringent types is edible either

Each group can be further subdivided, based on their response to pollination [18]. The amount of dark flesh coloration around the seeds varies in cultivars and changes in flesh color are related to seed formation, not pollination. In pollination variant types (PV), the flesh is dark and streaked around the seeds, but clear orange when seedless. When pollination is poor and only one or several seeds are formed, a dark area develops around the seeds, but the remaining flesh is light colored. The pollination variant types include cultivars that are astringent when they have several seeds or seedless (PVA), as well as partially or totally non-astringent when they have only one or a few seeds (PVNA). Also, in astringent cultivars of the pollination variant type, fruit which has a great degree of the dark flesh is non-astringent even when

Pollination constant (PC) types lack the dark streaking regardless of seed formation. The flesh color of pollination constant astringent (PCA) cultivars is not influenced by pollination and it does not develop dark flesh around the seeds. Pollination constant non-astringent (PCNA) persimmons are always edible when still firm, regardless of whether or not pollina-

PVA types can vary to either PCA or PVNA depending on several situations. If PVA type does not have any seed for some reason or when PVA persimmon varieties are cultivated without pollinators, the fruit has clear orange flesh and remains astringent (PCA) such as the Spainish variety 'Rojo Brillante' and Japanese variety 'Hiratanenashi'. Similarly, when PVA type has enough seeds (usually four or five) after pollination, the fruit has a great degree of dark flesh

and loses astringency (PVNA) such as 'Nishimura Wase'.

types with *2*n = 60 and 90 [15, 17], while *D. lotus* is diploid (*2*n = 2x =30) [14].

**4. Pomological classification**

the flesh is firm or soft [10, 12, 18].

the fruit flesh is firm [10, 18].

tion has occurred.

#### **3. Botany of persimmon**

The genus *Diospyros* contains hundreds of plant species and are distributed in the tropical and subtropical regions of the world. Four species of them have commercial importance. *D. kaki* L. is the most widely cultivated species of the *Diospyros* genus. *D. kaki* is also known as the persimmon, Japanese persimmon, Oriental persimmon, Japanese persimmon, Kaki, Asian persimmon. It has been reported that wild type *D. kaki* exists in the forests of China [8, 9]. The other species are *D. lotus* L. (the date plum), *D. viriginiaina* L. (native American persimmon) and *D. oleifera* Cheng [10].

The origin of *D. kaki* and its relationship to other *Diospyros* species is not well understood. The persimmon culture was known to occur in the fifth or sixth century in China [9]. In addition to *D. kaki, D. lotus* and *D. oleifera* also have been cultivated as fruit crop. *D. lotus* has been consumed as a fresh as well as dried fruit, and it is a source for tannin [9, 10]. Another important species known as a fruit crop is *D. virginiana*, of the eastern United States origin. This species which is consumed as fresh and processed is grown on a much smaller scale and is not yet considered a commercial crop [11]. These species are quite important as horticultural crops among the *Diospyros* species of temperate origin. On the other hands, *D. rhombifolia* originated from China is an ornamental plant which bears tiny attractive-colored fruit on a dwarf tree [9]. There are other species such as *D. digyna* (black sapote), *D. discolor* and *D. decandra* that have originated in the tropics and subtropics and produce edible fruits.

In the genus *Diospyros,* there are species and varieties having diploid (2n = 2x = 30), tetraploid (2n = 4x = 60), hexaploid (2n = 6x = 90), nonaploid *(2n* = 9x =135) and dodecaploid (2n = 12x = 180) chromosome number. Therefore, it is thought that the basic chromosome number of the genus *Diospyros* is 15 [9, 12]. The chromosome numbers of some wild species of genus *Diospyros*  *(D. oleifera*, *D. glandulosa*, *D. confertiflora*, *D. discolorare*, *D. ehretioides*, *D. lycioides*, *D. mollis*, *D. rhodocalyx* and *D. sumatrana*) are *2*n = 30, except for *2*n = 60 for *D. rhombifolia* and *2*n = 90 for *D. ebenum* [13–15]. *D. kaki* L. is a hexaploid (2n = 6x = 90). However, octoploid (2n = 8x = 120) cultivars such as Hasshu and nonaploid (*2*n = 9x =135) cultivars such as 'Hiratanenashi' and 'Tonewase' have also been reported [14–16]. On the other hand, *D. virginiana* has two karyotypes with *2*n = 60 and 90 [15, 17], while *D. lotus* is diploid (*2*n = 2x =30) [14].

#### **4. Pomological classification**

**2. Origin and history**

28 Breeding and Health Benefits of Fruit and Nut Crops

some European countries.

and *D. oleifera* Cheng [10].

**3. Botany of persimmon**

Zeven and Zhukovsky [5] suggested that persimmon (*D. kaki*) has a primary center of genetic origin in the mountains of central China and a secondary center in Japan. Persimmon cultivation in China began more than 2000 years ago, and it is also scientifically known as *D. chinensis*. In China, it is found wild at altitudes up to 6000–8000 ft. [6]. It spread from China to Korea and to Japan many years ago. Since from prehistoric times, permission is consumed as food source in these countries. There are some trees that are 400–500 years old. It was imported in Europe (South France) for the first time in 1760. Thereafter it spread to the Mediterranean coast (Italy, Spain, Greece, Turkey and Algeria). The persimmon plant was introduced in North America (California, Florida), South America (Brazil) and Australia in the mid-1800s. Early in the fourteenth century, the explorer Marco Polo recorded the Chinese trade in persimmons [7]. Its cultivation has recent traditions in western countries where it is present only since the second half of the nineteenth century. Currently, persimmon is one of the most important fruit crops in Asian countries and, there is also steady increase in its production in

The genus *Diospyros* contains hundreds of plant species and are distributed in the tropical and subtropical regions of the world. Four species of them have commercial importance. *D. kaki* L. is the most widely cultivated species of the *Diospyros* genus. *D. kaki* is also known as the persimmon, Japanese persimmon, Oriental persimmon, Japanese persimmon, Kaki, Asian persimmon. It has been reported that wild type *D. kaki* exists in the forests of China [8, 9]. The other species are *D. lotus* L. (the date plum), *D. viriginiaina* L. (native American persimmon)

The origin of *D. kaki* and its relationship to other *Diospyros* species is not well understood. The persimmon culture was known to occur in the fifth or sixth century in China [9]. In addition to *D. kaki, D. lotus* and *D. oleifera* also have been cultivated as fruit crop. *D. lotus* has been consumed as a fresh as well as dried fruit, and it is a source for tannin [9, 10]. Another important species known as a fruit crop is *D. virginiana*, of the eastern United States origin. This species which is consumed as fresh and processed is grown on a much smaller scale and is not yet considered a commercial crop [11]. These species are quite important as horticultural crops among the *Diospyros* species of temperate origin. On the other hands, *D. rhombifolia* originated from China is an ornamental plant which bears tiny attractive-colored fruit on a dwarf tree [9]. There are other species such as *D. digyna* (black sapote), *D. discolor* and *D. decandra* that have

In the genus *Diospyros,* there are species and varieties having diploid (2n = 2x = 30), tetraploid (2n = 4x = 60), hexaploid (2n = 6x = 90), nonaploid *(2n* = 9x =135) and dodecaploid (2n = 12x = 180) chromosome number. Therefore, it is thought that the basic chromosome number of the genus *Diospyros* is 15 [9, 12]. The chromosome numbers of some wild species of genus *Diospyros* 

originated in the tropics and subtropics and produce edible fruits.

Persimmon fruit is highly astringent due to soluble tannins in the vacuoles of the fruit flesh. However, some cultivars lose astringency naturally on the tree as fruit ripens, while others retain astringency until maturity. Therefore, persimmons are classified into two major groups (based on the presence or absence of astringency in the fruit at maturity) as astringent (A) and non-astringent or sweet (NA) cultivars. Water-soluble tannins which cause astringency in the flesh of astringent types decrease as the fruit softens and becomes edible. However, astringency can be removed by various chemical treatments. Carbon dioxide gas or alcohol can be used to remove astringency, while the fruit remains firm. If ethylene is used for removing astringency, the fruit softens very quickly. Fruit of the non-astringent types naturally loses astringency, while the fruit is still firm. Thus, the fruit of non-astringent types is edible either the flesh is firm or soft [10, 12, 18].

Each group can be further subdivided, based on their response to pollination [18]. The amount of dark flesh coloration around the seeds varies in cultivars and changes in flesh color are related to seed formation, not pollination. In pollination variant types (PV), the flesh is dark and streaked around the seeds, but clear orange when seedless. When pollination is poor and only one or several seeds are formed, a dark area develops around the seeds, but the remaining flesh is light colored. The pollination variant types include cultivars that are astringent when they have several seeds or seedless (PVA), as well as partially or totally non-astringent when they have only one or a few seeds (PVNA). Also, in astringent cultivars of the pollination variant type, fruit which has a great degree of the dark flesh is non-astringent even when the fruit flesh is firm [10, 18].

Pollination constant (PC) types lack the dark streaking regardless of seed formation. The flesh color of pollination constant astringent (PCA) cultivars is not influenced by pollination and it does not develop dark flesh around the seeds. Pollination constant non-astringent (PCNA) persimmons are always edible when still firm, regardless of whether or not pollination has occurred.

PVA types can vary to either PCA or PVNA depending on several situations. If PVA type does not have any seed for some reason or when PVA persimmon varieties are cultivated without pollinators, the fruit has clear orange flesh and remains astringent (PCA) such as the Spainish variety 'Rojo Brillante' and Japanese variety 'Hiratanenashi'. Similarly, when PVA type has enough seeds (usually four or five) after pollination, the fruit has a great degree of dark flesh and loses astringency (PVNA) such as 'Nishimura Wase'.

## **5. Commercial and recently improved persimmon varieties**

The fruits of two species (*D. kaki* L. and *D. virginiana*) in the genus *Diospyros* have commercial importance. In China, using native persimmon germplasm, several common persimmon varieties were developed. However, they all are PCA with the exception of 'Luo Tian Shi' [19, 20]. Persimmon is the main species cultured for edible fruit production in northern China. Recently, 'Jirou', 'Youhou', 'Taishuu' and 'Fuyu' among PCNA cultivars are gaining popularity. Persimmon growing regions are also spreading widely in Japan and Korea, thus some old well-known Persimmon varieties which were still being produced were selected from these countries. Persimmon has been a major fruit crop in Japan for many years [21] and for Japanese persimmon commercial production, 'Fuyu' (PCNA), 'Hiratanenashi' (PVA) and 'Tonewase' have been the three important cultivars. About 57% of the total area is devoted to these varieties [22]. Other varieties growing in Japan are 'Kosyu Hyakume', 'Matsumotowase Fuyu', 'Early ripening Jiro', 'Ichidagaki', 'Jiro Dojohachiya' and 'Taishu'. However, newly released cultivars such as 'Reigyoku' and 'Taiho' are also available. In Korea, non-astringent varieties have increased, while astringent varieties have decreased. In the recent years, amount of production of non-astringent varieties are higher than those of astringent varieties. Major cultivar of non-astringent type is 'Fuyu', which accounts for almost 82% in total production of persimmon, and 'Jiro' with 9.8% [23].

In Taiwan, 'Suzchou', 'Niouhsin' and 'Shihshih' local PCA are the major commercial varieties used. 'Fuyu' and' Jiro' are the main PCNA varieties [24]. Countries such as Azerbaijan and Uzbekistan focused on local astringent cultivars. In Brazil, the most cultivated persimmon cultivars include 'Rama Forte' and 'Giombo', which belong to the PVA group, 'Taubate' which is continually astringent with yellow flesh either with or without seeds (PCA), and 'Fuyu', which belongs to the PCNA group [25, 26].

a long time [28]. In Italy, almost 90% of the persimmon production is Kaki Tipo (PVNA), the rest of production is other PVNA varieties (Vainiglia, Mercatelli and Moro) and PCNA culti-

Genetic Diversity and Breeding of Persimmon http://dx.doi.org/10.5772/intechopen.74977 31

Israel has its own cultivar, Triumph (**Figure 2**) which is sold under the name of Sharon fruit, and it is planted on 95% of the total area devoted to persimmon [27]. Also, persimmon production in South Africa is based on Triumph [30]. In Turkey, a great amount of production is PCA and PVNA varieties, which are selected from Turkey. However, recently introduced PCNA cultivars such as Fuyu, Hana Fuyu, Jiro and Izu have become popular with the grow-

Persimmon originated from China, but it has been cultivated and produced mostly in Japan [31]. Persimmon has limited amount of production in the rest of the World. However, Spain, Italy, Israel and Brazil are now producing important amounts and these countries have developed their own cultivars such as 'Rojo Brillante' in Spain, 'Kaki Tipo' in Italy, 'Triumph' in Israel and 'Lama Forte' in Brazil. Recently, Australia and New Zealand have started to produce persimmon mainly for export, and the USA is also producing persimmon on a small scale.

Greene and Morris [32] indicated that germplasm collections are a source of genetic diversity to support crop improvement and botanical research as well as to support conservation efforts [33]. For the specific breeding objectives, these variations can either be created spontaneously or artificially by budwood mutations or cross breeding. The importance of germplasm can be explained by the variation of plant material. Therefore, recording and registration of genetic

resources is critical for breeders in terms of improving new varieties.

ers. The new orchards with 'Fuyu' and 'Hachiya' cultivars have been established.

vars such as Hana Fuyu, Jiro and Gosho [29].

**Figure 2.** Fruits and tree of 'Triumph' cultivar.

**6. Persimmon germplasm resources**

'Fuyu', 'Hana Fuyu' and 'Ichikikei Jiro' cultivars which are PCNA and 'Hachiya' (PCA) are commonly produced in the USA [10, 18]. In new persimmon growing countries such as New Zealand and Australia, most of the cultivation area is devoted to 'Fuyu' [27]. In Spain, the most produced cultivars are 'Rojo Brillante' (**Figure 1**) and 'Triumph' which can be stored for

**Figure 1.** Fruits and tree of 'Rojo Brillante' cultivar.

**Figure 2.** Fruits and tree of 'Triumph' cultivar.

**5. Commercial and recently improved persimmon varieties**

almost 82% in total production of persimmon, and 'Jiro' with 9.8% [23].

which belongs to the PCNA group [25, 26].

30 Breeding and Health Benefits of Fruit and Nut Crops

**Figure 1.** Fruits and tree of 'Rojo Brillante' cultivar.

The fruits of two species (*D. kaki* L. and *D. virginiana*) in the genus *Diospyros* have commercial importance. In China, using native persimmon germplasm, several common persimmon varieties were developed. However, they all are PCA with the exception of 'Luo Tian Shi' [19, 20]. Persimmon is the main species cultured for edible fruit production in northern China. Recently, 'Jirou', 'Youhou', 'Taishuu' and 'Fuyu' among PCNA cultivars are gaining popularity. Persimmon growing regions are also spreading widely in Japan and Korea, thus some old well-known Persimmon varieties which were still being produced were selected from these countries. Persimmon has been a major fruit crop in Japan for many years [21] and for Japanese persimmon commercial production, 'Fuyu' (PCNA), 'Hiratanenashi' (PVA) and 'Tonewase' have been the three important cultivars. About 57% of the total area is devoted to these varieties [22]. Other varieties growing in Japan are 'Kosyu Hyakume', 'Matsumotowase Fuyu', 'Early ripening Jiro', 'Ichidagaki', 'Jiro Dojohachiya' and 'Taishu'. However, newly released cultivars such as 'Reigyoku' and 'Taiho' are also available. In Korea, non-astringent varieties have increased, while astringent varieties have decreased. In the recent years, amount of production of non-astringent varieties are higher than those of astringent varieties. Major cultivar of non-astringent type is 'Fuyu', which accounts for

In Taiwan, 'Suzchou', 'Niouhsin' and 'Shihshih' local PCA are the major commercial varieties used. 'Fuyu' and' Jiro' are the main PCNA varieties [24]. Countries such as Azerbaijan and Uzbekistan focused on local astringent cultivars. In Brazil, the most cultivated persimmon cultivars include 'Rama Forte' and 'Giombo', which belong to the PVA group, 'Taubate' which is continually astringent with yellow flesh either with or without seeds (PCA), and 'Fuyu',

'Fuyu', 'Hana Fuyu' and 'Ichikikei Jiro' cultivars which are PCNA and 'Hachiya' (PCA) are commonly produced in the USA [10, 18]. In new persimmon growing countries such as New Zealand and Australia, most of the cultivation area is devoted to 'Fuyu' [27]. In Spain, the most produced cultivars are 'Rojo Brillante' (**Figure 1**) and 'Triumph' which can be stored for a long time [28]. In Italy, almost 90% of the persimmon production is Kaki Tipo (PVNA), the rest of production is other PVNA varieties (Vainiglia, Mercatelli and Moro) and PCNA cultivars such as Hana Fuyu, Jiro and Gosho [29].

Israel has its own cultivar, Triumph (**Figure 2**) which is sold under the name of Sharon fruit, and it is planted on 95% of the total area devoted to persimmon [27]. Also, persimmon production in South Africa is based on Triumph [30]. In Turkey, a great amount of production is PCA and PVNA varieties, which are selected from Turkey. However, recently introduced PCNA cultivars such as Fuyu, Hana Fuyu, Jiro and Izu have become popular with the growers. The new orchards with 'Fuyu' and 'Hachiya' cultivars have been established.

#### **6. Persimmon germplasm resources**

Persimmon originated from China, but it has been cultivated and produced mostly in Japan [31]. Persimmon has limited amount of production in the rest of the World. However, Spain, Italy, Israel and Brazil are now producing important amounts and these countries have developed their own cultivars such as 'Rojo Brillante' in Spain, 'Kaki Tipo' in Italy, 'Triumph' in Israel and 'Lama Forte' in Brazil. Recently, Australia and New Zealand have started to produce persimmon mainly for export, and the USA is also producing persimmon on a small scale.

Greene and Morris [32] indicated that germplasm collections are a source of genetic diversity to support crop improvement and botanical research as well as to support conservation efforts [33]. For the specific breeding objectives, these variations can either be created spontaneously or artificially by budwood mutations or cross breeding. The importance of germplasm can be explained by the variation of plant material. Therefore, recording and registration of genetic resources is critical for breeders in terms of improving new varieties.

Currently, more than 950 cultivars of persimmon exist from the subtropical to temperate regions of China [34]. There is only 1 genus and 63 species in persimmon family and most of them are distributed in tropic and sub-tropic regions of Hainan, Yunnan, Guangdong, Guangxi and Fujian provinces in China. The 63 species originated from this genus in China. Among these species *D. kaki* Thunb., *D. oleifera* Cheng., *D. lobata* L., *D. discilir* Willd., *D. potingensis* Merr. et Chun., *D. lotus* Linn., *D. glaucifolia* Metc., *D. rhombifolia* Hemsl. and *D. morrisiana* Hance. have been cultivated as fruit crops [35]. There are 550 accessions including most cultivars native to China and some native to Japan and Korea.

Persimmon was introduced into Brazil's São Paulo state in 1890. However, its cultivation expanded around 1920 with Japanese immigration. São Paulo is the main persimmon producing state. Rio Grande do Sul state has the second largest persimmon production of Brazil. In recent years, the persimmon acreage has increased and the trend is to continue crop expansion. 'Fuyu', 'Rama Forte', 'Giombo' and 'Taubaté' are the cultivars grown in Brazil. In Azerbaijan, persimmon production is widely spread since 1998, although its history has deep roots [2]. In terms of persimmon genetic resources in Israel, only high commercial value

Genetic Diversity and Breeding of Persimmon http://dx.doi.org/10.5772/intechopen.74977 33

Although the exact date of the introduction of persimmon to Anatolia is unknown, it is clear that it dates back to rather old times [39, 40]. Persimmon was introduced to Turkey from Russia via the Black Sea region. Turkey has main persimmon species (*D. kaki*, *D. lotus* and *D. oleifera*). *D. oleifera* can be seen only in the Mediterranean region of Turkey, while *D. lotus* grows as wild in Northern Anatolia and is used as dried fruits in this region. *D. kaki* and *D. oleifera* have been introduced from other countries at least 200 years ago. During this time, continuous propagation of persimmon by its seeds resulted in genetic diversity in *D. kaki* trees due to the high heterozygosity. Therefore, in the northeastern part of Turkey, persimmon trees differ from another in terms of fruit productivity, yield, shape, size, astringency and plant growth. This diversity in persimmon population in Turkey provided a great opportunity to the breeders for selection programs. As a result, the breeders were able to identify many promising clones in different parts of Turkey. A germplasm collection in the Black Sea

First studies on persimmon in Turkey were started to introduce the foreign cultivars by the Ministry of Agriculture in 1967. Then, some selection studies were done in different parts of Turkey. After 1989, the total number of the known cultivars and types reached up to 74. Most of these varieties were introduced from Italy and some of them were from Israel, Japan, France and Pakistan especially after the attempts made by the Cukurova University, in Eastern

Selection of different genotypes was started by the Department of Horticulture of Cukurova University, by Department of Horticulture of Ondokuzmayıs University in Black Sea and by Citrus Research Institute of Antalya belonging to the Ministry of Agriculture in Western Mediterranean regions. Recently, the wider selections have been carried out especially on Black Sea coast by Citrus Research Institute of Antalya for 1 year and they presently have 43 promising candidate clones. Most even totality of the selections is astringent type. It seems to be rather difficult to find non-astringent types in Turkey. Yilmaz et al. [41] established a characterization study on persimmon genetic resources collected from Turkey. These germplasms were preserved with commercial cultivars in an *ex situ* germplasm preservation orchard located at the Cukurova University, Turkey. Persimmon genotypes were characterized based on their morphological traits. The collection comprising traditional genotypes, local accessions and also global varieties were collected from five different provinces of the Mediterranean region of Turkey where persimmon is widely produced. A total of 48 persimmon genotypes and cultivars were morphologically characterized, using 59 morphological

region in Turkey with selected promising genotypes has been established.

cultivars are collected.

Mediterranean, Turkey.

and agronomic traits (**Table 5**).

Aside from this exceptional existence of a PCNA type cultivar in China, almost all non-astringent type cultivars were developed in Japan. Historical records show that 'Zenjimaru', known to be the oldest PVNA type cultivar, was found in the beginning of the twentieth century, and that 'Gosho', was the first PCNA type cultivar, which was recorded in the seventeenth century [36]. In the beginning of the nineteenth century, 'Fuyu' and 'Jiro' were recorded as the most popular PCNA type cultivars. According to a nationwide survey on persimmon cultivars in Japan (Agricultural Research Station 1912), there were only 6 PCNA type cultivars in contrast to 401 PVNA type cultivars among more than 1000 cultivars collected from all over Japan. This means that in addition to its more recent appearance, the PCNA type probably has very narrow genetic variability. A total of 40 PCNA cultivars, including bud sports, which may cover almost all PCNA type cultivars currently existing in Japan, are now preserved at the National Institute of the Fruit Tree Science (NIFTS) in Akitsu, Hiroshima [9]. There are many astringent and PVNA local cultivars throughout Japan. The current conservation in Japan consists of approx. 600 genotypes [37].

In Korea, 233 local cultivars were collected at the branch of Experimental Station at Kim-hae during 1959–1969, and 74 superior cultivars were selected for persimmon cultivation after identifying the name of 188 cultivars among these local cultivars. In Korea, interest in persimmon cultivation is increasing and two experimental stations for persimmon have been established, the one for non-astringent persimmon was established in 1994 and the other for astringent persimmon established in 1995. In addition, a breeding program for obtaining new PCNA cultivars was started in 1995 by crosses among PCNA cultivars that were introduced from Japan. The breeding objectives in Korea are focused on obtaining superior PCNA cultivars with good eating qualities, large fruit and early ripening characteristics [9].

In Europe, persimmon is considered a secondary fruit tree species; only few countries, located in the Mediterranean area, are interested in a large-scale production.

Persimmon was introduced in Italy at the end of the ninteenth century. Later in Tuscany, the interest for this new species was increased and the genotypes were collected together with exotic and local varieties of fruit tree species (citrus, peaches and plums among others). As early as 1940, the University of Florence collected 11 accessions from the USA and France, or as local varieties and characterized them. The persimmon collection of Florence consisted of 52 cultivars and was totally destroyed by winter frost in 1985. Then a new germplasm orchard was established by introducing new accessions from Japan. A French germplasm was recorded at the beginning of the twentieth century. The Spanish collection was created in 1993 with material from Italy (54.2% of accessions) and from Spanish institutions and nurseries (45.8%) [38].

Persimmon was introduced into Brazil's São Paulo state in 1890. However, its cultivation expanded around 1920 with Japanese immigration. São Paulo is the main persimmon producing state. Rio Grande do Sul state has the second largest persimmon production of Brazil. In recent years, the persimmon acreage has increased and the trend is to continue crop expansion. 'Fuyu', 'Rama Forte', 'Giombo' and 'Taubaté' are the cultivars grown in Brazil. In Azerbaijan, persimmon production is widely spread since 1998, although its history has deep roots [2]. In terms of persimmon genetic resources in Israel, only high commercial value cultivars are collected.

Currently, more than 950 cultivars of persimmon exist from the subtropical to temperate regions of China [34]. There is only 1 genus and 63 species in persimmon family and most of them are distributed in tropic and sub-tropic regions of Hainan, Yunnan, Guangdong, Guangxi and Fujian provinces in China. The 63 species originated from this genus in China. Among these species *D. kaki* Thunb., *D. oleifera* Cheng., *D. lobata* L., *D. discilir* Willd., *D. potingensis* Merr. et Chun., *D. lotus* Linn., *D. glaucifolia* Metc., *D. rhombifolia* Hemsl. and *D. morrisiana* Hance. have been cultivated as fruit crops [35]. There are 550 accessions including most

Aside from this exceptional existence of a PCNA type cultivar in China, almost all non-astringent type cultivars were developed in Japan. Historical records show that 'Zenjimaru', known to be the oldest PVNA type cultivar, was found in the beginning of the twentieth century, and that 'Gosho', was the first PCNA type cultivar, which was recorded in the seventeenth century [36]. In the beginning of the nineteenth century, 'Fuyu' and 'Jiro' were recorded as the most popular PCNA type cultivars. According to a nationwide survey on persimmon cultivars in Japan (Agricultural Research Station 1912), there were only 6 PCNA type cultivars in contrast to 401 PVNA type cultivars among more than 1000 cultivars collected from all over Japan. This means that in addition to its more recent appearance, the PCNA type probably has very narrow genetic variability. A total of 40 PCNA cultivars, including bud sports, which may cover almost all PCNA type cultivars currently existing in Japan, are now preserved at the National Institute of the Fruit Tree Science (NIFTS) in Akitsu, Hiroshima [9]. There are many astringent and PVNA local cultivars throughout Japan. The current conservation in

In Korea, 233 local cultivars were collected at the branch of Experimental Station at Kim-hae during 1959–1969, and 74 superior cultivars were selected for persimmon cultivation after identifying the name of 188 cultivars among these local cultivars. In Korea, interest in persimmon cultivation is increasing and two experimental stations for persimmon have been established, the one for non-astringent persimmon was established in 1994 and the other for astringent persimmon established in 1995. In addition, a breeding program for obtaining new PCNA cultivars was started in 1995 by crosses among PCNA cultivars that were introduced from Japan. The breeding objectives in Korea are focused on obtaining superior PCNA culti-

In Europe, persimmon is considered a secondary fruit tree species; only few countries, located

Persimmon was introduced in Italy at the end of the ninteenth century. Later in Tuscany, the interest for this new species was increased and the genotypes were collected together with exotic and local varieties of fruit tree species (citrus, peaches and plums among others). As early as 1940, the University of Florence collected 11 accessions from the USA and France, or as local varieties and characterized them. The persimmon collection of Florence consisted of 52 cultivars and was totally destroyed by winter frost in 1985. Then a new germplasm orchard was established by introducing new accessions from Japan. A French germplasm was recorded at the beginning of the twentieth century. The Spanish collection was created in 1993 with material from Italy (54.2% of accessions) and from Spanish institutions and nurseries (45.8%) [38].

vars with good eating qualities, large fruit and early ripening characteristics [9].

in the Mediterranean area, are interested in a large-scale production.

cultivars native to China and some native to Japan and Korea.

32 Breeding and Health Benefits of Fruit and Nut Crops

Japan consists of approx. 600 genotypes [37].

Although the exact date of the introduction of persimmon to Anatolia is unknown, it is clear that it dates back to rather old times [39, 40]. Persimmon was introduced to Turkey from Russia via the Black Sea region. Turkey has main persimmon species (*D. kaki*, *D. lotus* and *D. oleifera*). *D. oleifera* can be seen only in the Mediterranean region of Turkey, while *D. lotus* grows as wild in Northern Anatolia and is used as dried fruits in this region. *D. kaki* and *D. oleifera* have been introduced from other countries at least 200 years ago. During this time, continuous propagation of persimmon by its seeds resulted in genetic diversity in *D. kaki* trees due to the high heterozygosity. Therefore, in the northeastern part of Turkey, persimmon trees differ from another in terms of fruit productivity, yield, shape, size, astringency and plant growth. This diversity in persimmon population in Turkey provided a great opportunity to the breeders for selection programs. As a result, the breeders were able to identify many promising clones in different parts of Turkey. A germplasm collection in the Black Sea region in Turkey with selected promising genotypes has been established.

First studies on persimmon in Turkey were started to introduce the foreign cultivars by the Ministry of Agriculture in 1967. Then, some selection studies were done in different parts of Turkey. After 1989, the total number of the known cultivars and types reached up to 74. Most of these varieties were introduced from Italy and some of them were from Israel, Japan, France and Pakistan especially after the attempts made by the Cukurova University, in Eastern Mediterranean, Turkey.

Selection of different genotypes was started by the Department of Horticulture of Cukurova University, by Department of Horticulture of Ondokuzmayıs University in Black Sea and by Citrus Research Institute of Antalya belonging to the Ministry of Agriculture in Western Mediterranean regions. Recently, the wider selections have been carried out especially on Black Sea coast by Citrus Research Institute of Antalya for 1 year and they presently have 43 promising candidate clones. Most even totality of the selections is astringent type. It seems to be rather difficult to find non-astringent types in Turkey. Yilmaz et al. [41] established a characterization study on persimmon genetic resources collected from Turkey. These germplasms were preserved with commercial cultivars in an *ex situ* germplasm preservation orchard located at the Cukurova University, Turkey. Persimmon genotypes were characterized based on their morphological traits. The collection comprising traditional genotypes, local accessions and also global varieties were collected from five different provinces of the Mediterranean region of Turkey where persimmon is widely produced. A total of 48 persimmon genotypes and cultivars were morphologically characterized, using 59 morphological and agronomic traits (**Table 5**).


**No Cultivar and selections Scientific Name Origin Type of astringency**

Genetic Diversity and Breeding of Persimmon http://dx.doi.org/10.5772/intechopen.74977 **Figure 3.** Dendrogram of persimmon accessions collected from Turkey obtained from cluster analysis of 59 agro-

 Jiro C – 24,276 *Diospyros kaki* L. Italy PCNA Kawabata O'Gosho *Diospyros kaki* L. Italy PCNA Giant Fuyu *Diospyros kaki* L. Israel PCNA Tipo Kaki *Diospyros kaki* L. Pakistan PVNA Shogatsu *Diospyros kaki* L. Italy PVNA Giboshi *Diospyros kaki* L. Italy PVNA Thiene *Diospyros kaki* L. Italy PVNA Moro *Diospyros kaki* L. Italy PVNA Brazzale *Diospyros kaki* L. Italy PVNA Kirakaki *Diospyros kaki* L. Italy PVNA Akouman Kaki *Diospyros kaki* L. Italy PVNA Kurokuma *Diospyros kaki* L. Italy PVNA Hyakume *Diospyros kaki* L. Italy PVNA

**Table 5.** Origins of persimmon accessions.

morphological traits using average method.


**Table 5.** Origins of persimmon accessions.

**No Cultivar and selections Scientific Name Origin Type of astringency**

 Shakoku *Diospyros kaki* L. France PCA *Diospyros lotus Diospyros lotus* France PCA *Diospyros virginiana Diospyros virginiana* Israel PCA Seedless Mardan *Diospyros kaki* L. Pakistan PCA Yesil Hurma *Diospyros oleifera* Selection from Adana-ME-Turkey PCA 07 TH 13 *Diospyros kaki* L. Selection from Antalya-ME-Turkey PCA 07 TH 14 *Diospyros kaki* L. Selection from Antalya-ME-Turkey PVA 07 TH 17 *Diospyros kaki* L. Selection from Antalya-ME-Turkey PVA 07 TH 18 *Diospyros kaki* L. Selection from Antalya-ME-Turkey PVA 31 TH 01 *Diospyros kaki* L. Selection from Hatay-ME-Turkey PVA 31 TH 03 *Diospyros kaki* L. Selection from Hatay-ME-Turkey PCA 55 TH 05 *Diospyros kaki* L. Selection from Samsun-BS-Turkey PVA Fatsa-1 *Diospyros kaki* L. Selection from Ordu-BS-Turkey PVA Sarı Yenen *Diospyros kaki* L. Selection from Istanbul-MR-Turkey PCA Cekirdekli *Diospyros kaki* L. Selection from Adana-ME-Turkey PVA Saijo *Diospyros kaki* L. Israel PCA Hachiya *Diospyros kaki* L. Italy PCA Guilbecky *Diospyros kaki* L. Italy PCA BST-29 *Diospyros kaki* L. Italy PCA Fennio *Diospyros kaki* L. Italy PCA Lycopersicon *Diospyros kaki* L. Italy PCA Farmacista Honorati *Diospyros kaki* L. Italy PCA Fujiwara O'Gosho *Diospyros kaki* L. USA PCNA Triumph *Diospyros kaki* L. Israel PCA Vainiglia *Diospyros kaki* L. Pakistan PVNA Aman Kaki-1 *Diospyros kaki* L. Pakistan PVNA Sirin Hurma *Diospyros kaki* L. Iran PVA Nishimura wase *Diospyros kaki* L. Italy PVA Mikatani O'Gosho *Diospyros kaki* L. Italy PVNA Mandarino *Diospyros kaki* L. Italy PVNA Bruniquel *Diospyros kaki* L. Italy PVNA Aman Kaki-2 *Diospyros kaki* L. Italy PVNA Koshu Hyakume *Diospyros kaki* L. Japan PVA Mizushima O'Gosho *Diospyros kaki* L. Italy PVNA Chienting *Diospyros kaki* L. USA PVA

Breeding and Health Benefits of Fruit and Nut Crops

**Figure 3.** Dendrogram of persimmon accessions collected from Turkey obtained from cluster analysis of 59 agromorphological traits using average method.

From them, 9 traits were related with plant growth, 5 with leaves, 7 with flowers, 32 with fruits and 6 with seeds. As a result of the morphological characterization, persimmon varieties and types were classified by PCA, PVA, PCNA and PVNA. Besides, data obtained by characterization were subjected to similarity coefficient, principal components and cluster analyses to study phenotypic relationships among these genotypes. As a result of their study, the 12 factor scores represented 74.75% of the total multivariate variation, and cluster analysis indicated that the similarity index of the population consisting of the investigated genotypes ranged between 81.09 and 17.32% (**Figure 3**).

Main objective of persimmon breeding has been to produce commercially attractive cultivars of the PCNA type which can be eaten without any postharvest treatment [46]. Therefore, PCNA fruit are the most desirable for fresh consumption because it is not necessary to apply any postharvest treatment in order to remove the astringency. Hence, the breeding of new PCNA cultivars is the most popular objective in the entire persimmon growing countries.

Genetic Diversity and Breeding of Persimmon http://dx.doi.org/10.5772/intechopen.74977 37

Although persimmon is produced in Brazil, Israel, Italy, Spain, Azerbaijan, Uzbekistan, New Zealand and Australia, new persimmon cultivars developed by cross breeding have been released only in Japan and Korea [21] and also in these countries persimmon cultivars have been selected over time for commercial production. Hybridization method can also be used in persimmon breeding. In Japan, hybridization method has been used for fruit ripening time,

Persimmon breeding is complex, and results are not always as expected, especially when working on the PCNA group [42]. Researchers found that fruit ripening time is under additive and quantitative control. The tendencies of persimmon fruit are quantitatively inherited traits, the non-cracking cultivars are homozygous, whereas cultivars with cracking are heterozygous. Also, it has been claimed that fruit weight is a quantitative characteristic with high broad-sense heritability [21]. Many crosses performed using large and small fruit size parents indicated that small fruit alleles were dominant to large size alleles [48]. Complete loss of astringency is important for commercial persimmon production. Generally, a little astringency remains in PCNA fruit at maturity in cooler regions, so that they are commercially produced in warm regions. Incomplete loss of astringency results from not only environmental factors but also genetic factors [21]. According to the criteria established for persimmon cultivars, persimmon can be categorized into two major groups, PCNA type consisting of two subcategories, Chinese PCNA (CPCNA) and Japanese PCNA (J-PCNA). The second group is non-PCNA type consisting of three subcategories: PCA, PVNA and PVA [49]. Japanese PCNA cultivars are based on a recessive character and their genetic resources are very few. Repeated crossings within the narrow gene pool cause inbreeding depression, which hinders tree vigor, fruit yield and size. Therefore, studies have been ongoing to obtain new cultivars through the backcross (PCNA × non-PCNA) × PCNA since 1990. In 2007, 'Taiten' and 'Taigetsu', which are PVA cultivars, were derived from the cross of 'Kurokuma' (a local PVNA cultivar in Japan) × PCNA cultivar 'Taishu'. Parthenocarpy in 'Taigetsu' is high [19]. The trait of natural astringency loss is dominant and controlled by the single locus *CPCNA* in Chinese PCNA persimmon [50]. At the end of the hybridization studies in Japan, PCNA cultivars 'Shinshuu', 'Soshu', 'Kanshu' and 'Kishu' were released as early ripening cultivars while 'Suruga', 'Youhou', 'Taishuu' and 'Yubeni' were released as medium to late ripening cultivars [21]. Other persimmon hybrids are 'Fuyuhana' and 'Ito'. 'Fuyuhana' were developed from a 'Fuyu' × 'Hanogosho' cross as an alternative to 'Fuyu' and 'Jiro'. Ito is another

In persimmon, different genus can pollinate each other. Native American persimmon (*D. virgininiana*) and Japanese persimmon (*D. kaki*) hybridization would set a goal of stabilizing and improving the variable flavors and cold hardiness of the native American persimmon. *D. kaki* and *D. virginiana* are apparently cross-incompatible; however a hybrid 'Rossiyanka' has been developed through embryo culture technique [51]. Rossiyanka is cold hardy, nearly seedless and it is smooth textured with Asian persimmon flavor [52]. Nikita's gift hybrid persimmon is unique hybrid of Asian and American persimmon; the fruit is sweet and flavorful [53].

crack-resistance and large fruit size in persimmon.

hybrid obtained by crossing 'Fuyu' × 'Oku-Ogosho' [51].

#### **7. Persimmon breeding and genetic improvement**

Breeding aims for persimmon emphasized on enhanced fruit quality such as fruit weight, shape, color, soluble solids content, fruit cracking, fruit ripening time, high productivity, long shelf life, parthenocarpy and sex expression. Selection breeding is the most common breeding technique in persimmon due to the fact that persimmon breeding is mainly hindered by its high ploidy level and by its complex sex expression [42]. Because somatic and bud sport mutations affect the fruit traits, new lines of persimmon are frequently improved by bud mutations [43, 44]. Also, there are many seed propagated populations in persimmon growing countries of the world especially in native Asian countries. Many established cultivars are chance seedlings selected by growers or researchers in Japan, China and Korea. There are also many selections from introduced genotypes or seedling populations in the USA, Israel, India, Australia, New Zealand, Taiwan, Malaysia and some other countries. 'Kaki Tipo' in Italy, 'Lama Forte' in Brazil, 'Triumph' in Israel and 'Rojo Brillante' in Spain are cultivars that developed from bud sports [45]. 'Fuyu', 'Hachiya', 'Hiratatenashi', 'Izu', 'Jiro' and 'Saijo' cultivars were selected from shoot of bud sports in Japan and these selected cultivars are extensively growing all over the world. Early ripening bud sports of 'Fuyu' (Matsumotowase-Fuyu) and 'Jiro' (Maekawa-Jiro) were also found in a farmer's orchard. Early ripening Matsumotowase-Fuyu showed fruit cracking tendency as 'Fuyu'. Recently, a small fruit mutant, 'Totsutanenashi' (TTN), was discovered in Japan as a bud sport mutant of the leading cultivar Hiratanenashi (HTN) [44]. Following cultivars were also obtained by bud sports in Japan: 'Uenishiwase' (PCNA) and 'Kyi-joh' (PCNA) are bud mutation of 'Matsumotowase-Fuyu'; 'Sunami' (PCNA) and 'Tanbawase-Fuyu' (PCNA) are bud mutation of 'Fuyu'; 'Aisyuhou' (PCNA) is bud mutation of 'Maekawa-Jiro'; 'Tonewase' (PVA), 'Ohtanenashi' (PVA) and 'Kohshimaru' (PVA) are bud mutation of 'Hiratanenashi' [42]. 'Nantongxiaofangshi' variety having dwarfness character is a persimmon that has been found in Nantong. 'Nantong small persimmon' (*D. kaki* Linn. cv. Nantongxiaofangshi) is a rare and dwarf variety of persimmon found in 1982 in Jiangsu Province Nantong City Fruit resource survey.

The height of the adult tree is only about 2 m, which is approximately equal to 60% that of the standard type growing under the same conditions [46]. 'Hasshu' persimmon (*D. kaki* Thunb.) is a dwarf cultivar originated by a bud sport from the leading persimmon cultivar 'Hiratanenashi' in Hiroshima prefecture, Japan in 2005. Its somatic polyploidy (2n = 120 = 8x) was confirmed by flow cytometric analysis and chromosome observation. Although nonaploid 'Hiratanenashi' and some of its bud sports are known to be seedless, 'Hasshu' produces regular seeds with the ability to germinate [16].

Main objective of persimmon breeding has been to produce commercially attractive cultivars of the PCNA type which can be eaten without any postharvest treatment [46]. Therefore, PCNA fruit are the most desirable for fresh consumption because it is not necessary to apply any postharvest treatment in order to remove the astringency. Hence, the breeding of new PCNA cultivars is the most popular objective in the entire persimmon growing countries.

From them, 9 traits were related with plant growth, 5 with leaves, 7 with flowers, 32 with fruits and 6 with seeds. As a result of the morphological characterization, persimmon varieties and types were classified by PCA, PVA, PCNA and PVNA. Besides, data obtained by characterization were subjected to similarity coefficient, principal components and cluster analyses to study phenotypic relationships among these genotypes. As a result of their study, the 12 factor scores represented 74.75% of the total multivariate variation, and cluster analysis indicated that the similarity index of the population consisting of the investigated genotypes

Breeding aims for persimmon emphasized on enhanced fruit quality such as fruit weight, shape, color, soluble solids content, fruit cracking, fruit ripening time, high productivity, long shelf life, parthenocarpy and sex expression. Selection breeding is the most common breeding technique in persimmon due to the fact that persimmon breeding is mainly hindered by its high ploidy level and by its complex sex expression [42]. Because somatic and bud sport mutations affect the fruit traits, new lines of persimmon are frequently improved by bud mutations [43, 44]. Also, there are many seed propagated populations in persimmon growing countries of the world especially in native Asian countries. Many established cultivars are chance seedlings selected by growers or researchers in Japan, China and Korea. There are also many selections from introduced genotypes or seedling populations in the USA, Israel, India, Australia, New Zealand, Taiwan, Malaysia and some other countries. 'Kaki Tipo' in Italy, 'Lama Forte' in Brazil, 'Triumph' in Israel and 'Rojo Brillante' in Spain are cultivars that developed from bud sports [45]. 'Fuyu', 'Hachiya', 'Hiratatenashi', 'Izu', 'Jiro' and 'Saijo' cultivars were selected from shoot of bud sports in Japan and these selected cultivars are extensively growing all over the world. Early ripening bud sports of 'Fuyu' (Matsumotowase-Fuyu) and 'Jiro' (Maekawa-Jiro) were also found in a farmer's orchard. Early ripening Matsumotowase-Fuyu showed fruit cracking tendency as 'Fuyu'. Recently, a small fruit mutant, 'Totsutanenashi' (TTN), was discovered in Japan as a bud sport mutant of the leading cultivar Hiratanenashi (HTN) [44]. Following cultivars were also obtained by bud sports in Japan: 'Uenishiwase' (PCNA) and 'Kyi-joh' (PCNA) are bud mutation of 'Matsumotowase-Fuyu'; 'Sunami' (PCNA) and 'Tanbawase-Fuyu' (PCNA) are bud mutation of 'Fuyu'; 'Aisyuhou' (PCNA) is bud mutation of 'Maekawa-Jiro'; 'Tonewase' (PVA), 'Ohtanenashi' (PVA) and 'Kohshimaru' (PVA) are bud mutation of 'Hiratanenashi' [42]. 'Nantongxiaofangshi' variety having dwarfness character is a persimmon that has been found in Nantong. 'Nantong small persimmon' (*D. kaki* Linn. cv. Nantongxiaofangshi) is a rare and dwarf variety of persimmon found in 1982 in Jiangsu Province Nantong City Fruit resource survey.

The height of the adult tree is only about 2 m, which is approximately equal to 60% that of the standard type growing under the same conditions [46]. 'Hasshu' persimmon (*D. kaki* Thunb.) is a dwarf cultivar originated by a bud sport from the leading persimmon cultivar 'Hiratanenashi' in Hiroshima prefecture, Japan in 2005. Its somatic polyploidy (2n = 120 = 8x) was confirmed by flow cytometric analysis and chromosome observation. Although nonaploid 'Hiratanenashi' and some of its bud sports are known to be seedless, 'Hasshu' pro-

duces regular seeds with the ability to germinate [16].

ranged between 81.09 and 17.32% (**Figure 3**).

36 Breeding and Health Benefits of Fruit and Nut Crops

**7. Persimmon breeding and genetic improvement**

Although persimmon is produced in Brazil, Israel, Italy, Spain, Azerbaijan, Uzbekistan, New Zealand and Australia, new persimmon cultivars developed by cross breeding have been released only in Japan and Korea [21] and also in these countries persimmon cultivars have been selected over time for commercial production. Hybridization method can also be used in persimmon breeding. In Japan, hybridization method has been used for fruit ripening time, crack-resistance and large fruit size in persimmon.

Persimmon breeding is complex, and results are not always as expected, especially when working on the PCNA group [42]. Researchers found that fruit ripening time is under additive and quantitative control. The tendencies of persimmon fruit are quantitatively inherited traits, the non-cracking cultivars are homozygous, whereas cultivars with cracking are heterozygous. Also, it has been claimed that fruit weight is a quantitative characteristic with high broad-sense heritability [21]. Many crosses performed using large and small fruit size parents indicated that small fruit alleles were dominant to large size alleles [48]. Complete loss of astringency is important for commercial persimmon production. Generally, a little astringency remains in PCNA fruit at maturity in cooler regions, so that they are commercially produced in warm regions. Incomplete loss of astringency results from not only environmental factors but also genetic factors [21]. According to the criteria established for persimmon cultivars, persimmon can be categorized into two major groups, PCNA type consisting of two subcategories, Chinese PCNA (CPCNA) and Japanese PCNA (J-PCNA). The second group is non-PCNA type consisting of three subcategories: PCA, PVNA and PVA [49]. Japanese PCNA cultivars are based on a recessive character and their genetic resources are very few. Repeated crossings within the narrow gene pool cause inbreeding depression, which hinders tree vigor, fruit yield and size. Therefore, studies have been ongoing to obtain new cultivars through the backcross (PCNA × non-PCNA) × PCNA since 1990. In 2007, 'Taiten' and 'Taigetsu', which are PVA cultivars, were derived from the cross of 'Kurokuma' (a local PVNA cultivar in Japan) × PCNA cultivar 'Taishu'. Parthenocarpy in 'Taigetsu' is high [19]. The trait of natural astringency loss is dominant and controlled by the single locus *CPCNA* in Chinese PCNA persimmon [50]. At the end of the hybridization studies in Japan, PCNA cultivars 'Shinshuu', 'Soshu', 'Kanshu' and 'Kishu' were released as early ripening cultivars while 'Suruga', 'Youhou', 'Taishuu' and 'Yubeni' were released as medium to late ripening cultivars [21]. Other persimmon hybrids are 'Fuyuhana' and 'Ito'. 'Fuyuhana' were developed from a 'Fuyu' × 'Hanogosho' cross as an alternative to 'Fuyu' and 'Jiro'. Ito is another hybrid obtained by crossing 'Fuyu' × 'Oku-Ogosho' [51].

In persimmon, different genus can pollinate each other. Native American persimmon (*D. virgininiana*) and Japanese persimmon (*D. kaki*) hybridization would set a goal of stabilizing and improving the variable flavors and cold hardiness of the native American persimmon. *D. kaki* and *D. virginiana* are apparently cross-incompatible; however a hybrid 'Rossiyanka' has been developed through embryo culture technique [51]. Rossiyanka is cold hardy, nearly seedless and it is smooth textured with Asian persimmon flavor [52]. Nikita's gift hybrid persimmon is unique hybrid of Asian and American persimmon; the fruit is sweet and flavorful [53].

In persimmon breeding programs, mutation breeding technique has also used. The main objectives of persimmon mutation breeding were focused on obtaining new cultivars with the positive agronomic features but with more diversity in ripening date, astringency and fruit characteristics from the PCNA types. However, obtaining PCNA type varieties is difficult due to the dominant inheritance of astringency, the limited number of cultivars which bear male flowers and the hexaploid inheritance of basic persimmon cultivars. Therefore, the PCNA type cultivars have low genetic diversity and crossing among these generally result in negative effects of inbreeding. Mutation breeding has been used as an alternative method for generating diversity in persimmon [54]. Some researchers studied to determine which gamma ray doses can be used in persimmon. Ray [51] claimed that 5–10 kR gamma doses obtained widest range of viability on cuttings, seeds and pollen of persimmon. In Spain, shoot buds of the persimmon 'Rojo Brillante' were subjected to various doses of gamma rays, 15 and 20 gray from a 60Co source. In this study, Naval et al. [55] found that the most favorable gamma irradiation dose combining survival and mutation induction was 20 gray. Two new varieties with similar fruit quality to 'Rojo Brillante', that allow to enlarge the persimmon harvest season in Spain, were selected [56].

that the placement of several Japanese cultivars within the European cultivar group suggests that European cultivars were developed from Japanese germplasm relatively recent and differences among cultivars are much greater than differences among cultivar groups regarding AFLP markers. In addition, Guo and Luo [60] indicated that SSR markers are a valuable tool for

Genetic Diversity and Breeding of Persimmon http://dx.doi.org/10.5772/intechopen.74977 39

The main *in vitro* tissue culture techniques developed for persimmon deal with direct regeneration (from dormant buds and root tips) and indirect regeneration through callus from dormant buds, apexes and leaves. Kochanová et al. [61] indicated that in the genus *Diospyros* L., biotechnological researches focused on quality improvement and preservation of the cultivars that has been economically cultivated. The authors also remarked that the genetic variability had been lost as only those limited cultivars that are popular among growers are grown. In recent years, studies were conducted on *in vitro* micro-propagation of persimmon [62], especially on Jiro [63] and Rojo Brillante [64]. Choi et al. [65] recorded an efficient and simple plant regeneration via organogenesis from leaf segment cultures of persimmon (*D. kaki* Thunb.). The authors indicated that the frequencies of adventitious shoot regeneration by 'Nishimurawase' and 'Fuyu' reached up to 100% and the regenerated shoots rooted successfully with over 80% efficiency. Yokoyama et al. [66] suggested that the meristematic nodule is a promising material for propagation and long-term conservation of 'Fuyu' variety. Naval et al. [67] recorded a protocol for plant regeneration of *D. kaki* Thunb. cv. 'Rojo Brillante' via organogenesis from leaf explants by using combined phytohormones and dosages. In addition, Naval et al. [67] studied somaclonal variation of 'Rojo Brillante' as a breeding tool by using various combinations of cytokinin (Z or BA) with different auxins (IAA or NAA). Furthermore, Palla et al. [68] studied *in vitro* culture and rooting of *D. virginiana* L. from nodal root explants by using several phytohormones and culture media. The authors indicated that the presence of auxins was not essential but slightly accelerated the organogenic callus formation and organogenesis. Cryopreservation is recognized as having the distinctive advantage of allowing long-term conservation with minimum space and maintenance [69]. Matsumoto et al. [70] studied cryopreservation of persimmon (*D. kaki* Thunb.) by vitrification of dormant shoot tips. The authors recorded that using dormant shoot tips was

promising as a routine method for the cryopreservation of *Diospyros* germplasm.

After the great progress in *in vitro* regeneration of plants from protoplasts, several researches focused on plant somatic hybridization which allows combining protoplasts from different cultivars, species or genera for variety improvement [71]. Tao et al. [72] reported plant regeneration from callus protoplast of *D. kaki*. They used callus as the protoplast source derived from leaf primordia excised from dormant winter buds of adult Japanese persimmon (*D. kaki* L. cv. Jiro) for plant regeneration. Tamura et al. [73] studied protoplast culture and plant regeneration of *D. kaki* L. and reported that plantlets could be obtained from the protoplast-derived calli. Tamura et al. [74] indicated that somatic hybrids of Japanese persimmon (*D. kaki* L.) were obtained by electrofusion of protoplasts. Callus protoplasts of Jiro and Suruga were fused electrically and cultured in modified KM8p medium using agarose-bead culture. The authors recorded that the fused products had the dodecaploid chromosome number of around 2n = 180, which is twice the number of parental plants (2n = 90x = 15). In addition, Tamura et al. [75] recorded interspecific somatic hybrids between *D. glandulosa* (2n = 2x = 30) and *D. kaki* cv. Jiro (2n = 6x = 90) produced by electrofusion of protoplasts. Colchicine treatment of actively dividing cells can induce chromosome doubling and has been used to make plants with

the estimation of genetic diversity and divergence in *Diospyros*.

#### **8. Biotechnology and genomics**

Biotechnology refers to the use of living organisms or their components to provide useful products in its broadest sense. Using biotechnology in plant breeding has become the most attractive method due to increasing knowledge in plant biotechnology and genomics. Improvements in the field of genomics have resulted in the development of huge quantities of useful new knowledge that greatly assists scientific plant breeding. Also, improvements in biotechnological techniques like plant tissue culture provided new methods for rapid production of high-quality, disease-free and true to type planting material.

In persimmon, biotechnological advances and molecular biology have been used for the classification of *Diospyros* species, *in vitro* propagation, regeneration from callus, root, protoplast and endosperm, ploidy manipulations, agrobacterium-mediated genetic transformation and markerassisted selection. Molecular markers have been widely used for investigating the genetic relationships among persimmon genotypes. Akbulut et al. [57] compared persimmon genotypes by using random amplified polymorphic DNA (RAPD) and fatty acid methyl esters (FAME) data. The results showed that RAPD analyses could differentiate the relationship of persimmon (*D. kaki* Thunb.) genotypes used in their study. The authors suggested that more cultivars were needed as plant materials in terms of determining the degree of relationships of RAPD and FAME data which could help delimiting taxonomic classes within persimmon. Raddová et al. [58] indicated that RAPD and inter-primer binding site (i-PBS) were reliable enough to detect differences between the genetically close cultivars of persimmon. In addition, Badenes et al. [45] studied the genetic diversity of introduced and local Spanish persimmon cultivars as revealed by RAPD markers. The authors suggested that a correct identification of germplasm material from persimmon collections should be the first step in projects related to breeding or management of cultivar aimed at improving the crop. They also indicated that RAPD technology is adequate for fingerprinting persimmon. Yonemori et al. [59] studied the relationship between the European persimmon (*D. kaki* Thunb.) cultivars and Asian cultivars using AFLPs. The authors indicated that the placement of several Japanese cultivars within the European cultivar group suggests that European cultivars were developed from Japanese germplasm relatively recent and differences among cultivars are much greater than differences among cultivar groups regarding AFLP markers. In addition, Guo and Luo [60] indicated that SSR markers are a valuable tool for the estimation of genetic diversity and divergence in *Diospyros*.

In persimmon breeding programs, mutation breeding technique has also used. The main objectives of persimmon mutation breeding were focused on obtaining new cultivars with the positive agronomic features but with more diversity in ripening date, astringency and fruit characteristics from the PCNA types. However, obtaining PCNA type varieties is difficult due to the dominant inheritance of astringency, the limited number of cultivars which bear male flowers and the hexaploid inheritance of basic persimmon cultivars. Therefore, the PCNA type cultivars have low genetic diversity and crossing among these generally result in negative effects of inbreeding. Mutation breeding has been used as an alternative method for generating diversity in persimmon [54]. Some researchers studied to determine which gamma ray doses can be used in persimmon. Ray [51] claimed that 5–10 kR gamma doses obtained widest range of viability on cuttings, seeds and pollen of persimmon. In Spain, shoot buds of the persimmon 'Rojo Brillante' were subjected to various doses of gamma rays, 15 and 20 gray from a 60Co source. In this study, Naval et al. [55] found that the most favorable gamma irradiation dose combining survival and mutation induction was 20 gray. Two new varieties with similar fruit quality to 'Rojo Brillante',

that allow to enlarge the persimmon harvest season in Spain, were selected [56].

tion of high-quality, disease-free and true to type planting material.

Biotechnology refers to the use of living organisms or their components to provide useful products in its broadest sense. Using biotechnology in plant breeding has become the most attractive method due to increasing knowledge in plant biotechnology and genomics. Improvements in the field of genomics have resulted in the development of huge quantities of useful new knowledge that greatly assists scientific plant breeding. Also, improvements in biotechnological techniques like plant tissue culture provided new methods for rapid produc-

In persimmon, biotechnological advances and molecular biology have been used for the classification of *Diospyros* species, *in vitro* propagation, regeneration from callus, root, protoplast and endosperm, ploidy manipulations, agrobacterium-mediated genetic transformation and markerassisted selection. Molecular markers have been widely used for investigating the genetic relationships among persimmon genotypes. Akbulut et al. [57] compared persimmon genotypes by using random amplified polymorphic DNA (RAPD) and fatty acid methyl esters (FAME) data. The results showed that RAPD analyses could differentiate the relationship of persimmon (*D. kaki* Thunb.) genotypes used in their study. The authors suggested that more cultivars were needed as plant materials in terms of determining the degree of relationships of RAPD and FAME data which could help delimiting taxonomic classes within persimmon. Raddová et al. [58] indicated that RAPD and inter-primer binding site (i-PBS) were reliable enough to detect differences between the genetically close cultivars of persimmon. In addition, Badenes et al. [45] studied the genetic diversity of introduced and local Spanish persimmon cultivars as revealed by RAPD markers. The authors suggested that a correct identification of germplasm material from persimmon collections should be the first step in projects related to breeding or management of cultivar aimed at improving the crop. They also indicated that RAPD technology is adequate for fingerprinting persimmon. Yonemori et al. [59] studied the relationship between the European persimmon (*D. kaki* Thunb.) cultivars and Asian cultivars using AFLPs. The authors indicated

**8. Biotechnology and genomics**

38 Breeding and Health Benefits of Fruit and Nut Crops

The main *in vitro* tissue culture techniques developed for persimmon deal with direct regeneration (from dormant buds and root tips) and indirect regeneration through callus from dormant buds, apexes and leaves. Kochanová et al. [61] indicated that in the genus *Diospyros* L., biotechnological researches focused on quality improvement and preservation of the cultivars that has been economically cultivated. The authors also remarked that the genetic variability had been lost as only those limited cultivars that are popular among growers are grown. In recent years, studies were conducted on *in vitro* micro-propagation of persimmon [62], especially on Jiro [63] and Rojo Brillante [64]. Choi et al. [65] recorded an efficient and simple plant regeneration via organogenesis from leaf segment cultures of persimmon (*D. kaki* Thunb.). The authors indicated that the frequencies of adventitious shoot regeneration by 'Nishimurawase' and 'Fuyu' reached up to 100% and the regenerated shoots rooted successfully with over 80% efficiency. Yokoyama et al. [66] suggested that the meristematic nodule is a promising material for propagation and long-term conservation of 'Fuyu' variety. Naval et al. [67] recorded a protocol for plant regeneration of *D. kaki* Thunb. cv. 'Rojo Brillante' via organogenesis from leaf explants by using combined phytohormones and dosages. In addition, Naval et al. [67] studied somaclonal variation of 'Rojo Brillante' as a breeding tool by using various combinations of cytokinin (Z or BA) with different auxins (IAA or NAA). Furthermore, Palla et al. [68] studied *in vitro* culture and rooting of *D. virginiana* L. from nodal root explants by using several phytohormones and culture media. The authors indicated that the presence of auxins was not essential but slightly accelerated the organogenic callus formation and organogenesis. Cryopreservation is recognized as having the distinctive advantage of allowing long-term conservation with minimum space and maintenance [69]. Matsumoto et al. [70] studied cryopreservation of persimmon (*D. kaki* Thunb.) by vitrification of dormant shoot tips. The authors recorded that using dormant shoot tips was promising as a routine method for the cryopreservation of *Diospyros* germplasm.

After the great progress in *in vitro* regeneration of plants from protoplasts, several researches focused on plant somatic hybridization which allows combining protoplasts from different cultivars, species or genera for variety improvement [71]. Tao et al. [72] reported plant regeneration from callus protoplast of *D. kaki*. They used callus as the protoplast source derived from leaf primordia excised from dormant winter buds of adult Japanese persimmon (*D. kaki* L. cv. Jiro) for plant regeneration. Tamura et al. [73] studied protoplast culture and plant regeneration of *D. kaki* L. and reported that plantlets could be obtained from the protoplast-derived calli. Tamura et al. [74] indicated that somatic hybrids of Japanese persimmon (*D. kaki* L.) were obtained by electrofusion of protoplasts. Callus protoplasts of Jiro and Suruga were fused electrically and cultured in modified KM8p medium using agarose-bead culture. The authors recorded that the fused products had the dodecaploid chromosome number of around 2n = 180, which is twice the number of parental plants (2n = 90x = 15). In addition, Tamura et al. [75] recorded interspecific somatic hybrids between *D. glandulosa* (2n = 2x = 30) and *D. kaki* cv. Jiro (2n = 6x = 90) produced by electrofusion of protoplasts. Colchicine treatment of actively dividing cells can induce chromosome doubling and has been used to make plants with doubled chromosome number. Colchicine treatment to a protoplast at the very beginning of its division could be one method to overcome the problem because plants can be regenerated from a single cell with doubled chromosome number. Tamura et al. [74] reported production of dodecaploid plants of Japanese persimmon by colchicine treatment of protoplasts.

Selection breeding is the most common breeding technique in persimmon because persimmon breeding is mainly hindered by its high ploidy and by its complex sex expression. Fuyu, Hachiya, Hiratatenashi, Izu, Jiro and Saijo cultivars which are extensively growing all over the world were selected from shoots of bud sports in Japan. We should continue the screening of plants coming from spontaneous mutations. Hybridization method can also be used in persimmon breeding. Hybridization studies among *D. kaki* in Japan have led to the release of a lot of PCNA cultivars which ripen at different times. There is also a unique hybrid of Asian and American persimmon. In Spain, studies on induced mutation have also led to the devel-

Genetic Diversity and Breeding of Persimmon http://dx.doi.org/10.5772/intechopen.74977 41

There are a number of collections including many accessions in institutions of the various persimmon growing countries. The morphological and molecular characterization of all the persimmon accessions needs to be achieved. The information developed from this will be highly beneficial for screening against biotic and abiotic stress factors. Genomics and transcriptomic resources need to be developed for persimmon. It will also lead to the development of new

opment of new cultivars.

**Author details**

**References**

and improved cultivars of persimmon.

10.1007/978-3-540-34533-6

\*Address all correspondence to: tyesil@cu.edu.tr

www.tuik.gov.tr [Accessed: 2017-09-23]

The Netherlands: Wageningen; 1975

1987. pp. 411-416

ture. 2003

Turgut Yesiloglu\*, Berken Cimen, Meral Incesu and Bilge Yilmaz

Faculty of Agriculture, Department of Horticulture, Cukurova University, Turkey

[1] Kanzaki S, Persimmon YK. Genome mapping and molecular breeding in plants. In: Kole C, editor. Fruits and Nuts. Berlin Heidelberg: Springer; 2007. pp. 353-358. DOI:

[2] FAO Stat. [Internet]. 2017. Available from: http://www.faostat.com [Accessed: 2017-11-12] [3] TUİK. Turkish Statistic Council records (Ankara: TUIK) [Internet]. 2017. Avaible from:

[4] Llácer G, Martínez-Calvo J, Naval M, Badenes ML. From germplasm to fruit export: The case of 'Rojo Brillante' persimmon. Advances in Horticultural Science. 2008;**22**(4):281-285 [5] Zeven AC, Zhukovsky PM. Dictionary of Cultivated Plants and their Centres of Diversity.

[6] Morton J. Japanese Persimmon. In: Fruits of warm climates. Miami, FL: Julia F. Morton;

[7] Ullio L. Agfacts: Persimmon Growing in New South Wales. New South Wales Agricul-

Improved genomic research and resources, in recent years, have resulted in the development of screening tools via marker-assisted selection (MAS). Using MAS has led to more efficient selections and has increased the efficiency in persimmon breeding programs hastening the release of new cultivar. In order to obtain PCNA offspring in breeding programs, the parental materials considered for choosing the cross combinations have to be PCNA type regarding the inheritance of astringency. However, repeated crosses among PCNA cultivars/selections has led to inbreeding depression for tree vigor, productivity and fruit weight [36]. In these situations, marker-assisted selection should be developed for selecting PCNA offspring efficiently. Recently, Kanzaki et al. [47] have developed molecular markers associated with the trait of natural astringency loss in persimmon fruit and the markers are practically useful in persimmon breeding programs. In addition, Mitani et al. [76] studied if the SCAR markers could reliably distinguish PCNA and non-PCNA genotypes in a large number of offspring derived from backcross between 'Taigetsu' and PCNA 'Kanshu'. The authors indicated that PCNA offspring can be selected by two PCR primers in the progeny derived from 'Taigetsu' × 'Kanshu'. Yonemori et al. [77] reported molecular marker for selecting PCNA type persimmon progenies at the juvenile stage. Yonemori et al. [77] constructed a reliable PCR marker for selecting PCNA type offspring among breeding population of persimmon. In addition Kanzaki et al. [47] and Mitani et al. [76] reported that SCAR markers can practically be used in application of marker-assisted selection in persimmon breeding.

Genetic transformation is also an alternative technique for persimmon genetic improvement. Transgenic persimmon cultivars thus produced have potential for commercial success and grower acceptance because the unique genetic constitution of the cultivars has not been disturbed. Tao et al. [78] reported genetic transformation of persimmon by A*agrobacterium rhizogenes*. Phenotypic alterations such as dwarfness and decrease in rooting ability were observed in the transformants. In addition, Gao et al. [79] transformed 'Jiro' persimmon with Arabidopsis FT gene (*AtFT*) and *PmTFL1* gene, a *Prunus mume* ortholog of Arabidopsis *TFL1* gene. The authors indicated that the PmTFL1 transgenic *in vitro* shoots did not show a different appearance compared with non-transformed 'Jiro' shoots, however, the AtFT transgenic shoots indicated a 'bushy' phenotype having the short internodes.

#### **9. Conclusions**

Persimmon can adapt to a wide range of climatic conditions. Production in many countries having subtropical and tropical climates satisfies domestic demand and creates new export opportunities. Increasing the world persimmon production has been very successful since 1995. Recently, the applications of the technique for removing astringency without losing fruit firmness have been significantly promoted to increase the production. It is expected that the production will significantly increase over the next few decades.

Selection breeding is the most common breeding technique in persimmon because persimmon breeding is mainly hindered by its high ploidy and by its complex sex expression. Fuyu, Hachiya, Hiratatenashi, Izu, Jiro and Saijo cultivars which are extensively growing all over the world were selected from shoots of bud sports in Japan. We should continue the screening of plants coming from spontaneous mutations. Hybridization method can also be used in persimmon breeding. Hybridization studies among *D. kaki* in Japan have led to the release of a lot of PCNA cultivars which ripen at different times. There is also a unique hybrid of Asian and American persimmon. In Spain, studies on induced mutation have also led to the development of new cultivars.

There are a number of collections including many accessions in institutions of the various persimmon growing countries. The morphological and molecular characterization of all the persimmon accessions needs to be achieved. The information developed from this will be highly beneficial for screening against biotic and abiotic stress factors. Genomics and transcriptomic resources need to be developed for persimmon. It will also lead to the development of new and improved cultivars of persimmon.

#### **Author details**

doubled chromosome number. Colchicine treatment to a protoplast at the very beginning of its division could be one method to overcome the problem because plants can be regenerated from a single cell with doubled chromosome number. Tamura et al. [74] reported production of

Improved genomic research and resources, in recent years, have resulted in the development of screening tools via marker-assisted selection (MAS). Using MAS has led to more efficient selections and has increased the efficiency in persimmon breeding programs hastening the release of new cultivar. In order to obtain PCNA offspring in breeding programs, the parental materials considered for choosing the cross combinations have to be PCNA type regarding the inheritance of astringency. However, repeated crosses among PCNA cultivars/selections has led to inbreeding depression for tree vigor, productivity and fruit weight [36]. In these situations, marker-assisted selection should be developed for selecting PCNA offspring efficiently. Recently, Kanzaki et al. [47] have developed molecular markers associated with the trait of natural astringency loss in persimmon fruit and the markers are practically useful in persimmon breeding programs. In addition, Mitani et al. [76] studied if the SCAR markers could reliably distinguish PCNA and non-PCNA genotypes in a large number of offspring derived from backcross between 'Taigetsu' and PCNA 'Kanshu'. The authors indicated that PCNA offspring can be selected by two PCR primers in the progeny derived from 'Taigetsu' × 'Kanshu'. Yonemori et al. [77] reported molecular marker for selecting PCNA type persimmon progenies at the juvenile stage. Yonemori et al. [77] constructed a reliable PCR marker for selecting PCNA type offspring among breeding population of persimmon. In addition Kanzaki et al. [47] and Mitani et al. [76] reported that SCAR markers can practically be used

Genetic transformation is also an alternative technique for persimmon genetic improvement. Transgenic persimmon cultivars thus produced have potential for commercial success and grower acceptance because the unique genetic constitution of the cultivars has not been disturbed. Tao et al. [78] reported genetic transformation of persimmon by A*agrobacterium rhizogenes*. Phenotypic alterations such as dwarfness and decrease in rooting ability were observed in the transformants. In addition, Gao et al. [79] transformed 'Jiro' persimmon with Arabidopsis FT gene (*AtFT*) and *PmTFL1* gene, a *Prunus mume* ortholog of Arabidopsis *TFL1* gene. The authors indicated that the PmTFL1 transgenic *in vitro* shoots did not show a different appearance compared with non-transformed 'Jiro' shoots, however, the AtFT transgenic

Persimmon can adapt to a wide range of climatic conditions. Production in many countries having subtropical and tropical climates satisfies domestic demand and creates new export opportunities. Increasing the world persimmon production has been very successful since 1995. Recently, the applications of the technique for removing astringency without losing fruit firmness have been significantly promoted to increase the production. It is expected that

dodecaploid plants of Japanese persimmon by colchicine treatment of protoplasts.

40 Breeding and Health Benefits of Fruit and Nut Crops

in application of marker-assisted selection in persimmon breeding.

shoots indicated a 'bushy' phenotype having the short internodes.

the production will significantly increase over the next few decades.

**9. Conclusions**

Turgut Yesiloglu\*, Berken Cimen, Meral Incesu and Bilge Yilmaz

\*Address all correspondence to: tyesil@cu.edu.tr

Faculty of Agriculture, Department of Horticulture, Cukurova University, Turkey

#### **References**


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[32] Greene SL, Morris JB. The case for multiple-use plant germplasm collections and a strat-

[33] Naval M, Zuriaga E, Pecchioli S, Llácer G, Giordani E, Badenes ML. Analysis of genetic diversity among persimmon cultivars using microsatellite markers. Tree Genetics and

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[37] Yamada M. Persimmon genetic resources and breeding in Japan. Acta Horticulturae.

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[8] Grubov VI. Family Ebenaceae. In: Shishkin BK, Bobrov EG, editors. Flora of the U.S.S.R. Vol. 18. Jerusalem: Israel Program for Scientific Translations; 1967. pp. 349-355

[9] Yonemori K, Sugiura A, Yamada M. Persimmon genetics and breeding. Plant Breeding

[10] Kitagawa H, Glucina P. Persimmon Culture in New Zealand. Wellington: Dept. Science

[11] Darrow GM. Minor temperate fruits. In: Janick J, Moore JN, editors. Advances in Fruit

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**Chapter 3**

**Provisional chapter**

**Peach Breeding Studies in Turkey and the Evaluation of**

Peach (*Prunus persica* [L.] Batsch) is widely cultivated due to its easy adaptability to different ecological conditions, early fruit set and a long period of harvest. Peach cultivation extends along 30–45° North and South parallels of latitude. Around 21,638,953 tons of peaches are produced in an area of 1538,174 ha across the world. Turkey ranks sixth with a production of 637,573 tons/year in 29,092 ha area. Early fruiting habit and correlative characteristics of peaches encouraged fruit breeders to study on this fruit species. The main aims of the study are new cultivar or rootstock breeding, resistance to diseases, late ripening season, fruit quality improvement, fruit shape changes, new tree shapes, and low chilling cultivars. Breeding studies have been carried out at the Department of Horticulture at the University of Cukurova since 1990s. In these, peach and nectarine breeding programs with different aims such as late ripening, quality improvement, Sharka resistance, and low chilling cultivars were studied. In this chapter, some of the results on late ripening peach and nectarine breeding program have also been presented. **Keywords:** molecular markers, embryo rescue, low and high chilling cultivars, Sharka

**Peach Breeding Studies in Turkey and the Evaluation of** 

DOI: 10.5772/intechopen.73440

© 2016 The Author(s). Licensee InTech. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution,

© 2018 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use,

distribution, and reproduction in any medium, provided the original work is properly cited.

and reproduction in any medium, provided the original work is properly cited.

Iwata et al. [1] stated that in genomic analysis technologies, many new advancements promote the efficiency of plant breeding. They also determined that genome-wide association studies (GWAS) and genomic selection (GS) are very helpful, especially in various fruit tree breeding research programs. Breeding of fruit crops is not so easy, as it takes a very long time to get the quality fruits with good size of a tree, to overcome the juvenility period as well as to obtain

**Peach and Nectarine Hybrids**

**Peach and Nectarine Hybrids**

Ayzin Baykam Kuden, Songul Comlekcioglu, Kadir Sarier, Burhanettin Imrak and Ali Kuden

Ayzin Baykam Kuden, Songul Comlekcioglu, Kadir Sarier,

Additional information is available at the end of the chapter

Additional information is available at the end of the chapter

http://dx.doi.org/10.5772/intechopen.73440

resistance, fruit quality characteristics

**Abstract**

**1. Introduction**

Burhanettin Imrak and Ali Kuden


**Provisional chapter**

#### **Peach Breeding Studies in Turkey and the Evaluation of Peach and Nectarine Hybrids Peach and Nectarine Hybrids**

**Peach Breeding Studies in Turkey and the Evaluation of** 

DOI: 10.5772/intechopen.73440

Ayzin Baykam Kuden, Songul Comlekcioglu, Kadir Sarier, Burhanettin Imrak and Ali Kuden Burhanettin Imrak and Ali Kuden Additional information is available at the end of the chapter

Ayzin Baykam Kuden, Songul Comlekcioglu, Kadir Sarier,

Additional information is available at the end of the chapter

http://dx.doi.org/10.5772/intechopen.73440

#### **Abstract**

[67] Naval MM, Llacer G, Badenes ML, Giordani E. Adventitious shoot regeneration from leaf explants of the persimmon (*Diospyros kaki* Thunb.) cv. 'Rojo Brillante'. Acta

[68] Palla KJ, Beasley RR. In vitro culture and rooting of *Diospyros virginiana* L. Hortscience.

[69] Sakai A. Potentially valuable cryogenic procedures for cryopreservation of cultured plant meristems. In: Lazdan MK, Cocking EC, editors. Conservation of Plant Genetic

[70] Matsumoto T, Mochida K,·Itamura H, Sakai A.Cryopreservation of persimmon (*Diospyros kaki* Thunb.) by vitrification of dormant shoot tips. Plant Cell Reports. 2001;**20**:398-402.

[71] Cimen B, Yesiloglu T. 2016. Rootstock Breeding for Abiotic Stress Tolerance in Citrus, Abiotic and Biotic Stress in Plants—Recent Advances and Future Perspectives. In:

[72] Tao R, Tamura M, Yonemori K, Sugiura A. Plant regeneration from callus protoplast of adult Japanese persimmon (*Diospyros kaki* L.). Plant Science. 1991;**79**:119-125

[73] Tamura M, Tao R, Sugiura A. Improved protoplast culture and plant regeneration of Japanese persimmon (*Diospyros kaki* L.). Japanese Journal of Breeding. 1993;**43**:239-245

[74] Tamura M, Tao R, Sugiura A. Regeneration of somatic hybrids from electrofused protoplast of Japanese persimmon (*Diospyros kaki* L.). Plant Science. 1995;**108**:101-107

[75] Tamura M, Tao R, Suguira A. Production of somatic hybrids between *Diospyros gladulosa* and *D. kaki* by protoplast fusion. Plant Cell, Tissue and Organ Culture. 1998;**54**:85-91 [76] Mitani N, Kono A, Yamada M, Sato A, Kobayashi S, Ban Y, Ueno T, Shiraishi M, Kanzaki S, Tsujimoto T. Application of marker-assisted selection in persimmon breeding of PCNA offspring using Scar markers among the population from the cross between non-

[77] Yonemori K, Akagi T, Kanzaki S. Construction of a reliable PCR marker for selecting pollination constant and non-astringent (PCNA) type offspring among breeding population

of persimmon (*Diospyros kaki* Thunb.). Acta Horticulturae. 2009;**839**(839):625-630

[78] Tao R, Handa T, Tamyra M, Sugiura A. Genetic transformation of Japanese persimmon (*Diospyros kaki* L.) by agrobacterium rhizogenes wild type strain A4. Journal of the

[79] Gao M, Takeishi H, Katayama A, Tao R. Genetic transformation of Japanese persimmon with flowering locus T (FT) gene and terminal flower 1 (TfFL1) homologues gene. Acta

PCNA 'Taigetsu' and PCNA 'Kanshu'. Hortscience. 2014;**49**:1132-1135

Horticulturae. 2013;**996**:159-167. DOI: 10.17660/ActaHortic.2013.996.20

Japanese Society for Horticultural Science. 1994;**63**:283-289

Horticulturae. 2009;**833**:183-186

46 Breeding and Health Benefits of Fruit and Nut Crops

DOI 10.1007/s002990100350

Resources In Vitro. USA: Science; 1997. pp. 53-66

Shanker A, editor. InTech. DOI: 10.5772/62047

2013;**48**(6):747-749

Peach (*Prunus persica* [L.] Batsch) is widely cultivated due to its easy adaptability to different ecological conditions, early fruit set and a long period of harvest. Peach cultivation extends along 30–45° North and South parallels of latitude. Around 21,638,953 tons of peaches are produced in an area of 1538,174 ha across the world. Turkey ranks sixth with a production of 637,573 tons/year in 29,092 ha area. Early fruiting habit and correlative characteristics of peaches encouraged fruit breeders to study on this fruit species. The main aims of the study are new cultivar or rootstock breeding, resistance to diseases, late ripening season, fruit quality improvement, fruit shape changes, new tree shapes, and low chilling cultivars. Breeding studies have been carried out at the Department of Horticulture at the University of Cukurova since 1990s. In these, peach and nectarine breeding programs with different aims such as late ripening, quality improvement, Sharka resistance, and low chilling cultivars were studied. In this chapter, some of the results on late ripening peach and nectarine breeding program have also been presented.

**Keywords:** molecular markers, embryo rescue, low and high chilling cultivars, Sharka resistance, fruit quality characteristics

#### **1. Introduction**

Iwata et al. [1] stated that in genomic analysis technologies, many new advancements promote the efficiency of plant breeding. They also determined that genome-wide association studies (GWAS) and genomic selection (GS) are very helpful, especially in various fruit tree breeding research programs. Breeding of fruit crops is not so easy, as it takes a very long time to get the quality fruits with good size of a tree, to overcome the juvenility period as well as to obtain

Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. © 2018 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

© 2016 The Author(s). Licensee InTech. This chapter is distributed under the terms of the Creative Commons

desired fruits according to the aim of the breeding program [1]. Whole-genome sequences have been recently released for apple, peach, strawberry, Japanese apricot, and Chinese and European pear. After these developments, in breeding programs, marker-assisted selection has been used in wide genomics studies. These studies have caused to develop genome scale SNPs and SSR markers and to place reference linkage maps in Rosacea family to allow the identification of evolutionary relationships, which can be found in Genome Database in Rosaceae website. Yamamoto and Terakami [2] reviewed the recent advances in genomic studies and their practical applications for Rosaceae fruit trees, particularly in pear, apple, peach, and cherry.

Various regions of Turkey are suitable to grow peaches under different ecological conditions. The Marmara region can grow high chilling requiring peaches, but the ripening time of the latest peach cultivars is about mid-September. A peach breeding study was carried out by Eroglu [11] in this important peach growing area. In this project, five foreign cultivars (Rio Oso Gem, Fortuna, Monroe, Jungerman and Vivian) and four local types (Bayramiç Tüysüzü, Alyanak Hulu, Sarı Papa and Takunyacı I) were hybridized to obtain fresh market peaches and two foreign and one local type peaches were hybridized for processing peaches. Eroglu et al. [12] stated the fruit quality performances of 121 peach genotypes for fresh market and 35 genotypes for processing. Adana province in Cukurova plain at the Mediterranean region has a subtropical climate. It is located very close to the Mediterranean Sea, thus this condition is suitable to grow only low or midseason chilling requiring peach cultivars. Within 1 h distance to this area, in Taurus Mountains (1100 m elevation) region, high chilling requiring cultivars

Peach Breeding Studies in Turkey and the Evaluation of Peach and Nectarine Hybrids

http://dx.doi.org/10.5772/intechopen.73440

49

**a.** The first peach breeding study in Turkey was conducted by Tanriver and Kuden [13] on 'Ustun', a very late ripening (in mid-October) peach cultivar from the beginning of the 1990s. This study was carried out for [1] breeding of early cultivars for subtropical regions and [2] breeding of late ripening cultivars for cold regions, during 1995–1999. In the breeding experiments, carried out in Adana for early peach cultivars, Springtime, Suncrest, Flavorcrest and Redcap peach cultivars were used. Embryo rescue method was used with the combinations, where Springtime was used as a mother parent. As a result, embryo size was found to be important for the success of embryo rescue, and early cultivars should be used as pollinators. In the breeding studies to obtain late ripening peach cultivars, 'Monroe', 'Rio-Oso-Gem' and 'Ustun' were crossed reciprocally, while 'J.H. Hale' (pollen sterile) was used only as a mother plant at Pozantı Agricultural Research Center of Cukurova University. Seeds from crosses were grown in the orchards, of the Horticultural Department in Adana, under subtropical conditions. Morphological, pomological and isozyme

Among the peach hybrids, especially Rio-Oso-Gem × Ustun combination yielded very good results, and six hybrids of this combination were placed among 20 promising candidate cultivars. As a result of this study, several high-quality, very late ripening hybrids were identified [15]. The best results for late ripening, yield and fruit quality characteristics were obtained from the hybrids of Rio-Oso-Gem × Ustun combination, Nos. 24 and 19. These were followed by Ustun × Monroe 14, J.H. Hale × Rio-Oso-Gem 14, Rio-Oso-Gem × Ustun 21, and Independence open pollinated hybrid No. 8. The promising genotypes were planted at the orchards of Pozanti Agricultural Research and Application Center to see the real performances of these high chilling requiring peach and nectarine genotypes. According to the observations among the hybrids, some of them were found to be resistant to *Taphrina deformans*. The band profiles of 12 enzyme systems of parent cultivars were investigated and polymorphism obtained in 7 enzyme systems (MDH, PRX, ADH, AMY, IDH, PEP and ACP). In 31 hybrids, polymorphism in enzyme systems was found to be suitable to Mendel Segregation rates. As a result of this experiment, two very late ripening peach cultivars from Rio Oso Gem × Ustun combination and two peach cultivars from J.H. Hale open pollination were registered and patented. All these new cultivars are of good quality and high chilling requiring cultivars.

could be grown efficiently [5].

analysis were also carried out in Adana [14].

Peach (*Prunus persica* [L.] Batsch) and nectarine (*P. persica* var. nectarine Maxim) belong to the *Rosaceae* family. Peach is widely cultivated due to its easy adaptability to different ecological conditions, early fruit set and a long period of harvest. Peach cultivation extends along 30–45° north and south parallels of latitude. At higher elevations, low winter temperatures and late spring frosts are limiting factors for peaches [3]. Around 21,638,953 tons of peaches are produced in an area of 1,538,174 ha across the world. Turkey ranks number 6, with a production of 637,573 tons/year in 29,092 ha area, following China (11,924,085 tons), Italy (1,401,795 tons), Spain (1,329,800 tons), the USA (964,890 tons), and Greece (666,200 tons) [4].

Fruit breeders preferred to work on peach breeding for its early fruiting habit and correlative characteristics [5]. In cultivar or rootstock breeding programs, disease tolerant genotypes, early or late ripening habit, improvement of fruit quality characteristics such as shape, aroma, flesh firmness, especially low chilling requirements and to improve tree shape were aimed [6]. Important characteristics on peach breeding have already been determined to be; white fruit flesh is dominant to yellow fruit flesh; pubescence to smooth skin; soft fruit flesh to firm and freestone to clingstone. In crossbreeding of peaches and nectarines, since pubescence characteristics are controlled by single gene, generally heterozygote variation of 3:1 is seen. In addition, several correlations between fruit flesh color and receptacle and the color of leaves are seen. These correlative characteristics reveal that early selection is available in peach breeding [7, 8].

Classical plant breeding depends on the phenotypic selection of superior genotypes obtained as a result of crossbreeding. However, genotype × environment interaction causes time consumption and is quite difficult. Moreover, phenotypic selection is expensive and most of the time is not feasible for some characteristics like tolerance to abiotic stress conditions. Markerassisted selection is an approach developed as an alternative to these problems faced in classical plant breeding [9]. Usually, fruit demand of the consumers in peach is large weight and big-sized fruits, which are quantitative characters (QTLs) affected by several genes and various environmental conditions. Linge et al. [10] carried out an experiment on these characteristics and determined a genetic map of an F2 progeny with 117 individuals of PI91459 ('NJ Weeping') 9 'Bounty' with SNP markers. The fruit quality characters such as fruit weight, height, width, and depth of the progeny and parents were determined in 2011 and in 2012 and were compared with SNP markers. They found a positive correlation between characteristics of fruit weight and characteristics of fruit size. They also constructed a SNP map obtained from 1148 markers distributed across more than eight linkage groups. With this study, they identified 28 QTLs for these characters in which 11 of them were stable in both2011 and 2012 [10].

Various regions of Turkey are suitable to grow peaches under different ecological conditions. The Marmara region can grow high chilling requiring peaches, but the ripening time of the latest peach cultivars is about mid-September. A peach breeding study was carried out by Eroglu [11] in this important peach growing area. In this project, five foreign cultivars (Rio Oso Gem, Fortuna, Monroe, Jungerman and Vivian) and four local types (Bayramiç Tüysüzü, Alyanak Hulu, Sarı Papa and Takunyacı I) were hybridized to obtain fresh market peaches and two foreign and one local type peaches were hybridized for processing peaches. Eroglu et al. [12] stated the fruit quality performances of 121 peach genotypes for fresh market and 35 genotypes for processing. Adana province in Cukurova plain at the Mediterranean region has a subtropical climate. It is located very close to the Mediterranean Sea, thus this condition is suitable to grow only low or midseason chilling requiring peach cultivars. Within 1 h distance to this area, in Taurus Mountains (1100 m elevation) region, high chilling requiring cultivars could be grown efficiently [5].

desired fruits according to the aim of the breeding program [1]. Whole-genome sequences have been recently released for apple, peach, strawberry, Japanese apricot, and Chinese and European pear. After these developments, in breeding programs, marker-assisted selection has been used in wide genomics studies. These studies have caused to develop genome scale SNPs and SSR markers and to place reference linkage maps in Rosacea family to allow the identification of evolutionary relationships, which can be found in Genome Database in Rosaceae website. Yamamoto and Terakami [2] reviewed the recent advances in genomic studies and their practical applications for Rosaceae fruit trees, particularly in pear, apple,

Peach (*Prunus persica* [L.] Batsch) and nectarine (*P. persica* var. nectarine Maxim) belong to the *Rosaceae* family. Peach is widely cultivated due to its easy adaptability to different ecological conditions, early fruit set and a long period of harvest. Peach cultivation extends along 30–45° north and south parallels of latitude. At higher elevations, low winter temperatures and late spring frosts are limiting factors for peaches [3]. Around 21,638,953 tons of peaches are produced in an area of 1,538,174 ha across the world. Turkey ranks number 6, with a production of 637,573 tons/year in 29,092 ha area, following China (11,924,085 tons), Italy (1,401,795 tons),

Fruit breeders preferred to work on peach breeding for its early fruiting habit and correlative characteristics [5]. In cultivar or rootstock breeding programs, disease tolerant genotypes, early or late ripening habit, improvement of fruit quality characteristics such as shape, aroma, flesh firmness, especially low chilling requirements and to improve tree shape were aimed [6]. Important characteristics on peach breeding have already been determined to be; white fruit flesh is dominant to yellow fruit flesh; pubescence to smooth skin; soft fruit flesh to firm and freestone to clingstone. In crossbreeding of peaches and nectarines, since pubescence characteristics are controlled by single gene, generally heterozygote variation of 3:1 is seen. In addition, several correlations between fruit flesh color and receptacle and the color of leaves are seen. These correlative characteristics reveal that early selection is available in peach breeding [7, 8].

Classical plant breeding depends on the phenotypic selection of superior genotypes obtained as a result of crossbreeding. However, genotype × environment interaction causes time consumption and is quite difficult. Moreover, phenotypic selection is expensive and most of the time is not feasible for some characteristics like tolerance to abiotic stress conditions. Markerassisted selection is an approach developed as an alternative to these problems faced in classical plant breeding [9]. Usually, fruit demand of the consumers in peach is large weight and big-sized fruits, which are quantitative characters (QTLs) affected by several genes and various environmental conditions. Linge et al. [10] carried out an experiment on these characteristics and determined a genetic map of an F2 progeny with 117 individuals of PI91459 ('NJ Weeping') 9 'Bounty' with SNP markers. The fruit quality characters such as fruit weight, height, width, and depth of the progeny and parents were determined in 2011 and in 2012 and were compared with SNP markers. They found a positive correlation between characteristics of fruit weight and characteristics of fruit size. They also constructed a SNP map obtained from 1148 markers distributed across more than eight linkage groups. With this study, they identified 28 QTLs for

Spain (1,329,800 tons), the USA (964,890 tons), and Greece (666,200 tons) [4].

these characters in which 11 of them were stable in both2011 and 2012 [10].

peach, and cherry.

48 Breeding and Health Benefits of Fruit and Nut Crops

**a.** The first peach breeding study in Turkey was conducted by Tanriver and Kuden [13] on 'Ustun', a very late ripening (in mid-October) peach cultivar from the beginning of the 1990s. This study was carried out for [1] breeding of early cultivars for subtropical regions and [2] breeding of late ripening cultivars for cold regions, during 1995–1999. In the breeding experiments, carried out in Adana for early peach cultivars, Springtime, Suncrest, Flavorcrest and Redcap peach cultivars were used. Embryo rescue method was used with the combinations, where Springtime was used as a mother parent. As a result, embryo size was found to be important for the success of embryo rescue, and early cultivars should be used as pollinators. In the breeding studies to obtain late ripening peach cultivars, 'Monroe', 'Rio-Oso-Gem' and 'Ustun' were crossed reciprocally, while 'J.H. Hale' (pollen sterile) was used only as a mother plant at Pozantı Agricultural Research Center of Cukurova University. Seeds from crosses were grown in the orchards, of the Horticultural Department in Adana, under subtropical conditions. Morphological, pomological and isozyme analysis were also carried out in Adana [14].

Among the peach hybrids, especially Rio-Oso-Gem × Ustun combination yielded very good results, and six hybrids of this combination were placed among 20 promising candidate cultivars. As a result of this study, several high-quality, very late ripening hybrids were identified [15]. The best results for late ripening, yield and fruit quality characteristics were obtained from the hybrids of Rio-Oso-Gem × Ustun combination, Nos. 24 and 19. These were followed by Ustun × Monroe 14, J.H. Hale × Rio-Oso-Gem 14, Rio-Oso-Gem × Ustun 21, and Independence open pollinated hybrid No. 8. The promising genotypes were planted at the orchards of Pozanti Agricultural Research and Application Center to see the real performances of these high chilling requiring peach and nectarine genotypes. According to the observations among the hybrids, some of them were found to be resistant to *Taphrina deformans*. The band profiles of 12 enzyme systems of parent cultivars were investigated and polymorphism obtained in 7 enzyme systems (MDH, PRX, ADH, AMY, IDH, PEP and ACP). In 31 hybrids, polymorphism in enzyme systems was found to be suitable to Mendel Segregation rates. As a result of this experiment, two very late ripening peach cultivars from Rio Oso Gem × Ustun combination and two peach cultivars from J.H. Hale open pollination were registered and patented. All these new cultivars are of good quality and high chilling requiring cultivars.

**b.** The aim of the second peach breeding program was to improve some quality characteristics of Ustun peach to obtain high chilling, late ripening, good quality and good yielding peach, as well as nectarine cultivars. In this study, the pollen of Ustun cultivar was crossed with Venus and Stark Red Gold nectarines [16]. By crossing Venus and Stark Red Gold nectarines, with Ustun peach cultivar, 61 genotypes from VxU combination and 115 genotypes from SRGxU combination were obtained. A total of 176 genotypes were investigated for their morphological as well as phenological characteristics and were analyzed pomologically. Also, some pomological characters were compared by BPPCT009, MA014, MA040, and STS-OPAG8 SSR primer pairs to investigate the effectiveness of the marker-assisted selection in F1 genotypes.

The Sharka disease is a race such as PPV-D, PPV-M, and PPV-Rec, and new breeds can be taken out from a combination of these races. PPV-T is a combination of PPV-M and PPV-D races, and it was found to be a race belonging to Turkey. In this study, local apricot cultivars Hacıhaliloglu and Kabaası were crossed with foreign apricot cultivars such as Stark Early Orange, Rojo Pasion, Murciana, and P 1908 (peach clone from *Prunus davidiana*), which are known to be resistant to PPV. For peaches, commercial peach cultivars such as Flored and Carolina were crossed with PPV resistant Stark Early Orange (apricot) and P 1908 peach clone. In the hybridization studies, embryo rescue was performed with the combinations in which Flored peach variety was used as a mother parent, and in other combinations, the seeds were folded. Murashige & Skoog (MS) and Woody Plant Medium (WPM) nutrient media were used for embryo rescue combinations. Molecular studies were used to determine early resistance to Sharka disease in the hybridized individuals. Studies of the PGS1.21, PGS1.24, and ZP002 markers in hybrid subjects revealed the presence of resistance, tolerance, and susceptibility alleles. A total of 365 genotypes from crossing among 12 combinations of apricot and peach were tested with SSR markers (P GS1.21, PGS1.24, and ZP002). Approximately, 138 genotypes were found to be candidates for PPV resistance in future studies [18]. Individuals with endurance allele at the next stage of the study will be protected for other tests and observations to be made by

Peach Breeding Studies in Turkey and the Evaluation of Peach and Nectarine Hybrids

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51

**d.** Current peach breeding work is focused on breeding of low chilling, good quality genotypes, and to obtain flat and nonacid peaches (*Prunus persica* var. platycarpa). For this aim, Venus, Maycrest, Early Silver, and Gransun peach and nectarine cultivars and four flat and nonacid local peach genotypes, which were obtained by a selection study (Flat peach genotype 1, Flat peach genotype 2, Flat peach genotype 4, and Flat peach genotype 5), were used as parents. Phenological observations (blistering, green tip, pink tip, balloon, full bloom, and fall of petals) and pomological analyses (fruit weight, fruit height, fruit length, fruit width, total acid, pH, firmness, fruit top color, ground color, fruit pulp color, freestone state, fruit shape, and pubesence) were also studied. Wang et al. [19] stated that the nonacid peaches are preferred in the market, and this trait is usually selected in the commercial breeding programs. A major gene (D/d) located on chromosome 5 of peach has been described for this character, where the nonacid character is determined by the dominant D allele. Flavor analysis in fruit juice samples taken from genotypes will be carried out using the HS-GCMS technique, while sugar and organic acids will be carried out by the HPLC technique. Thus, the aroma, sugar, and organic acid levels of each individual

In this chapter, brief information on the results of the second (b) peach breeding experiment

In this breeding study, Ustun peach cultivar was used as a father parent, and Venus and Stark

hybrids obtained

Red Gold nectarine cultivars were used as mother parent. In the trial, 61 F1

grafting on the clone rootstock.

will be determined.

**2. Material and method**

at Cukurova University is provided.

In the weighted ranking method, VxU-55, VxU-41, VxU-34, VxU-14, VxU-1, VxU-13, VxU-24, VxU-26 peach genotypes and VxU-31, VxU-42, VxU-53, VxU-15 nectarine genotypes in VxU combination gave the highest points. In SRGxU combination, SRGxU-101, SRGxU-28, SRGxU-88, SRGxU-84, SRGxU-36, SRGxU-57, SRGxU-23, SRGxU-92, SRGxU-93 peach, and SRGxU-5 nectarine genotypes gave the highest points for the same scaling method.

No gene amplification was obtained from the PCR reactions among VxU and SRGxU populations analyzed by MA040 and STS-OPAG8 SSR primer pairs. BPPCT009 and MA014SSR primer pairs were also insufficient to determine fruit shape and free stone characteristics for VxU and SRGxU populations. These selected genotypes will eventually be taken under registration very soon.

Detailed information on this research is given below.

**c.** Another peach and apricot breeding program resistant to Sharka has been completed with a cooperative research between BETA Private Research Center, Malatya Fruit Research Institute and Cukurova University. This study aimed to obtain peach, nectarine, and apricot genotypes resistant to "Sharka" with crossbreeding method. Plum pox virus (PPV) causing Sharka disease is the most important viral agent for stone fruits. This disease is most harmful to apricot, plum, and peach trees. In Valencian Institute of Agricultural Research (IVIA), peach breeding program on apricot and on peach was started in 1993 and 1997, respectively. The aim of the study on apricot was to adapt to Southern Europe and to obtain high-quality cultivars resistant to the Sharka disease. Sharka or PPV is the most important limiting factor in the production of apricot. It was first described in Spain in 1984, causing serious loss of fruit and destruction of more than 1.5 million trees. The disease resistance breeding program is based on the transfer of resistance from local cultivars of Sharka disease to other cultivars by crossbreeding experiments. With 15 genotypes selected in accordance with the program's objectives, they were worked on apricot production areas in Spain. The aim of the peach breeding program was to obtain new cultivars of peach and nectarine that ripen early and provide good quality cultivars found in the market. The main market in Spain is the European countries, just as the big world producers are in the countries. In some parts of Valencia, Murcia, and Andulacia, climatic conditions allow the production of early cultivars that mature so as not to conflict with other European countries. In this study, 15 apricot genotypes and 12 peach genotypes selected for the purposes of the peach breeding program have been defined as resistant to the Sharka disease [17].

The Sharka disease is a race such as PPV-D, PPV-M, and PPV-Rec, and new breeds can be taken out from a combination of these races. PPV-T is a combination of PPV-M and PPV-D races, and it was found to be a race belonging to Turkey. In this study, local apricot cultivars Hacıhaliloglu and Kabaası were crossed with foreign apricot cultivars such as Stark Early Orange, Rojo Pasion, Murciana, and P 1908 (peach clone from *Prunus davidiana*), which are known to be resistant to PPV. For peaches, commercial peach cultivars such as Flored and Carolina were crossed with PPV resistant Stark Early Orange (apricot) and P 1908 peach clone. In the hybridization studies, embryo rescue was performed with the combinations in which Flored peach variety was used as a mother parent, and in other combinations, the seeds were folded. Murashige & Skoog (MS) and Woody Plant Medium (WPM) nutrient media were used for embryo rescue combinations. Molecular studies were used to determine early resistance to Sharka disease in the hybridized individuals. Studies of the PGS1.21, PGS1.24, and ZP002 markers in hybrid subjects revealed the presence of resistance, tolerance, and susceptibility alleles. A total of 365 genotypes from crossing among 12 combinations of apricot and peach were tested with SSR markers (P GS1.21, PGS1.24, and ZP002). Approximately, 138 genotypes were found to be candidates for PPV resistance in future studies [18]. Individuals with endurance allele at the next stage of the study will be protected for other tests and observations to be made by grafting on the clone rootstock.

**d.** Current peach breeding work is focused on breeding of low chilling, good quality genotypes, and to obtain flat and nonacid peaches (*Prunus persica* var. platycarpa). For this aim, Venus, Maycrest, Early Silver, and Gransun peach and nectarine cultivars and four flat and nonacid local peach genotypes, which were obtained by a selection study (Flat peach genotype 1, Flat peach genotype 2, Flat peach genotype 4, and Flat peach genotype 5), were used as parents. Phenological observations (blistering, green tip, pink tip, balloon, full bloom, and fall of petals) and pomological analyses (fruit weight, fruit height, fruit length, fruit width, total acid, pH, firmness, fruit top color, ground color, fruit pulp color, freestone state, fruit shape, and pubesence) were also studied. Wang et al. [19] stated that the nonacid peaches are preferred in the market, and this trait is usually selected in the commercial breeding programs. A major gene (D/d) located on chromosome 5 of peach has been described for this character, where the nonacid character is determined by the dominant D allele. Flavor analysis in fruit juice samples taken from genotypes will be carried out using the HS-GCMS technique, while sugar and organic acids will be carried out by the HPLC technique. Thus, the aroma, sugar, and organic acid levels of each individual will be determined.

In this chapter, brief information on the results of the second (b) peach breeding experiment at Cukurova University is provided.

#### **2. Material and method**

**b.** The aim of the second peach breeding program was to improve some quality characteristics of Ustun peach to obtain high chilling, late ripening, good quality and good yielding peach, as well as nectarine cultivars. In this study, the pollen of Ustun cultivar was crossed with Venus and Stark Red Gold nectarines [16]. By crossing Venus and Stark Red Gold nectarines, with Ustun peach cultivar, 61 genotypes from VxU combination and 115 genotypes from SRGxU combination were obtained. A total of 176 genotypes were investigated for their morphological as well as phenological characteristics and were analyzed pomologically. Also, some pomological characters were compared by BPPCT009, MA014, MA040, and STS-OPAG8 SSR primer pairs to investigate the effectiveness of the marker-assisted

In the weighted ranking method, VxU-55, VxU-41, VxU-34, VxU-14, VxU-1, VxU-13, VxU-24, VxU-26 peach genotypes and VxU-31, VxU-42, VxU-53, VxU-15 nectarine genotypes in VxU combination gave the highest points. In SRGxU combination, SRGxU-101, SRGxU-28, SRGxU-88, SRGxU-84, SRGxU-36, SRGxU-57, SRGxU-23, SRGxU-92, SRGxU-93 peach, and SRGxU-5 nectarine genotypes gave the highest points for the same scaling method.

No gene amplification was obtained from the PCR reactions among VxU and SRGxU populations analyzed by MA040 and STS-OPAG8 SSR primer pairs. BPPCT009 and MA014SSR primer pairs were also insufficient to determine fruit shape and free stone characteristics for VxU and SRGxU populations. These selected genotypes will eventually

**c.** Another peach and apricot breeding program resistant to Sharka has been completed with a cooperative research between BETA Private Research Center, Malatya Fruit Research Institute and Cukurova University. This study aimed to obtain peach, nectarine, and apricot genotypes resistant to "Sharka" with crossbreeding method. Plum pox virus (PPV) causing Sharka disease is the most important viral agent for stone fruits. This disease is most harmful to apricot, plum, and peach trees. In Valencian Institute of Agricultural Research (IVIA), peach breeding program on apricot and on peach was started in 1993 and 1997, respectively. The aim of the study on apricot was to adapt to Southern Europe and to obtain high-quality cultivars resistant to the Sharka disease. Sharka or PPV is the most important limiting factor in the production of apricot. It was first described in Spain in 1984, causing serious loss of fruit and destruction of more than 1.5 million trees. The disease resistance breeding program is based on the transfer of resistance from local cultivars of Sharka disease to other cultivars by crossbreeding experiments. With 15 genotypes selected in accordance with the program's objectives, they were worked on apricot production areas in Spain. The aim of the peach breeding program was to obtain new cultivars of peach and nectarine that ripen early and provide good quality cultivars found in the market. The main market in Spain is the European countries, just as the big world producers are in the countries. In some parts of Valencia, Murcia, and Andulacia, climatic conditions allow the production of early cultivars that mature so as not to conflict with other European countries. In this study, 15 apricot genotypes and 12 peach genotypes selected for the purposes of the

peach breeding program have been defined as resistant to the Sharka disease [17].

selection in F1

genotypes.

50 Breeding and Health Benefits of Fruit and Nut Crops

be taken under registration very soon.

Detailed information on this research is given below.

In this breeding study, Ustun peach cultivar was used as a father parent, and Venus and Stark Red Gold nectarine cultivars were used as mother parent. In the trial, 61 F1 hybrids obtained from Venus × Ustun crossbreeding and 115 F1 hybrids obtained from Stark Red Gold × Ustun crossbreeding, and a total of 176 F1 hybrids were used as plant material.

**3. Results and discussion**

**3.1. The morphological, phenological, and pomological analyses**

hybrids showed the characteristics of rosaceae and campanula flower [16].

obtained from SRGxU-32 (154.30 g) and SRGxU-82 (150.90 g) genotypes.

Harvest date 20 Fruit shape 7 Fruit weight 20 Fruit ground color 7 Brix 13 Fruit tip state 5 Attractiveness 12 Red color under skin 3 Freestone state 10 Red color around seed 3

obtained; In VxU combination, the highest value (87.65 mm) was obtained from VxU-18 genotype, whereas the lowest one (33.54 mm) was obtained from VxU-56 genotype. In SRGxU combination, the highest trunk diameter value (99.44 mm) was determined in SRGxU-9 genotype, while the lowest one (21.92 mm) was determined in SRGxU-100 genotype. The genotypes reached full bloom phase on March 11, the earliest and on April 6, the latest. The

Peach Breeding Studies in Turkey and the Evaluation of Peach and Nectarine Hybrids

under the subtropical conditions of Adana at the Cukurova University (50 m elevation). The genotypes could not show their best performances in Adana, but they gave us early selection opportunities. Although to the low chilling and sea-level conditions in Adana, some genotypes showed very late ripening habit such as, September 3–18. If we consider Ustun late peach cultivar is ripening at the end of September in Pozanti, some of these genotypes could

The highest mean fruit weights in VxU combination were obtained from VxU-58 (137.30 g) and VxU-55 (136.95 g) genotypes. In SRGxU combination, the highest fruit weights were

Brix values among all hybrids (**Table 1**) were the highest in VxU-4 (14.40), SRGxU-110 (13.0), and SRGxU-111 (13.0) genotypes. These results are in accordance with the results of Tanriver and Kuden [13] and Monet [7, 20, 23] who stated that the hybrids showed their best performances (fruit weight, color, Brix value, and especially for their harvesting dates) in the areas convenient for their chilling requirements. Harvesting dates and fruit characteristics of the

The data obtained from the observations and analyzes were weighed out in the individuals in both combinations. According to results, VxU-55, VxU-41, VxU-34, VxU-14, VxU-1, Vx-13, VxU-24, and VxU-26 genotypes gave the best results among the peach genotypes in VxU combination. In the same combination, VxU-31, VxU-42, VxU-53, and VxU-15 were found to be the best nectarine genotypes. In the other combination (SRGxU), the performances were obtained from SRGxU-101, SRGxU-28, SRGxU-88, SRGxU-84, SRGxU-36, SRGxU-57, SRGxU-23, SRGxU-92, and SRGxU-93 peach genotypes and SRGxU-5 nectarine genotype (**Table 1**).

hybrids, following results were

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53

seeds of the genotypes were sown and grown

If we consider the trunk diameter measurements on F1

be ripen later than the parents under Pozanti conditions.

To get faster growth and earlier fruit set, F1

selected genotypes are given in **Tables 2** and **3**.

**Table 1.** Weighted grading types and scores.

Ustun peach cultivar has been emerged as a result of bud mutation from peach cultivar of J.H.H. It matures between the first week and the third week of October (1400–1500 altitude). Fruit peel is fairly hairy, red cheek on yellow ground, fruit stalk is short, flesh is hard and yellow color and it has a very good aroma (**Figure 1**).

Venus and Stark Red Gold are kinds of nectarine with a yellow flesh. They have quite good properties in terms of fruit weight, color, and taste. Venus matures 10 days later than the Stark Red Gold.

#### **2.1. Phenological, morphological, and pomological analyses**

A total of 176 F1 hybrids, that is, 61 from Venus × Ustun crosses and 115 from Stark Red Gold × Ustun crosses were compared for their correlative, phenological (pink tip, balloon, first bloom, full bloom, harvest date), morphological (plant length and trunk diameter, tree form, flower property, flower color), and pomological (fruit weight, fruit height, fruit length, fruit width, seed weight, Brix, % acidity, pH, pulp/seed rate, fruit shape, fruit tip, rupture state of the fruit, pubesence, fruit top color, fruit ground color, red blush in flesh, fruit attractiveness, firmness, taste, freestone, and fruit flesh color) characteristics in this experiment [20, 21]. Also, the marker-assisted selection effectiveness of F<sup>1</sup> peach hybrids was determined by two SSR markers.

#### **2.2. Molecular analysis (SSR)**

High molecular weight genomic DNA was extracted from the leaf samples of each F1 hybrids for SSR analysis. SSR analysis was performed in accordance with Aka-Kacar et al. [22]. Two primer pairs (BPPCT009/b, MA014a) were used to generate the SSR genotyping. DNA profiles of parents and F1 hybrids were recorded, and their allelic profiles were compared. Results obtained from the SSR analysis were evaluated with the pomological features of genotypes.

**Figure 1.** Ustun local peach cultivar.

## **3. Results and discussion**

from Venus × Ustun crossbreeding and 115 F1

low color and it has a very good aroma (**Figure 1**).

the marker-assisted selection effectiveness of F<sup>1</sup>

**2.2. Molecular analysis (SSR)**

**Figure 1.** Ustun local peach cultivar.

**2.1. Phenological, morphological, and pomological analyses**

crossbreeding, and a total of 176 F1

52 Breeding and Health Benefits of Fruit and Nut Crops

Red Gold.

markers.

A total of 176 F1

hybrids obtained from Stark Red Gold × Ustun

peach hybrids was determined by two SSR

hybrids

hybrids were used as plant material.

hybrids, that is, 61 from Venus × Ustun crosses and 115 from Stark Red

Ustun peach cultivar has been emerged as a result of bud mutation from peach cultivar of J.H.H. It matures between the first week and the third week of October (1400–1500 altitude). Fruit peel is fairly hairy, red cheek on yellow ground, fruit stalk is short, flesh is hard and yel-

Venus and Stark Red Gold are kinds of nectarine with a yellow flesh. They have quite good properties in terms of fruit weight, color, and taste. Venus matures 10 days later than the Stark

Gold × Ustun crosses were compared for their correlative, phenological (pink tip, balloon, first bloom, full bloom, harvest date), morphological (plant length and trunk diameter, tree form, flower property, flower color), and pomological (fruit weight, fruit height, fruit length, fruit width, seed weight, Brix, % acidity, pH, pulp/seed rate, fruit shape, fruit tip, rupture state of the fruit, pubesence, fruit top color, fruit ground color, red blush in flesh, fruit attractiveness, firmness, taste, freestone, and fruit flesh color) characteristics in this experiment [20, 21]. Also,

High molecular weight genomic DNA was extracted from the leaf samples of each F1

for SSR analysis. SSR analysis was performed in accordance with Aka-Kacar et al. [22]. Two primer pairs (BPPCT009/b, MA014a) were used to generate the SSR genotyping. DNA profiles of parents and F1 hybrids were recorded, and their allelic profiles were compared. Results obtained from the SSR analysis were evaluated with the pomological features of genotypes.

#### **3.1. The morphological, phenological, and pomological analyses**

If we consider the trunk diameter measurements on F1 hybrids, following results were obtained; In VxU combination, the highest value (87.65 mm) was obtained from VxU-18 genotype, whereas the lowest one (33.54 mm) was obtained from VxU-56 genotype. In SRGxU combination, the highest trunk diameter value (99.44 mm) was determined in SRGxU-9 genotype, while the lowest one (21.92 mm) was determined in SRGxU-100 genotype. The genotypes reached full bloom phase on March 11, the earliest and on April 6, the latest. The hybrids showed the characteristics of rosaceae and campanula flower [16].

To get faster growth and earlier fruit set, F1 seeds of the genotypes were sown and grown under the subtropical conditions of Adana at the Cukurova University (50 m elevation). The genotypes could not show their best performances in Adana, but they gave us early selection opportunities. Although to the low chilling and sea-level conditions in Adana, some genotypes showed very late ripening habit such as, September 3–18. If we consider Ustun late peach cultivar is ripening at the end of September in Pozanti, some of these genotypes could be ripen later than the parents under Pozanti conditions.

The highest mean fruit weights in VxU combination were obtained from VxU-58 (137.30 g) and VxU-55 (136.95 g) genotypes. In SRGxU combination, the highest fruit weights were obtained from SRGxU-32 (154.30 g) and SRGxU-82 (150.90 g) genotypes.

Brix values among all hybrids (**Table 1**) were the highest in VxU-4 (14.40), SRGxU-110 (13.0), and SRGxU-111 (13.0) genotypes. These results are in accordance with the results of Tanriver and Kuden [13] and Monet [7, 20, 23] who stated that the hybrids showed their best performances (fruit weight, color, Brix value, and especially for their harvesting dates) in the areas convenient for their chilling requirements. Harvesting dates and fruit characteristics of the selected genotypes are given in **Tables 2** and **3**.

The data obtained from the observations and analyzes were weighed out in the individuals in both combinations. According to results, VxU-55, VxU-41, VxU-34, VxU-14, VxU-1, Vx-13, VxU-24, and VxU-26 genotypes gave the best results among the peach genotypes in VxU combination. In the same combination, VxU-31, VxU-42, VxU-53, and VxU-15 were found to be the best nectarine genotypes. In the other combination (SRGxU), the performances were obtained from SRGxU-101, SRGxU-28, SRGxU-88, SRGxU-84, SRGxU-36, SRGxU-57, SRGxU-23, SRGxU-92, and SRGxU-93 peach genotypes and SRGxU-5 nectarine genotype (**Table 1**).


**Table 1.** Weighted grading types and scores.


**Table 2.** Harvesting dates and the fruit characteristics of the selected genotypes.

#### **3.2. The molecular analysis**

A total of 176 F1 hybrids were examined by using two different SSR primer pairs for early marker-assisted selection. The fruit characteristics of some F1 hybrids are found to be different as compared to their parents, while some of them were almost found to be the same.

correct results. As a conclusion, for early selection criteria, SSR primer pairs are good molecular markers to be used for this aim. However, in this research, we could not obtain a very good

Dirlewanger et al. [24] stated that MA014a SSR primer pair is associated with fruit shape and flat fruit. In our experiments, we found that MA014a primer pair was more suitable to VxU population, but not for SRGxU population. For freestone character, BPPCT009/b SSR primer pair was used (**Figure 2**) to determine the freestone character. This primer was not compatible

not match each other properly. The photos of some of the selected genotypes are shown in

populations. Freestone characters of hybrids and allelic profiles did

**) Freestone Fruit shape Pulp color Pubescence**

Peach Breeding Studies in Turkey and the Evaluation of Peach and Nectarine Hybrids

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55

VxU-55 2.92 Semi freestone Oval Yellow Medium VxU-41 2.67 Semi freestone Round Yellow Medium VxU-34 2.19 Cling stone Round Yellow Medium VxU-14 2.06 Cling stone Round Yellow Medium VxU-1 3.12 Freestone Oval Yellow Medium VxU-13 1.88 Cling stone Oval Yellow Medium VxU-24 2.68 Freestone Oval Yellow Medium VxU-26 2.9 Semi freestone Oval Yellow Medium VxU-31 1.5 Freestone Oval Yellow Nectarine VxU-42 3.10 Cling stone Oval Yellow Nectarine VxU-53 3.06 Cling stone Oval White-Red Nectarine VxU-15 2.28 Semi freestone Round White-Red Nectarine SRGxU-101 2.07 Freestone Oval Yellow Medium SRGxU-28 3.26 Freestone Round Yellow Medium SRGxU-88 2.92 Semi freestone Oval Yellow Medium SRGxU-84 2.65 Cling stone Oval Yellow Medium SRGxU-36 1.86 Freestone Round Yellow Medium SRGxU-57 2.75 Freestone Round Yellow Medium SRGxU-23 2.45 Freestone Round Yellow Medium SRGxU-92 2.34 Semi freestone Oval Yellow Medium SRGxU-93 3.47 Cling stone Round Yellow Medium SRGxU-5 2.75 Freestone Round Yellow Nectarine

relationship among our hybrids.

**Table 3.** The fruit characteristics of the selected genotypes.

**Genotypes Firmness (kg/cm2**

for SRGxU and VxU F1

**Figures 3**–**16**.

Among the molecular data determined from MA014a primer pair, with pomological analysis of hybrids, more correct results were obtained from MA014a primer pair in the genotypes of SRGxU combination (42.02%) and in the genotypes of VxU combination (85.29%). Also, BPPCT009/b primer pair was used to determine freestone characteristics of the hybrids. SRGxU combination gave 38.35% of correct results, and VxU combination gave 30.50% of


**Table 3.** The fruit characteristics of the selected genotypes.

**3.2. The molecular analysis**

54 Breeding and Health Benefits of Fruit and Nut Crops

marker-assisted selection. The fruit characteristics of some F1

**Table 2.** Harvesting dates and the fruit characteristics of the selected genotypes.

as compared to their parents, while some of them were almost found to be the same.

**Genotypes Harvest date Fruit weight (g) TSS (%) Acidity** VxU-55 08/14–27 136.95 12 0.73 VxU-41 08/27 173.20 12 0.68 VxU-34 08/26 163.10 15 0.64 VxU-14 08/26 171.28 12 0.68 VxU-1 07/12–21 108.60 12.9 0.75 VxU-13 08/20–27 120.12 13.15 0.82 VxU-24 08/1–13 142.25 11 0.54 VxU-26 08/14–09/18 129.15 12.9 0.66 VxU-31 09/03 79.8 13 0.77 VxU-42 08/02–09/03 108.23 11.5 0.51 VxU-53 06/28 131.74 10 1.03 VxU-15 06/10–29 87.42 10 0.60 SRGxU-101 08/14 161.75 12 0.53 SRGxU-28 07/24 109.8 12 0.50 SRGxU-88 08/14–27 136.95 12 0.48 SRGxU-84 07/04 164.21 12 0.67 SRGxU-36 08/12 152.89 10 0.38 SRGxU-57 07/27 106.6 10.5 0.66 SRGxU-23 07/23–08/27 118.37 11 0.69 SRGxU-92 07/12 110.64 10 0.58 SRGxU-93 07/08 209.17 12 0.46 SRGxU-5 08/24 108.09 11.5 0.49

Among the molecular data determined from MA014a primer pair, with pomological analysis of hybrids, more correct results were obtained from MA014a primer pair in the genotypes of SRGxU combination (42.02%) and in the genotypes of VxU combination (85.29%). Also, BPPCT009/b primer pair was used to determine freestone characteristics of the hybrids. SRGxU combination gave 38.35% of correct results, and VxU combination gave 30.50% of

hybrids were examined by using two different SSR primer pairs for early

hybrids are found to be different

A total of 176 F1

correct results. As a conclusion, for early selection criteria, SSR primer pairs are good molecular markers to be used for this aim. However, in this research, we could not obtain a very good relationship among our hybrids.

Dirlewanger et al. [24] stated that MA014a SSR primer pair is associated with fruit shape and flat fruit. In our experiments, we found that MA014a primer pair was more suitable to VxU population, but not for SRGxU population. For freestone character, BPPCT009/b SSR primer pair was used (**Figure 2**) to determine the freestone character. This primer was not compatible for SRGxU and VxU F1 populations. Freestone characters of hybrids and allelic profiles did not match each other properly. The photos of some of the selected genotypes are shown in **Figures 3**–**16**.

**Figure 6.** Fruits of VxU-1 genotype.

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**Figure 7.** Fruits of VxU-13 genotype.

**Figure 8.** Fruits of VxU-24 genotype.

**Figure 9.** Fruits of VxU-26 genotype.

**Figure 2.** Polyacrylamide gel image of SSR bands of BPPCT009 primer pair with SRGxU population.

**Figure 3.** Fruits of VxU-55 genotype.

**Figure 4.** Fruits of VxU-41genotype.

**Figure 5.** Fruits of VxU-14 genotype.

Peach Breeding Studies in Turkey and the Evaluation of Peach and Nectarine Hybrids http://dx.doi.org/10.5772/intechopen.73440 57

#### **Figure 6.** Fruits of VxU-1 genotype.

**Figure 7.** Fruits of VxU-13 genotype.

**Figure 3.** Fruits of VxU-55 genotype.

56 Breeding and Health Benefits of Fruit and Nut Crops

**Figure 4.** Fruits of VxU-41genotype.

**Figure 5.** Fruits of VxU-14 genotype.

**Figure 2.** Polyacrylamide gel image of SSR bands of BPPCT009 primer pair with SRGxU population.

**Figure 8.** Fruits of VxU-24 genotype.

**Figure 9.** Fruits of VxU-26 genotype.

**Figure 10.** Fruits of SRGxU-101 genotype.

**Figure 11.** Fruits of SRGxU-28 genotype.

**Figure 12.** Fruits of SRGxU-88 genotype.

**4. Conclusion**

**Figure 14.** Fruits of VxU-53 genotype.

**Figure 15.** Fruits of VxU-15 genotype.

**Figure 16.** Fruits of SRGxU-5 genotype.

higher elevation conditions (1100 m).

This peach breeding study was carried out to obtain late ripening, high chilling, good quality peaches, and nectarines suitable for more continental climates. As a result of the experiment, some genotypes were found to be later fruit ripening producers than their parents (September 3–18) under Adana subtropical climatic conditions (23 m elevation). The results showed that these late genotypes could ripen later under the continental climatic conditions. This means that these genotypes could have better performances at Taurus Mountains in Pozanti under

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**Figure 13.** Fruits of SRGxU-84 genotype.

Peach Breeding Studies in Turkey and the Evaluation of Peach and Nectarine Hybrids http://dx.doi.org/10.5772/intechopen.73440 59

**Figure 14.** Fruits of VxU-53 genotype.

**Figure 15.** Fruits of VxU-15 genotype.

**Figure 16.** Fruits of SRGxU-5 genotype.

#### **4. Conclusion**

**Figure 13.** Fruits of SRGxU-84 genotype.

**Figure 10.** Fruits of SRGxU-101 genotype.

58 Breeding and Health Benefits of Fruit and Nut Crops

**Figure 11.** Fruits of SRGxU-28 genotype.

**Figure 12.** Fruits of SRGxU-88 genotype.

This peach breeding study was carried out to obtain late ripening, high chilling, good quality peaches, and nectarines suitable for more continental climates. As a result of the experiment, some genotypes were found to be later fruit ripening producers than their parents (September 3–18) under Adana subtropical climatic conditions (23 m elevation). The results showed that these late genotypes could ripen later under the continental climatic conditions. This means that these genotypes could have better performances at Taurus Mountains in Pozanti under higher elevation conditions (1100 m).

One of the parents of these genotypes was Ustun, late peach cultivar, which ripened at the end of September in Pozanti. This observation lead us to think that some of these genotypes ripen on 3rd-18th September. Under Adana subtropical climatic conditions, they could ripen later than their parents in Pozantı (may be at the end of September or at the beginning of October). Thus, this will be a very good opportunity to get very high market prices with these very late season peaches and nectarines.

[4] FAO, Food and Agricultural Organization. 2013. Available from: www.fao.org/statistical

Peach Breeding Studies in Turkey and the Evaluation of Peach and Nectarine Hybrids

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61

[5] Kuden AB, Kuden A. Şeftali Yetiştiriciliği [Peach Growing]. Yayınları: Tübitak Tarp; 2000. 20 p [6] Fideghelli C, Della Strada G, Grassi F, Morico G. The peach industry in the world:

[8] Küden AB, Tanrıver E. Şeftalilerde Değişik Islah Yöntemleriyle Yeni Çeşit Eldesi ve İzoenzim Analizleri İle Genotiplerin Tanımlanması. Ç.Ü.Z.F. Dergisi. 1996;**11**(4):183-192

[9] Francia E, Rızza F, Cattivelli L, Stanca A, Galiba G, Toth B, Hayes P, Skinner JS, Pecchioni N. Two loci on chromosome 5H determine low-temperature tolerance in a 'Nure' [winter] × 'Tremois' [spring] barley map. Theoretical and Applied Genetics. 2004;**108**:670-680.

[10] Linge CS, Bassi D, Bianco L, Pacheco I, Pirona R, Rossini L. Genetic dissection of fruit weight and size in an F2 peach (*Prunus persica* (L.) Batsch) progeny. Molecular Breeding.

[11] Eroglu ZO. Melezleme Yoluyla Şeftali Çeşit Islahı [Peach Cultivar Crossbreeding]. [PhD thesis] Izmir, Turkey: Ege Univ., Institute of Natural and Applied Science; 2012. p. 178

[12] Eroglu ZO, Misirli A, Kuden AB. Bazı Şeftali Çeşitlerinin Melezleme Performansları. [Cross Breeding Performances of Some Peach Cultivars]. Yüzüncü Yıl Üniversitesi Tarım

[13] Tanriver E, Kuden AB. Breeding studies with Ustun peach. Eucarpia symposium on fruit

[14] Kuden AB, Tanriver E, Gulen H, Buyukalaca S. Embryo Rescue of Peach Breds. In: Proceedings of the Eucarpia Symposium on Fruit Breeding and Genetics. Oxford,

[15] Tanriver E. Cross breeding studies on peaches. [PhD thesis] P 246. Adana, Turkey:

[16] Comlekcioglu S. Identification of Vegetable Properties of Nectarine and Peach Hybrids

[17] Martinez-Calvo J, Font A, Llacer G, Badenes ML. Apricot and peach breeding programs

[18] Unek C. Breeding Studies on Resistance to Sharka on Peach, Nectarine and Apricots. [PhD thesis]. P 151. Adana, Turkey: Institute of Natural and Applied Science; 2015 [19] Wang Q, Wang L, Zhu G, Cao K, Fang W, Chen Wang X. DNA marker-assisted evaluation of fruit acidity in diverse peach (*Prunus persica*) germplasm. Euphytica. 2016;**210**:

and Availability of Some SSR Markers in Early Selection. [PhD thesis] 2014

[7] Monet R. Le pêcher. Masson, Paris, France: Genetique et physiolgie; 1983. 133 p

Present situation and trend. Acta Horticulturae. 1998;**465**:29-40

(Institute of Natural and Applied Science, Adana, Turkey, p. 209)

breeding and genetics. Acta Horticulturae. 2004;**663**(2):733-738

G.Britain. Acta Horticulture. 2nd-5th September 1996;(486):1-6, 531-534

database

2015;**35**:71

413-426

Bilimleri Dergisi. 2016;**26**(1):89-97

Institute of Natural and Applied Science; 2000

from the IVIA. Acta Horticulturae. 2009;**814**(1):185-188

Considering the fruit quality characteristics of the genotypes, among F1 hybrids, VxU-18 and SRGxU-9 genotypes gave better trunk development than the others. For the fruit weight characteristics, SRGxU-32, SRGxU-82, VxU-58, and VxU-55 gave the biggest fruits among all the genotypes. The highest brix values were obtained from VxU-4, SRGxU-110, and SRGxU-111 genotypes.

In conclusion, the genotypes of VxU-55, VxU-41, VxU-34, VxU-14, VxU-1, Vx-13, VxU-24, VxU-26, SRGxU-101, SRGxU-28, SRGxU-88, SRGxU-84, SRGxU-36, SRGxU-57, SRGxU-23, SRGxU-92 and SRGxU-93 peaches and VxU-31, VxU-42, VxU-53, VxU-15 and SRGxU-5 nectarines were found to be promising ones.

The selected genotypes which were grafted onto GF-677 rootstock were taken to Pozanti Agricultural Research and Application Center at the Taurus Mountains in Pozanti to compare their performances at high chilling area under second selection. Also in the future studies, more locus specific molecular marker systems such as SSRs or SNPs could be used to find out better characterization of the hybrid populations.

### **Author details**

Ayzin Baykam Kuden<sup>1</sup> \*, Songul Comlekcioglu1 , Kadir Sarier<sup>2</sup> , Burhanettin Imrak<sup>2</sup> and Ali Kuden<sup>1</sup>

\*Address all correspondence to: abkuden@gmail.com

1 Department of Horticulture, University of Cukurova, Faculty of Agriculture, Saricam, Adana, Turkey

2 Pozanti Agricultural Research and Application Center, University of Cukurova, Saricam, Adana, Turkey

### **References**


One of the parents of these genotypes was Ustun, late peach cultivar, which ripened at the end of September in Pozanti. This observation lead us to think that some of these genotypes ripen on 3rd-18th September. Under Adana subtropical climatic conditions, they could ripen later than their parents in Pozantı (may be at the end of September or at the beginning of October). Thus, this will be a very good opportunity to get very high market prices with these very late

Considering the fruit quality characteristics of the genotypes, among F1 hybrids, VxU-18 and SRGxU-9 genotypes gave better trunk development than the others. For the fruit weight characteristics, SRGxU-32, SRGxU-82, VxU-58, and VxU-55 gave the biggest fruits among all the genotypes. The highest brix values were obtained from VxU-4, SRGxU-110, and SRGxU-111 genotypes. In conclusion, the genotypes of VxU-55, VxU-41, VxU-34, VxU-14, VxU-1, Vx-13, VxU-24, VxU-26, SRGxU-101, SRGxU-28, SRGxU-88, SRGxU-84, SRGxU-36, SRGxU-57, SRGxU-23, SRGxU-92 and SRGxU-93 peaches and VxU-31, VxU-42, VxU-53, VxU-15 and SRGxU-5 nec-

The selected genotypes which were grafted onto GF-677 rootstock were taken to Pozanti Agricultural Research and Application Center at the Taurus Mountains in Pozanti to compare their performances at high chilling area under second selection. Also in the future studies, more locus specific molecular marker systems such as SSRs or SNPs could be used to find out

1 Department of Horticulture, University of Cukurova, Faculty of Agriculture, Saricam,

2 Pozanti Agricultural Research and Application Center, University of Cukurova, Saricam,

[1] Iwata H, Minamikawa MF, Kajiya-Kanegae H, Ishimori M, Hayashi T. Genomics-assisted

[2] Yamamoto T, Terakami S. Genomics of pear and other Rosaceae fruit trees. Breeding

[3] Ozbek S. Özel Meyvecilik [Fruit Growing]. Ç.Ü.Z.F. Yayinlari: Ders Kitabi; 1978. 11 p

, Kadir Sarier<sup>2</sup>

, Burhanettin Imrak<sup>2</sup>

and

season peaches and nectarines.

60 Breeding and Health Benefits of Fruit and Nut Crops

tarines were found to be promising ones.

**Author details**

Ali Kuden<sup>1</sup>

Adana, Turkey

Adana, Turkey

**References**

Science. 2016;**66**:148-159

Ayzin Baykam Kuden<sup>1</sup>

better characterization of the hybrid populations.

\*Address all correspondence to: abkuden@gmail.com

\*, Songul Comlekcioglu1

breeding in fruit trees. Breeding Science. 2016;**66**:100-115


[20] Monet R. Peach Tree. Cropping Techniques and Nurseries. Bari: Instituto Agronomico Mediterraneo; 1995. 42 p

**Chapter 4**

**Provisional chapter**

**Genetic Apricot Resources and their Utilisation in**

**Genetic Apricot Resources and their Utilisation in** 

DOI: 10.5772/intechopen.77125

This chapter outlines the evolution of apricot which took place not only in its original gene centers but also after its domestication in new, secondary areas. During this process, Ice Age, migration of nations as well as the influence of mountains played a significant role in the diversity of this fruit species where many clones of genetically similar cultivars and ecological groups of apricots were formed. The chapter presents the list of donors of main biological and economic properties which are important in breeding to increase the adaptability of the species. The chapter summarizes some of the breeding results and inheritance of characters related to frost hardiness of blossom buds, fruits and plum pox

Man has participated in the evolution of cultivated plants by selection as well as by controlled evolution, that is, crossing. In the past, most or all wild fruit trees possessed certain properties that were beneficial and tempting for humans. Plants were also know for their changeability due to the influence of external conditions but also to breeding with related varieties. As a result, even in their wild form, hybrids with complex genetic bases were created. Their seeds gave rise to many distinct types which were preserved and in fruit trees this initiated either accidental or intentionally developed bud mutations which also led to a greater diversification of species. This method of propagation is being used in some areas of Central Asia and China to this day, and, as a result, even within the European group of apricots, we have been

**Keywords:** apricot, *Prunus armeniaca* L., germplasm, inheritance, breeding

© 2016 The Author(s). Licensee InTech. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

© 2018 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use,

distribution, and reproduction in any medium, provided the original work is properly cited.

Additional information is available at the end of the chapter

Additional information is available at the end of the chapter

http://dx.doi.org/10.5772/intechopen.77125

**Breeding**

**Breeding**

Boris Krška

Boris Krška

**Abstract**

virus (PPV).

**1. Introduction**


#### **Genetic Apricot Resources and their Utilisation in Breeding Genetic Apricot Resources and their Utilisation in Breeding**

DOI: 10.5772/intechopen.77125

#### Boris Krška Boris Krška

[20] Monet R. Peach Tree. Cropping Techniques and Nurseries. Bari: Instituto Agronomico

[21] Kuden AB, Kaska N. Bazı şeftali ve nektarin çeşitlerinin soğuklama gereksinimleri ve büyüme derece saatleri toplamının çeşitli yöntemlerle saptanması [Determination of chilling requirements and growth degree hours of some peach and nectarin cultivars].

[22] Aka-Kacar Y, Simsek O, Solmaz I, Sari N, Yalcın Mendi Y. Genetic diversity among melon accessions [*Cucumis melo* L.] from Turkey as revealed by SSR markers. Genetic

[23] Monet R, Guye A, Roy M, Dachary N. Peach Mendelian genetics: A short review and

[24] Dirlewanger E, Cosson P, Boudehri K, Renaud C, Capdeville G, Tauzin Y, Laigret F, Moing A. Development of a second-generation genetic linkage map for peach [*Prunus Persica* [L.] Batsch] and characterization of morphological traits affecting flower and

Mediterraneo; 1995. 42 p

62 Breeding and Health Benefits of Fruit and Nut Crops

Bahçe. 1995;**18**(1-2):35-44

and Molecular Research. 2012;**11**(4):4622-4631

new results. Agronomie. 1996;**16**:321-329

fruit. Tree Genetics & Genomes. 2006;**3**:1-13

Additional information is available at the end of the chapter Additional information is available at the end of the chapter

http://dx.doi.org/10.5772/intechopen.77125

#### **Abstract**

This chapter outlines the evolution of apricot which took place not only in its original gene centers but also after its domestication in new, secondary areas. During this process, Ice Age, migration of nations as well as the influence of mountains played a significant role in the diversity of this fruit species where many clones of genetically similar cultivars and ecological groups of apricots were formed. The chapter presents the list of donors of main biological and economic properties which are important in breeding to increase the adaptability of the species. The chapter summarizes some of the breeding results and inheritance of characters related to frost hardiness of blossom buds, fruits and plum pox virus (PPV).

**Keywords:** apricot, *Prunus armeniaca* L., germplasm, inheritance, breeding

#### **1. Introduction**

Man has participated in the evolution of cultivated plants by selection as well as by controlled evolution, that is, crossing. In the past, most or all wild fruit trees possessed certain properties that were beneficial and tempting for humans. Plants were also know for their changeability due to the influence of external conditions but also to breeding with related varieties. As a result, even in their wild form, hybrids with complex genetic bases were created. Their seeds gave rise to many distinct types which were preserved and in fruit trees this initiated either accidental or intentionally developed bud mutations which also led to a greater diversification of species. This method of propagation is being used in some areas of Central Asia and China to this day, and, as a result, even within the European group of apricots, we have been

© 2016 The Author(s). Licensee InTech. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. © 2018 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

able to select some interesting varieties with higher adaptability to the environment (Sucre de Bohutice, Rosa Early, Holubova, Pourtáleská, Keczkemete Rozsa, Kamenickyi and so on).

**2. Domestication and selection of apricots**

**2.1. Domestication and genetic diversity of apricots**

among both domesticated and wild apricot trees.

The origin of all eco-geographic groups of apricot dates to the beginning of the Tertiary Period and is associated with the Northern and Middle China centers, which gave origin to more than 100 species of stone fruit, particularly to cherry, peach and apricot. In the Tertiary Period, apricot trees were found abundantly in extensive mountainous parts of Northern and Middle China, where, because of changing conditions, a process of forming and creation of new ecotypes was developed through natural selection best adapted to the changeability of the environment.

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65

Ice Age, as stated by Kostina [2], played a significant part in the formation of individual apricot varieties mainly on the edges of apricot-growing areas. In the North, frost-resistant varieties were formed with shortest vegetation period, that is, the xerophytic dwarf variety of *Armeniaca sibirica* and its related variety of *A. Davidiana* and a forest tree, *A. mandshurica.* Kostina further suggests that the greatest diversity of varieties and forms can be found from the middle to northern area of the overall species region of China which is represented by *A. vulgaris, A. Davidiana, A. mandshurica* as well as *A. ansu* and *A.mume* varieties. The connection of different areas of several varieties with easy breeding potential facilitates great diversity

In relation to the presence of the only variety *A. vulgaris* in the western part of China (Tian Shan), and, with regard to comparable unity of these apricot forms, we can deduce that this area is of secondary and later origin as a result of migration of Common Apricot from the primary Chinese area westward. At the time, when the climate in these mountains was not so arid and the mountains were covered in woody vegetation, they could provide a sort of

The formation and movement of apricot was probably a lengthy one and commenced with the beginning of human agricultural activities. It was then when humans started to plant first orchards in the mountains by cutting the trees around their settlements, leaving only those who provided edible fruits. This activity had a considerable influence on the diversity of wild apricot trees with juicy and sweet flesh. The earlier natural biodiversity of more abundant varieties represents even today's abundance of apricot varieties in the mountainous parts of Tian Shan and Northeastern China. Subsequent introduction of apricots as a domesticated

Man has influenced the growth of the apricot for a significant period of time. Historical records prove that as early as 4000 B.C., apricot trees accounted for widespread cultivated fruit trees in China. Later, via Central Asia, the apricot spread to Western Asia. It was only at the beginning of our century that the apricot made its way from Armenia to ancient Rome and was called the Armenian apple at the time. Simultaneously, the apricot made its way from China to Central Asia where it was possible for it to be formed autonomously by domestication of local wild apricot trees from Tian Shan. This process of transmittal into areas of wild growing apricots can be observed in many regions of Eastern Kazakhstan and Kyrgyzstan where "mountain" apricots are regarded as the most frost-resistant types and less sensitive than cultivated species [2].

species means a new era of apricot tree evolution influenced by artificial selection.

bridge between woody flora of China and the eastern part of Tian Shan.

In all countries where apricots can be grown, apricots enjoy great attention. With the change of economic factors and growing influence of globalization, further development of apricotgrowing activities is determined mainly by the three following factors:


The overall trend of world apricot production is rising, currently at approximately 2.2 million tons of apricots a year. From 1950 to 2000 the worldwide production of apricots increased four times. Some major European countries growing apricots include Spain, Italy, France, Greece, Ukraine, Moldova and others. In addition to these main producers, there are also other world producers such as Turkey, Iran, Uzbekistan, Algeria, Pakistan, and Morocco, as shown in **Figure 1** [1].

**Figure 1.** Top ten counties by apricot production.

## **2. Domestication and selection of apricots**

able to select some interesting varieties with higher adaptability to the environment (Sucre de Bohutice, Rosa Early, Holubova, Pourtáleská, Keczkemete Rozsa, Kamenickyi and so on). In all countries where apricots can be grown, apricots enjoy great attention. With the change of economic factors and growing influence of globalization, further development of apricot-

**1.** Costs are on the increase, mainly the cost of labor which is not offset with corresponding yield per hectare and price, particularly where traditional varieties of apricots are grown. **2.** Due to the spread of plum pox virus (PPV)—Sharka and European stone fruit yellows (ESFY) phytoplasma into previously unaffected areas—the yields here are lower and so is

**3.** An unresolved problem in apricot growing persists which causes early death and therefore

The overall trend of world apricot production is rising, currently at approximately 2.2 million tons of apricots a year. From 1950 to 2000 the worldwide production of apricots increased four times. Some major European countries growing apricots include Spain, Italy, France, Greece, Ukraine, Moldova and others. In addition to these main producers, there are also other world producers such as Turkey, Iran, Uzbekistan, Algeria, Pakistan, and Morocco, as shown in **Figure 1** [1].

growing activities is determined mainly by the three following factors:

the quality of fruits.

results in significant economic losses.

64 Breeding and Health Benefits of Fruit and Nut Crops

**Figure 1.** Top ten counties by apricot production.

#### **2.1. Domestication and genetic diversity of apricots**

The origin of all eco-geographic groups of apricot dates to the beginning of the Tertiary Period and is associated with the Northern and Middle China centers, which gave origin to more than 100 species of stone fruit, particularly to cherry, peach and apricot. In the Tertiary Period, apricot trees were found abundantly in extensive mountainous parts of Northern and Middle China, where, because of changing conditions, a process of forming and creation of new ecotypes was developed through natural selection best adapted to the changeability of the environment.

Ice Age, as stated by Kostina [2], played a significant part in the formation of individual apricot varieties mainly on the edges of apricot-growing areas. In the North, frost-resistant varieties were formed with shortest vegetation period, that is, the xerophytic dwarf variety of *Armeniaca sibirica* and its related variety of *A. Davidiana* and a forest tree, *A. mandshurica.* Kostina further suggests that the greatest diversity of varieties and forms can be found from the middle to northern area of the overall species region of China which is represented by *A. vulgaris, A. Davidiana, A. mandshurica* as well as *A. ansu* and *A.mume* varieties. The connection of different areas of several varieties with easy breeding potential facilitates great diversity among both domesticated and wild apricot trees.

In relation to the presence of the only variety *A. vulgaris* in the western part of China (Tian Shan), and, with regard to comparable unity of these apricot forms, we can deduce that this area is of secondary and later origin as a result of migration of Common Apricot from the primary Chinese area westward. At the time, when the climate in these mountains was not so arid and the mountains were covered in woody vegetation, they could provide a sort of bridge between woody flora of China and the eastern part of Tian Shan.

The formation and movement of apricot was probably a lengthy one and commenced with the beginning of human agricultural activities. It was then when humans started to plant first orchards in the mountains by cutting the trees around their settlements, leaving only those who provided edible fruits. This activity had a considerable influence on the diversity of wild apricot trees with juicy and sweet flesh. The earlier natural biodiversity of more abundant varieties represents even today's abundance of apricot varieties in the mountainous parts of Tian Shan and Northeastern China. Subsequent introduction of apricots as a domesticated species means a new era of apricot tree evolution influenced by artificial selection.

Man has influenced the growth of the apricot for a significant period of time. Historical records prove that as early as 4000 B.C., apricot trees accounted for widespread cultivated fruit trees in China. Later, via Central Asia, the apricot spread to Western Asia. It was only at the beginning of our century that the apricot made its way from Armenia to ancient Rome and was called the Armenian apple at the time. Simultaneously, the apricot made its way from China to Central Asia where it was possible for it to be formed autonomously by domestication of local wild apricot trees from Tian Shan. This process of transmittal into areas of wild growing apricots can be observed in many regions of Eastern Kazakhstan and Kyrgyzstan where "mountain" apricots are regarded as the most frost-resistant types and less sensitive than cultivated species [2].

Cultivated types of the basic *A.vulgaris* variety went through significant changes and took on various characters when moving west and south from China to Central Asia. This happened because of many evolutionary factors (external conditions, environment, changeability of inheritance and natural or artificial selection). As the role of artificial selection was of most important significance, these changes were reflected mainly in the quality of fruits (size and taste) but also in some economic traits such as frost resistance, immunity to main fungal diseases and biology of propagation.

The simplicity of generative propagation and a commonly known method of vegetative apricot propagation have played a key role in the intensity of selection and vegetative propagation of most economically valuable differences. The comparison of the current cultivated range of apricot with wild varieties in Western and Central Tian Shan provides evidence of gradually acquired biodiversity of traits in apricots. Primary differences between these two groups are based on the size of the fruit. Wild grown apricots in Tian Shan weigh from 3.0 to 35 g (an average of 8–12 g), domesticated varieties of Central Asia from 5.5 to 55 g (an average of 15–30 g) and Irano-Caucasian and European domesticated varieties weigh from 10 to 165 g (an average of 30–55 g). Other traits include sugar and acidity contents, taste, kernel taste, skin pubescence and stone size [2].

The evolution of cultivated apricots in Europe took on a slightly different direction because of its shorter history in this region. The apricot first arrived from Iran to Ancient Greece and Rome and to southern Europe. A more or less substantial spread of the apricot in Europe was not achieved until the seventeenth century. At the same time, a brief period of growing and limited original material domination of vegetative propagation and a very low degree of seed propagation in the apricot have all led to a lot more limited diversity in European apricots. Direct consumption of apricot fruits initiated the selection and vegetative propagation of those varieties which had been formed as random seedlings in orchards and nurseries. Basic characteristics that presented value in introducing apricots as fruits used in their fresh form were the size of fruits, a comparable low stone-to-flesh ratio, excellent taste, harmonious constitution of sugars and acids, aroma and flesh firmness.

took place in two stages. Apricots were known in Italy and Greece as a result of the Roman-Persian wars in the first century BC. The species of *Armeniaca* suggests that apricots were first brought to Italy and Greece by Armenian traders. This happened much later when the apricot

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67

Because of lack of biological material, the specification of local varieties in North Africa and Spain is not highlighted. Here, apricots were brought mainly by the Arabs from Syria and they kept their Syrian names (mesh mesh and mush mush). For the character of its fruits and biological properties, this group can be assigned to the Irano-Caucasian ecological geographic group. However, these were classified as a North African group adapted to warmer climate [6].

The process of domestication in Mediterranean species results in a loss of diversity which is far greater in fruit species introduced into Mediterranean areas compared to the species native to this region (olive and grape). By comparing genetic diversity among regional apricot gene pools in several Mediterranean areas, the loss of genetic diversity associated with apricot selection and diffusion into the Mediterranean Basin was investigated [7]. Microsatellite markers were able to detect a marked domestication bottleneck in the Mediterranean apricot material. This led to the depiction of a global image of two diffusion routes from the "Irano-Caucasian" gene pool: North Mediterranean and Southwest Mediterranean. It also assessed a significant loss of genetic diversity from the "Irano-Caucasian" gene pool, considered as a secondary center of diversification, to the Northern and Southwestern Mediterranean Basin. A substantial proportion of shared alleles were specifically detected when comparing gene pools from the "North Mediterranean Basin" and "South Mediterranean Basin" to the secondary center of diversification. Based on the three main identified gene pools, we observed a significant and substantial loss of apricot genetic diversity, ranging from about 37 to 49%

was already grown in other parts of Southern Europe [5].

**Picture 1.** The mountains and valleys of Western Pamir (photo by Saodatkadamova T.M.).

Vavilov highlights the importance of mountains when clarifying the variety of cultivated plants [3, 4]. Vavilov states that apricots are grown in their three centers of origin. Key areas of these centers are Central China (Gansu Province, mountain areas of Northeastern and Central China and Northeastern Tibet) and Central Asia (mountain areas stretching from Northwestern Iran, the Caucasus Region to Central Turkey). Vavilov found that the center of Near East is a secondary center for apricots in terms of originally grown varieties. As these findings on the origin and evolution of growing apricots outline, the importance of mountains when clarifying biodiversity of apricot species and varieties is unquestionable. The connection of wild species and forms, old and formerly grown varieties with mountains is also confirmed when studying the terrain of regions in Central Asia [3] (**Picture 1**).

The apricot spread to Europe from Central Asia, through Iran and into the trans-Caucasus Region and then further West. This movement was conditional to war campaigns and economic and cultural exchange during the arrival of Alexander the Great in the area from Turkistan to the Fergana Valley in the fourth century BC.Another shift of the apricot westward

**Picture 1.** The mountains and valleys of Western Pamir (photo by Saodatkadamova T.M.).

Cultivated types of the basic *A.vulgaris* variety went through significant changes and took on various characters when moving west and south from China to Central Asia. This happened because of many evolutionary factors (external conditions, environment, changeability of inheritance and natural or artificial selection). As the role of artificial selection was of most important significance, these changes were reflected mainly in the quality of fruits (size and taste) but also in some economic traits such as frost resistance, immunity to main fungal dis-

The simplicity of generative propagation and a commonly known method of vegetative apricot propagation have played a key role in the intensity of selection and vegetative propagation of most economically valuable differences. The comparison of the current cultivated range of apricot with wild varieties in Western and Central Tian Shan provides evidence of gradually acquired biodiversity of traits in apricots. Primary differences between these two groups are based on the size of the fruit. Wild grown apricots in Tian Shan weigh from 3.0 to 35 g (an average of 8–12 g), domesticated varieties of Central Asia from 5.5 to 55 g (an average of 15–30 g) and Irano-Caucasian and European domesticated varieties weigh from 10 to 165 g (an average of 30–55 g). Other traits include sugar and acidity contents, taste, kernel taste, skin

The evolution of cultivated apricots in Europe took on a slightly different direction because of its shorter history in this region. The apricot first arrived from Iran to Ancient Greece and Rome and to southern Europe. A more or less substantial spread of the apricot in Europe was not achieved until the seventeenth century. At the same time, a brief period of growing and limited original material domination of vegetative propagation and a very low degree of seed propagation in the apricot have all led to a lot more limited diversity in European apricots. Direct consumption of apricot fruits initiated the selection and vegetative propagation of those varieties which had been formed as random seedlings in orchards and nurseries. Basic characteristics that presented value in introducing apricots as fruits used in their fresh form were the size of fruits, a comparable low stone-to-flesh ratio, excellent taste, harmonious

Vavilov highlights the importance of mountains when clarifying the variety of cultivated plants [3, 4]. Vavilov states that apricots are grown in their three centers of origin. Key areas of these centers are Central China (Gansu Province, mountain areas of Northeastern and Central China and Northeastern Tibet) and Central Asia (mountain areas stretching from Northwestern Iran, the Caucasus Region to Central Turkey). Vavilov found that the center of Near East is a secondary center for apricots in terms of originally grown varieties. As these findings on the origin and evolution of growing apricots outline, the importance of mountains when clarifying biodiversity of apricot species and varieties is unquestionable. The connection of wild species and forms, old and formerly grown varieties with mountains is also confirmed

The apricot spread to Europe from Central Asia, through Iran and into the trans-Caucasus Region and then further West. This movement was conditional to war campaigns and economic and cultural exchange during the arrival of Alexander the Great in the area from Turkistan to the Fergana Valley in the fourth century BC.Another shift of the apricot westward

eases and biology of propagation.

66 Breeding and Health Benefits of Fruit and Nut Crops

pubescence and stone size [2].

constitution of sugars and acids, aroma and flesh firmness.

when studying the terrain of regions in Central Asia [3] (**Picture 1**).

took place in two stages. Apricots were known in Italy and Greece as a result of the Roman-Persian wars in the first century BC. The species of *Armeniaca* suggests that apricots were first brought to Italy and Greece by Armenian traders. This happened much later when the apricot was already grown in other parts of Southern Europe [5].

Because of lack of biological material, the specification of local varieties in North Africa and Spain is not highlighted. Here, apricots were brought mainly by the Arabs from Syria and they kept their Syrian names (mesh mesh and mush mush). For the character of its fruits and biological properties, this group can be assigned to the Irano-Caucasian ecological geographic group. However, these were classified as a North African group adapted to warmer climate [6].

The process of domestication in Mediterranean species results in a loss of diversity which is far greater in fruit species introduced into Mediterranean areas compared to the species native to this region (olive and grape). By comparing genetic diversity among regional apricot gene pools in several Mediterranean areas, the loss of genetic diversity associated with apricot selection and diffusion into the Mediterranean Basin was investigated [7]. Microsatellite markers were able to detect a marked domestication bottleneck in the Mediterranean apricot material. This led to the depiction of a global image of two diffusion routes from the "Irano-Caucasian" gene pool: North Mediterranean and Southwest Mediterranean. It also assessed a significant loss of genetic diversity from the "Irano-Caucasian" gene pool, considered as a secondary center of diversification, to the Northern and Southwestern Mediterranean Basin. A substantial proportion of shared alleles were specifically detected when comparing gene pools from the "North Mediterranean Basin" and "South Mediterranean Basin" to the secondary center of diversification. Based on the three main identified gene pools, we observed a significant and substantial loss of apricot genetic diversity, ranging from about 37 to 49% from the secondary apricot diversification zone ("Irano-Caucasian") to the Southwestern Mediterranean Basin, depicting a genetic signature of apricot domestication and diffusion into the Mediterranean Basin. Unlike Kostina's assumptions, we proposed an evolutionary scenario in favor of two diffusion routes in Southern Europe and North Africa as revealed by a substantial proportion of shared alleles that were specifically detected along each of the two diffusion routes [8]. This study generated genetic insight that will be useful for management and conservation of Mediterranean apricot germ-plasm as well as genetic selection and breeding programs related to adaptive traits [7].

Michurin used the seedlings of Mongolian, Siberian and Manchurian apricots as donors for frost hardiness and increased adaptability. Abrikos No. 84, No. 86, No 241, No 246, Mongol, Sacer were utilized as Mongolian apricot seedlings and Lučšij Mičurinskiy and Tovarišč as the seedlings of *P. sibirica*. To increase frost hardiness, it is recommended to breed apricot varieties with varieties that are native to countries further away from the center of cultivated

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To increase the frost hardiness of flower buds, breeding apricots with *P. salicina* L. while creating several fertile types with transitional traits called "Plumcot" have been suggested. It has been highlighted that the Royal variety can be grown in many regions where other apricot varieties do not bear fruit and may prove to be a valuable material for further selection [10]. Hummel created a list of *Prunus* varieties which are known as frost hardy or grown with minimal damage in zones classified according to frost hardiness from 4b (from −30 to −20°F, which is −34.4 to −28.9°C) to zone 1 (below −50°F, below −45.6°C). For zone 4a varieties such as Anda, Harbin and Labin—all seedlings of *P. mandshurica* L.— and then Manchu and Moongold,

The basic requirement for breeding, selection and further introduction of new varieties is knowledge of a wide collection of genotypes, varieties, clones and landraces of all ecological groups of *P. armeniaca* L. This enables breeders to select donors of individual properties which they expect to appear in the progeny's genotype. At Horticulture Faculty in Lednice, studies have been carried out since 1985 on several apricot collections of genotypes, and several years of observations have helped in choosing the most important characters connected with adaptability to environment [12]. The analysis of these characters enables breeders to choose parents—donors of characters and conservation of genetic characters. Genetic improvement of apricots as a species is possible in all characters relating to increase in adaptability. However, it usually takes a longer period of time. The following donors of characters have been chosen.

In different years and in different collections, the frequency of genotypes having a high level of frost hardiness of flower buds varied from 8.6% to almost 18% of all observed apricot progeny. Genotypes having fruits presenting a good market value and a high frost hardiness had a frequency of occurrence varying from 2 to 48% (**Table 1**). Some of the frost-hardy cultivars are Harlayne, Harval, Leala, Lejuna, Leronda, Leskora, Lenova, Lejara, LE-498, LE-806, NJA 1, Vivagold, Volschebnyi, Vynoslivyi, Yulskyi, Strepet, Harrow Star, NJA 35, NJA 62, NJA 77, NJA 44, Veharda, Vegama, Harcot, Saldcot, Vyndrop, Alfred, Reliable, apricot seedlings of the serious M-LE-1, M-VA-3, M-VA-2, M-VA-1, Dzhankojskiy rannyi, Arzami,Henderson, Morden,Senetate, VS 023/187, Riland, Veecot, Vestar, Lunnik, Harglow, Stark Early Orange,

A higher frequency of phenotypes (16%) was observed with late ending of dormancy character, globally. From the breeding and practical points of view, genotypes presenting a combination

varieties, such as those of Asia Minor, Palestine and Persia [9].

Morden 604, Morden 601 and Scout and others have been chosen [11].

*2.2.1. Frost hardiness of flower buds*

Orange Red, Scout and Lel.

*2.2.2. Late termination of dormancy*

#### **2.2. Selection of donors of characters related to breeding aims**

Breeding programs of all countries with economically significant apricot production are based on the effort of obtaining varieties with wide or varied ecological adaptability regarding their production areas. The efforts are to create varieties that are adaptable to low temperatures, mainly to sudden temperature changes during the post-dormancy period, but also to meet requirements for a small number of chilling units in winter. Since the apricot requires specified eco-climatic conditions as outlined by its phylogenesis, it lacks the ability for wide environmental adaptability as opposed to, for example, the apple variety Golden Delicious or the peach variety Redhaven. The change in the traditional range of apricots is also caused by globalization, methods of sale, requirements of vendors and the fact that in the European market apricots are of increasingly higher importance as a fresh commodity.

Successful introduction of new apricot varieties should focus on main targets which are similar in most growing periods. Generally, it is possible to summarize breeding aims into several objectives:


It is possible to say that all the aforementioned breeding aims are of fundamental importance and are pursued by a majority of breeders worldwide. Complimentary breeding objectives include growth control, so called dwarfism or semi-dwarfism, the compatibility of varieties with diverse rootstock, crown habitat and its suitability for different modern breeding shapes and resistance to drought.

Michurin used the seedlings of Mongolian, Siberian and Manchurian apricots as donors for frost hardiness and increased adaptability. Abrikos No. 84, No. 86, No 241, No 246, Mongol, Sacer were utilized as Mongolian apricot seedlings and Lučšij Mičurinskiy and Tovarišč as the seedlings of *P. sibirica*. To increase frost hardiness, it is recommended to breed apricot varieties with varieties that are native to countries further away from the center of cultivated varieties, such as those of Asia Minor, Palestine and Persia [9].

To increase the frost hardiness of flower buds, breeding apricots with *P. salicina* L. while creating several fertile types with transitional traits called "Plumcot" have been suggested. It has been highlighted that the Royal variety can be grown in many regions where other apricot varieties do not bear fruit and may prove to be a valuable material for further selection [10].

Hummel created a list of *Prunus* varieties which are known as frost hardy or grown with minimal damage in zones classified according to frost hardiness from 4b (from −30 to −20°F, which is −34.4 to −28.9°C) to zone 1 (below −50°F, below −45.6°C). For zone 4a varieties such as Anda, Harbin and Labin—all seedlings of *P. mandshurica* L.— and then Manchu and Moongold, Morden 604, Morden 601 and Scout and others have been chosen [11].

The basic requirement for breeding, selection and further introduction of new varieties is knowledge of a wide collection of genotypes, varieties, clones and landraces of all ecological groups of *P. armeniaca* L. This enables breeders to select donors of individual properties which they expect to appear in the progeny's genotype. At Horticulture Faculty in Lednice, studies have been carried out since 1985 on several apricot collections of genotypes, and several years of observations have helped in choosing the most important characters connected with adaptability to environment [12]. The analysis of these characters enables breeders to choose parents—donors of characters and conservation of genetic characters. Genetic improvement of apricots as a species is possible in all characters relating to increase in adaptability. However, it usually takes a longer period of time. The following donors of characters have been chosen.

#### *2.2.1. Frost hardiness of flower buds*

from the secondary apricot diversification zone ("Irano-Caucasian") to the Southwestern Mediterranean Basin, depicting a genetic signature of apricot domestication and diffusion into the Mediterranean Basin. Unlike Kostina's assumptions, we proposed an evolutionary scenario in favor of two diffusion routes in Southern Europe and North Africa as revealed by a substantial proportion of shared alleles that were specifically detected along each of the two diffusion routes [8]. This study generated genetic insight that will be useful for management and conservation of Mediterranean apricot germ-plasm as well as genetic selection and breed-

Breeding programs of all countries with economically significant apricot production are based on the effort of obtaining varieties with wide or varied ecological adaptability regarding their production areas. The efforts are to create varieties that are adaptable to low temperatures, mainly to sudden temperature changes during the post-dormancy period, but also to meet requirements for a small number of chilling units in winter. Since the apricot requires specified eco-climatic conditions as outlined by its phylogenesis, it lacks the ability for wide environmental adaptability as opposed to, for example, the apple variety Golden Delicious or the peach variety Redhaven. The change in the traditional range of apricots is also caused by globalization, methods of sale, requirements of vendors and the fact that in the European

Successful introduction of new apricot varieties should focus on main targets which are similar in most growing periods. Generally, it is possible to summarize breeding aims into several

• Adaptability to both cold and warm areas: To achieve an appropriate level of adaptability, it is important to assure a good degree of frost resistance. Furthermore, it is a requirement for a small or large number of chilling units, long or short dormancy periods, slow development of pollen microsporogenesis during the post-dormancy period, late blooming period,

• Ideotype of fruits: For their utilization, it is important fruits comply with certain criteria, that is, the size and firmness of fruits, attractive appearance, taste, aroma, sugar and acid

• Resistance to diseases and early death: In Europe, for example, the main objective is breeding apricots resistant to Sharka—PPV, European stone fruit yellows (ESFY), brown rot, *Cytospora* and bacteria such as *Pseudonamas* spp. and *Xantonomas* spp. and early decline as

It is possible to say that all the aforementioned breeding aims are of fundamental importance and are pursued by a majority of breeders worldwide. Complimentary breeding objectives include growth control, so called dwarfism or semi-dwarfism, the compatibility of varieties with diverse rootstock, crown habitat and its suitability for different modern breeding shapes

ing programs related to adaptive traits [7].

68 Breeding and Health Benefits of Fruit and Nut Crops

objectives:

contents.

and resistance to drought.

frost hardiness and autogamy.

a result of pathological factors and environment.

**2.2. Selection of donors of characters related to breeding aims**

market apricots are of increasingly higher importance as a fresh commodity.

In different years and in different collections, the frequency of genotypes having a high level of frost hardiness of flower buds varied from 8.6% to almost 18% of all observed apricot progeny. Genotypes having fruits presenting a good market value and a high frost hardiness had a frequency of occurrence varying from 2 to 48% (**Table 1**). Some of the frost-hardy cultivars are Harlayne, Harval, Leala, Lejuna, Leronda, Leskora, Lenova, Lejara, LE-498, LE-806, NJA 1, Vivagold, Volschebnyi, Vynoslivyi, Yulskyi, Strepet, Harrow Star, NJA 35, NJA 62, NJA 77, NJA 44, Veharda, Vegama, Harcot, Saldcot, Vyndrop, Alfred, Reliable, apricot seedlings of the serious M-LE-1, M-VA-3, M-VA-2, M-VA-1, Dzhankojskiy rannyi, Arzami,Henderson, Morden,Senetate, VS 023/187, Riland, Veecot, Vestar, Lunnik, Harglow, Stark Early Orange, Orange Red, Scout and Lel.

#### *2.2.2. Late termination of dormancy*

A higher frequency of phenotypes (16%) was observed with late ending of dormancy character, globally. From the breeding and practical points of view, genotypes presenting a combination


*2.2.4. Late blooming*

and Harglow).

Gergana.

yellows.

*2.2.5. High level of self-fertility*

*2.2.6. Climatic adaptation*

*2.2.9. Attractiveness of fruits*

attractive varieties.

On the contrary, the character of late blooming was observed in the lowest number of phenotypes (0.061). The frequency of individuals presenting both a late blooming and a good market value of fruit appeared to be very rare, only about 0.8% of all evaluated genotypes of apricots (Early Gold, Machova, Marculesti, *P. brigantiaca* L. xOlymp, Re Umberto, Sulmona, Stella, Pozdněkvetoucí, Frostina, Farclo, Fardao, Dolgocvetna, Polyus yuznyi, Zard, Oranzevokrasnyi, Venus, Tilton, Selena, Badami, Kamenický, Rosa Late, Yulskyi, Ambrosia, Farbaly

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High frequency of the requested values of characters was achieved on self-fertility. Of the overall number of self-fertile genotypes, 25% showed significant pomological traits. Examples of such cultivars are Bergeron, Minaret, Vestar, Kostinskyi,Leala, Marlen, Pisana, Kioto and

Bergeron, Goldrich, Marculesti, Tilton, Kecskemete Rose, Leala, Lejuna, Leskora, Re Umberto, Rose Early, Vynosliviy, Bergarouge, Harrow Star, Harlayne, Tomcot, Bronzoviy and Kostinskyi show climatic adaptation. Harglow, NS-4 and Strepet also had a higher level of adaptability in the European eco-geographical groups of apricots. It includes some types bearing small fruits which come from a European subgroup as well as species from the groups Rosa Early, Holubova, Pourtal abricose, Sucre de Bohutice, Keczkemete Rozsa,

Vestar and Royer were the only two cultivars which were tolerant to European stone fruit

Stark Early Orange, Harlayne, Henderson, Orangered, Betinka, Adriana, Candela, Sophia, Veecot, Leronda,Harval, Sundrop and Harcot are some of the varieties resistant to PPV.

Harrow Joy, Bergarouge, Betinka, Orangered, Rubista, Pincot, Chuan Zhi Hong, Laycot, Bobcot, Roxana, Neptun, Gergana, Veselka, Gama, Robada, Big Red, Tsunami, Carmen Top, Harrow Star, Flavor Cot, Kioto, Harrow Blush, Harogem, Magic Cot, Cegledi Piroska, Sophia, Bergeval, Sephora,Orange Rubis,Pricia,Bergeval, Swired, Gilgat, Rougemont and Montier are some of the

Bergeron, Kamenickiy and other types of natural population.

*2.2.7. Field tolerance to European stone fruit yellows (ESFY)*

*2.2.8. Tolerance or resistance to plum pox virus (PPV)*

**Table 1.** Frequency of the most important characters of apricots connected to their adaptability [12].

of both characteristics (late dormancy time and fruit quality) are very interesting, but this was the case for only 7% of analyzed progeny (Vestar × Stark Early Orange). Henderson, ChuanZhi Hong, Lebela, LE-498, Oranzevo-krasnyi, Stark Early Orange, Vegama, Veharda, Zard, Vynosliviy, Reliable, NJA2 and Curtis are varieties exhibiting late termination of dormancy.

#### *2.2.3. Frost tolerance of juvenile fruits*

Almost 18% of observed genotypes showed a higher frost hardiness in young fruits than the control clone Velkopavlovická LE 12/2 /type Hungarian Best. Other genotypes having pomological characters answering market requirements were altogether 6.7% in the whole collection. For example, cultivars such as Leala, Lemira, Ledana, Leskora, Lejuna, Lefrosta, Neptun, Re Umberto, Henderson, Early Gold, Zard, Oranzevo-krasnyi, Marculešti 17/2, Horákova raná, Fara, Rakovskyi, Bergeron, Reumberto,Marlen, Detskyi, M 146, Neptun, Triumf severa, Patriarca tempráno, M 90B, Baneasa 16/3, Kamenickýi, Selena, Mari de Cenad, NJA 33, Morden 604, Sephora, Bergeval, Big Red, Orange Rubis, Anegat, Primaya and Orangered had juvenile fruits with frost tolerance.

#### *2.2.4. Late blooming*

On the contrary, the character of late blooming was observed in the lowest number of phenotypes (0.061). The frequency of individuals presenting both a late blooming and a good market value of fruit appeared to be very rare, only about 0.8% of all evaluated genotypes of apricots (Early Gold, Machova, Marculesti, *P. brigantiaca* L. xOlymp, Re Umberto, Sulmona, Stella, Pozdněkvetoucí, Frostina, Farclo, Fardao, Dolgocvetna, Polyus yuznyi, Zard, Oranzevokrasnyi, Venus, Tilton, Selena, Badami, Kamenický, Rosa Late, Yulskyi, Ambrosia, Farbaly and Harglow).

#### *2.2.5. High level of self-fertility*

High frequency of the requested values of characters was achieved on self-fertility. Of the overall number of self-fertile genotypes, 25% showed significant pomological traits. Examples of such cultivars are Bergeron, Minaret, Vestar, Kostinskyi,Leala, Marlen, Pisana, Kioto and Gergana.

#### *2.2.6. Climatic adaptation*

of both characteristics (late dormancy time and fruit quality) are very interesting, but this was the case for only 7% of analyzed progeny (Vestar × Stark Early Orange). Henderson, ChuanZhi Hong, Lebela, LE-498, Oranzevo-krasnyi, Stark Early Orange, Vegama, Veharda, Zard, Vynosliviy, Reliable, NJA2 and Curtis are varieties exhibiting late termination of dormancy.

**Characters Categories No. of** 

**Frost hardiness of flower buds 346 1**

fruits

**Termination of bud dormancy 82 1**

of fruits

**Blooming time 489 1**

**Self-fertility 124 1**

fruits

**Table 1.** Frequency of the most important characters of apricots connected to their adaptability [12].

Hardiness + market value of

More tolerant + market value

Self-fertile + market value of

**individuals**

56 0.042

**209 1**

14 0.066

31 0.250

High hardiness 182 0.135

Late 13 0.159 Late + market value of fruits 6 0.073

Than control 37 0.177

Late 29 0.059 Late + market-value of fruits 13 0.030

Self-fertile 44 0.354

**Frequency**

Almost 18% of observed genotypes showed a higher frost hardiness in young fruits than the control clone Velkopavlovická LE 12/2 /type Hungarian Best. Other genotypes having pomological characters answering market requirements were altogether 6.7% in the whole collection. For example, cultivars such as Leala, Lemira, Ledana, Leskora, Lejuna, Lefrosta, Neptun, Re Umberto, Henderson, Early Gold, Zard, Oranzevo-krasnyi, Marculešti 17/2, Horákova raná, Fara, Rakovskyi, Bergeron, Reumberto,Marlen, Detskyi, M 146, Neptun, Triumf severa, Patriarca tempráno, M 90B, Baneasa 16/3, Kamenickýi, Selena, Mari de Cenad, NJA 33, Morden 604, Sephora, Bergeval, Big Red, Orange Rubis, Anegat, Primaya and Orangered had

*2.2.3. Frost tolerance of juvenile fruits*

**Frost tolerance of juvenile fruits more** 

70 Breeding and Health Benefits of Fruit and Nut Crops

**tolerant**

juvenile fruits with frost tolerance.

Bergeron, Goldrich, Marculesti, Tilton, Kecskemete Rose, Leala, Lejuna, Leskora, Re Umberto, Rose Early, Vynosliviy, Bergarouge, Harrow Star, Harlayne, Tomcot, Bronzoviy and Kostinskyi show climatic adaptation. Harglow, NS-4 and Strepet also had a higher level of adaptability in the European eco-geographical groups of apricots. It includes some types bearing small fruits which come from a European subgroup as well as species from the groups Rosa Early, Holubova, Pourtal abricose, Sucre de Bohutice, Keczkemete Rozsa, Bergeron, Kamenickiy and other types of natural population.

#### *2.2.7. Field tolerance to European stone fruit yellows (ESFY)*

Vestar and Royer were the only two cultivars which were tolerant to European stone fruit yellows.

#### *2.2.8. Tolerance or resistance to plum pox virus (PPV)*

Stark Early Orange, Harlayne, Henderson, Orangered, Betinka, Adriana, Candela, Sophia, Veecot, Leronda,Harval, Sundrop and Harcot are some of the varieties resistant to PPV.

#### *2.2.9. Attractiveness of fruits*

Harrow Joy, Bergarouge, Betinka, Orangered, Rubista, Pincot, Chuan Zhi Hong, Laycot, Bobcot, Roxana, Neptun, Gergana, Veselka, Gama, Robada, Big Red, Tsunami, Carmen Top, Harrow Star, Flavor Cot, Kioto, Harrow Blush, Harogem, Magic Cot, Cegledi Piroska, Sophia, Bergeval, Sephora,Orange Rubis,Pricia,Bergeval, Swired, Gilgat, Rougemont and Montier are some of the attractive varieties.

#### *2.2.10. Big fruit size*

Gergana, Roxana, China, Hargrand, Senetate, Goldrich, Olymp, Gama, Velikiy, Exnerova, Agat, Jumbo Cot, Goldstrike are some of the varieties that have big fruit sizes.

and hybrid 252 had heavier fruits than 1 gram and the percentage of frozen fruits were from 69 to 100%. This means that the size of fruits is of lesser significance to sensitivity or hardiness

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Among hardy varieties, Djurič further included Sacharijstyi. Melitopol early, Keč-pšar, RuchiDžuvanon and Zemljaničniy (frost damage to fruits varied from 40.1 to 60%) were cat-

In Italy, Bassi et al. [14] evaluated a collection of apricots at a temperature of −5.8°C when in full bloom and for the next four consecutive nights. In 1993, when apricots were losing petals, frost occurred 6 times (from 0.0 to −4.8°C). What the team in Italy found in 1990 was that some most productive varieties had more than 80% of brown pistils and the relation between the detected percentage of brown pistils and crop yield (r = 0.04) was not therefore confirmed. Varieties that had fruits mostly in the upper part of the crown were Alfred, Bella di Casale, Farmingdale and Harlayne. Other varieties such as Bergeron, Precoce di Cesena, Bulida, Canino, Goldcot and Ivonne Liverani a Mandorlou showed a relatively uniform location of fruits. Very late blooming varieties from Hungary and Romania showed a very low count of buds during all the years as well as a high percentage of brown pistils even though they had bloomed after a period of frost. This defect is caused by bad adaptability of varieties from

Concerning the damage to the apricot fruits during spring frost at Mendel University in Brno,

**i.** The genotypes Leala, Lemira, Ledana, Leskora, Lejuna, Lefrosta, Henderson, Early Gold, Zard, Oranžovokrasniy, Marculešti 17/2, Horákova raná, Fara, Rakovského, Reumberto, Lednická, Detskij, Neptun, Triumf severa, Patriarca tempráno, Baneasa 16/3, Kamenický, Selena, Mari de Cenad and NJA 33 conclusively or highly conclusively increased tolerance of fruits to frost than the control variety (Velkopavlovická LE-19/2,) in 2, pos-

**ii.** The genotypes Orange Red, Legolda, Shalah, Nugget, Kecskemetr rozsa, Leronda, Skaha

It was possible to detect a highly conclusive impact of genotype, year and blooming period during these years of observation when the conditions differed each time either because of vegetation development or because of critical temperatures. The impact of blooming period was recorded as highly conclusive in relation to the percentage of frozen fruits (r = 0.41++). In all years, the fruits of late blooming varieties sustained less damage (Reumberto, Venus, Zard, Marculesti 17/2, Neptun and Leala). However, there were certain genotypes which, despite their early or mid-early blooming period, were not significantly damaged (Bukurija, Junskij,

Breeders also face another challenge, which is to single out the most harmful diseases and pests relevant to a particular area, recognize their biology and inheritance and explore the mechanism of resistance. They would then be able to, based on such selection, use disease-resistance donors in breeding programs with quality donors. When growing plants, resistance breeding is the most efficient, natural and widespread method of protection against pathogens [16].

Horticulture Faculty in Lednice, from 1989 to 1996, the following was observed:

than genetic foundation of a variety character.

Central to Eastern Europe which demands cooler conditions [14].

and Lerosa showed lower resistance of fruits to frost.

egorized as semi-sensitive varieties [13].

sibly 3 years.

Senetate and Henderson) [15].

#### *2.2.11. High sugar content*

Shalach, Sekerpare, Forum, Lakaniy, Strepet, Nadezda, Olymp, Kabaasi, Hacihaliloglu, Bronzoviy, Abutalibi, Vynosliviy, Hasanbey, Suphani and Isfarah are some of the varieties that have high sugar content.

#### *2.2.12. Excellent taste and apricot aroma*

Velkopavlovická, Sucre de Bohutice, Sabinovská, Hungarian Best, Klosterneuburg, Krasnoscokiy, Bronzoviy, Vynosliviy, Bobcot, Harrow Joy, Bergarouge, Hargrand, Skopljanska krupnoplodna, Royer, Paviot, Sucre de Holub, Nancy apricot, Luizet, Polonais, Betinka, Bergeval, Breda and Paviot are some of the varieties that have an excellent taste and aroma.

#### *2.2.13. Extension of the ripening time*

In relation to late ripening, many potentially interesting parents are also available. Pisana Bergeron, Tardif de Bordaneil, Tardicot, Anegat, Farbaly, Farclo, Fardao, Kechpsar,Vynoslivyi, Helena de Roussilon, Rosa Late, Keczkemetr rozsa and Borsi-félekései rózsa are some of them.

In relation to early ripening, many potentially interesting parents are also available such as Tomcot, Magic Cot, Wonder Cot, Pricia, Spring Blush, Big Red, Bukuriya, Early Samarkand, Tsunami and Banzai [11].

All the evaluated traits exhibited a different value of variability and frequency enabling the breeders and geneticists to make more informed choices on the selection of donors for a particular character. This variability allows the integration and combination of each single character in the breeding process which will lead to the production of hybrids with a higher level of adaptability and fruit market value [12].

Increased adaptability to environment is also essential for the selection of genotypes with frost-tolerant fruits. Late spring frost which damages small developing fruits after the blooming period is more common in the Mediterranean, but from time to time it can also occur in the Central European climate. As the climate changes (milder winters, early onset of spring), spring frost after the blooming period is more frequent.

A degree of hardiness in fruits of individual varieties is of the greatest difference as conditioned by the development phase and the length and time of the critical period.

Different authors suggest that the temperatures from −0.5 to −1.6°C are dangerous for apricots after the blooming period. Djurič found no relation between the development (size) of fruits and their frost hardiness. He found that varieties with the smallest percentage of frozen fruits (Hindukush, Overnskyi, Zimostojki and Novosadski clone BC-1) had fruits of lower weight than 1 gram; however, varieties Blenril, Nugget, Royal, SEO, Kostjuženskij and NJ 27, NJ 26 and hybrid 252 had heavier fruits than 1 gram and the percentage of frozen fruits were from 69 to 100%. This means that the size of fruits is of lesser significance to sensitivity or hardiness than genetic foundation of a variety character.

*2.2.10. Big fruit size*

*2.2.11. High sugar content*

that have high sugar content.

*2.2.12. Excellent taste and apricot aroma*

72 Breeding and Health Benefits of Fruit and Nut Crops

*2.2.13. Extension of the ripening time*

level of adaptability and fruit market value [12].

spring frost after the blooming period is more frequent.

Tsunami and Banzai [11].

Gergana, Roxana, China, Hargrand, Senetate, Goldrich, Olymp, Gama, Velikiy, Exnerova,

Shalach, Sekerpare, Forum, Lakaniy, Strepet, Nadezda, Olymp, Kabaasi, Hacihaliloglu, Bronzoviy, Abutalibi, Vynosliviy, Hasanbey, Suphani and Isfarah are some of the varieties

Velkopavlovická, Sucre de Bohutice, Sabinovská, Hungarian Best, Klosterneuburg, Krasnoscokiy, Bronzoviy, Vynosliviy, Bobcot, Harrow Joy, Bergarouge, Hargrand, Skopljanska krupnoplodna, Royer, Paviot, Sucre de Holub, Nancy apricot, Luizet, Polonais, Betinka, Bergeval, Breda and Paviot are some of the varieties that have an excellent taste and aroma.

In relation to late ripening, many potentially interesting parents are also available. Pisana Bergeron, Tardif de Bordaneil, Tardicot, Anegat, Farbaly, Farclo, Fardao, Kechpsar,Vynoslivyi, Helena de Roussilon, Rosa Late, Keczkemetr rozsa and Borsi-félekései rózsa are some of them. In relation to early ripening, many potentially interesting parents are also available such as Tomcot, Magic Cot, Wonder Cot, Pricia, Spring Blush, Big Red, Bukuriya, Early Samarkand,

All the evaluated traits exhibited a different value of variability and frequency enabling the breeders and geneticists to make more informed choices on the selection of donors for a particular character. This variability allows the integration and combination of each single character in the breeding process which will lead to the production of hybrids with a higher

Increased adaptability to environment is also essential for the selection of genotypes with frost-tolerant fruits. Late spring frost which damages small developing fruits after the blooming period is more common in the Mediterranean, but from time to time it can also occur in the Central European climate. As the climate changes (milder winters, early onset of spring),

A degree of hardiness in fruits of individual varieties is of the greatest difference as condi-

Different authors suggest that the temperatures from −0.5 to −1.6°C are dangerous for apricots after the blooming period. Djurič found no relation between the development (size) of fruits and their frost hardiness. He found that varieties with the smallest percentage of frozen fruits (Hindukush, Overnskyi, Zimostojki and Novosadski clone BC-1) had fruits of lower weight than 1 gram; however, varieties Blenril, Nugget, Royal, SEO, Kostjuženskij and NJ 27, NJ 26

tioned by the development phase and the length and time of the critical period.

Agat, Jumbo Cot, Goldstrike are some of the varieties that have big fruit sizes.

Among hardy varieties, Djurič further included Sacharijstyi. Melitopol early, Keč-pšar, RuchiDžuvanon and Zemljaničniy (frost damage to fruits varied from 40.1 to 60%) were categorized as semi-sensitive varieties [13].

In Italy, Bassi et al. [14] evaluated a collection of apricots at a temperature of −5.8°C when in full bloom and for the next four consecutive nights. In 1993, when apricots were losing petals, frost occurred 6 times (from 0.0 to −4.8°C). What the team in Italy found in 1990 was that some most productive varieties had more than 80% of brown pistils and the relation between the detected percentage of brown pistils and crop yield (r = 0.04) was not therefore confirmed.

Varieties that had fruits mostly in the upper part of the crown were Alfred, Bella di Casale, Farmingdale and Harlayne. Other varieties such as Bergeron, Precoce di Cesena, Bulida, Canino, Goldcot and Ivonne Liverani a Mandorlou showed a relatively uniform location of fruits. Very late blooming varieties from Hungary and Romania showed a very low count of buds during all the years as well as a high percentage of brown pistils even though they had bloomed after a period of frost. This defect is caused by bad adaptability of varieties from Central to Eastern Europe which demands cooler conditions [14].

Concerning the damage to the apricot fruits during spring frost at Mendel University in Brno, Horticulture Faculty in Lednice, from 1989 to 1996, the following was observed:


It was possible to detect a highly conclusive impact of genotype, year and blooming period during these years of observation when the conditions differed each time either because of vegetation development or because of critical temperatures. The impact of blooming period was recorded as highly conclusive in relation to the percentage of frozen fruits (r = 0.41++). In all years, the fruits of late blooming varieties sustained less damage (Reumberto, Venus, Zard, Marculesti 17/2, Neptun and Leala). However, there were certain genotypes which, despite their early or mid-early blooming period, were not significantly damaged (Bukurija, Junskij, Senetate and Henderson) [15].

Breeders also face another challenge, which is to single out the most harmful diseases and pests relevant to a particular area, recognize their biology and inheritance and explore the mechanism of resistance. They would then be able to, based on such selection, use disease-resistance donors in breeding programs with quality donors. When growing plants, resistance breeding is the most efficient, natural and widespread method of protection against pathogens [16].

Greek colleagues have worked very hard studying apricot varieties mainly in a stage after a strong natural infection. Syrgiannidis was first to describe the Stark Early Orange and Stella cultivars as resistant against *plum pox virus* based on his fieldwork between the years of 1967– 1970. He then sorted the varieties into very sensitive (ProimoTyrinthos, Rouge de Sernhac), quite sensitive (Blenril, Canino, Docteur Mascle, Bergeron, Moorpark, Nugget, Rouge de Fournes, Sungold and Tilton a Stavropoulos) and less sensitive (Blenheim Royal, Ricordo di Amic and Grossa del Giardino) [17].

(its intensity and length) ensures less dependence on external environmental conditions and is a crucial factor for frost hardiness. Based on its origin, the apricot is a mountainous plant and as such is typical for its adaptability to cold winter without fluctuating temperatures. Results of many authors have shown that buds of stone fruit experience important stages of

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Development of blossom buds at the time of dormancy is assured by individual changes and intensity of metabolism. Jablonskij and Elmanova found a relation between accumulated hydrocarbons and the pace of morphogenesis as well as a relation between morphological and anatomical changes and the dynamics of phenol contents in buds (r = 0,83). This occurrence relates to the fact that most phenols inhibit the growing process. They found that the highest contents of phenols in blossom buds were present (as detected in the beginning of

Kostina et al. compared the inheritance of dormancy length in reciprocal crossing of varieties Zard and Shalah, which belong to two different eco-geographical groups with different dormancy lengths. In this cross, 34.8% of progeny inherited the character of slow development of

The study on two progenies, where the late terminating dormancy variety Stark Early Orange (SEO) was the male variety for both of them and female varieties from early to mid-earlybreaking dormancy of Vestar and Velkopavlovická LE-19/2 were utilized, observed that in both the combinations 82.1% of hybrids terminated their deep vegetative dormancy after their parents and intermediates. New traits appeared in 17.9% of progeny, of which being positive

In her breeding work, Kostina established a dominant inheritance character in apricots from Central Asia regarding several biological traits including late blooming and increased frost hardiness if apricots in this group had been crossed with apricots in the Irano-Caucasian group or European group [8]. Crossa-Raynaud (when crossing Canino and Amerleuch showed the intermediate inheritance character of development rhythm in flower buds of blooming period [26]. Zagorodnaya also found that more than a half of observed progeny possessed frosthardiness properties of blossom buds among parents and quite a considerable number of progeny were drawing nearer to their frost-hardy parent, particularly if this parent was used

Oranževokrasniy is also recommended as a frost-hardy donor for crossing by Kostina together with Churmai variety (long dormancy period and late blooming) [24]. Hough also recommended (written statement, 1989) the crossing of Oranževokrasnyi × Orange Red with

*Plum pox virus* (PPV) resistance presents one of the most discussed topics in European breeding programs. This topic has been reviewed by Zhebentyayeva et al. [29]. All apricot cultivars of European origin are susceptible to PPV. Currently, most apricot breeding programs in Europe use the PPV-resistant North American cultivars to introduce this trait into European germplasm. Resistance among apricots has been found only in some North American cultivars such as "Goldrich," "Harlayne," "Stark Early Orange," (SEO) "Stella" and "Harcot" [30, 31].

the aim of obtaining frost-hardy individuals with firm fruits and early ripening [28].

micro- and macrosporogenesis at the time of autumn, winter and spring [22].

April) in the following varieties: Narjadniy, Vynosliviy and Orfyand Amur [23].

Zard and 21.9% of progeny inherited the character of its father variety Zard [24].

(late dormancy termination) appeared in 9.4% [25].

as a mother cultivar [27].

Murg et al. [18] recorded the results of observing 64 apricot species infected with PPV in the region of Oradea and found that Stark Early Orange, Manitoba, Farmingdale, Bolšoj Rozovoj, Blenril, King, Doty, Arzami, Pize, Smyrna, Sophie, Tarzii de Bucuresti, Selena, Favorit, Timpurii de Chisinau, Rozova Scora, Rosii de Banesa, Magyar Kayszi and Borozi Rozsa varieties were not infected. Varieties which had no symptoms in fruits and weak symptoms on leaves were Venus, Tuzla, Saturn, Calatis, Dalia, Neptun, Sulina, Excelsior, Čačak, Raske Carpo, Joubert, Toulon and Precoce de Italia [18].

#### **2.3. Achieved findings in apricot breeding**

Clone selection can be regarded as an evolution of varieties when clones with higher adaptability are selected during a long-term growing period in a relatively vast area, whereas clones with lower adaptability to the environment cease to exist or are only relevant to a smaller section of areas. This is called the plasticity of a given cultivar which, influenced by its environment and length of growing, provides an array of clones with various adaptabilities. This plasticity is conditioned by the age of varieties. Hence, clone selection provides one of the solutions for increased adaptability for varieties such as Velkopavlovická, Hungarian best and Sabinovská. Clone selection can increase environmental adaptability as can the selection of additional properties other than just fertility. In the scope of several cycles of clone selection of apricots which Professor Vachůn initiated in the 60s, variability of the blooming period was also detected (a 1–2-day delay of full bloom can in some years positively influence the harvest), as was the number of blossoms with two pistils making the clones with higher number of pistils more efficient. Of other variable properties, clone selection can be used to improve the health of trees (early death or economically significant viral diseases) when even in these traits conclusive differences were found [19].

In Hungary, Nyujtó chose large fruits of local cultivars and saw the European group of apricots as being suitable enough to achieve a faster and significant improvement in cultivar adaptability using clone selection as opposed to the creation of new cultivars through crossing. His program saw the creation of varieties such as Cegledi Biborkajzsi C.244, Kéczkei rozsa barack C.171, Cegledi oriás and Mandula kajzsi C.712. When selecting 532 items of apricot, it was discovered that the size of fruits and excellent taste are rarely related to high productivity and even less to frost hardiness and drought tolerance [20]. Later, Szabó showed that Keczkei rozsa and Mandula kajzsi varieties introduced frost hardiness into its progeny dominantly and irrespective of them being used as female or male parents [21].

The degree of apricot adaptability is, to a certain extent, conditioned by the level of frost hardiness in buds, blossoms and juvenile fruits. Therefore, the period of vegetative dormancy (its intensity and length) ensures less dependence on external environmental conditions and is a crucial factor for frost hardiness. Based on its origin, the apricot is a mountainous plant and as such is typical for its adaptability to cold winter without fluctuating temperatures. Results of many authors have shown that buds of stone fruit experience important stages of micro- and macrosporogenesis at the time of autumn, winter and spring [22].

Greek colleagues have worked very hard studying apricot varieties mainly in a stage after a strong natural infection. Syrgiannidis was first to describe the Stark Early Orange and Stella cultivars as resistant against *plum pox virus* based on his fieldwork between the years of 1967– 1970. He then sorted the varieties into very sensitive (ProimoTyrinthos, Rouge de Sernhac), quite sensitive (Blenril, Canino, Docteur Mascle, Bergeron, Moorpark, Nugget, Rouge de Fournes, Sungold and Tilton a Stavropoulos) and less sensitive (Blenheim Royal, Ricordo di

Murg et al. [18] recorded the results of observing 64 apricot species infected with PPV in the region of Oradea and found that Stark Early Orange, Manitoba, Farmingdale, Bolšoj Rozovoj, Blenril, King, Doty, Arzami, Pize, Smyrna, Sophie, Tarzii de Bucuresti, Selena, Favorit, Timpurii de Chisinau, Rozova Scora, Rosii de Banesa, Magyar Kayszi and Borozi Rozsa varieties were not infected. Varieties which had no symptoms in fruits and weak symptoms on leaves were Venus, Tuzla, Saturn, Calatis, Dalia, Neptun, Sulina, Excelsior, Čačak, Raske

Clone selection can be regarded as an evolution of varieties when clones with higher adaptability are selected during a long-term growing period in a relatively vast area, whereas clones with lower adaptability to the environment cease to exist or are only relevant to a smaller section of areas. This is called the plasticity of a given cultivar which, influenced by its environment and length of growing, provides an array of clones with various adaptabilities. This plasticity is conditioned by the age of varieties. Hence, clone selection provides one of the solutions for increased adaptability for varieties such as Velkopavlovická, Hungarian best and Sabinovská. Clone selection can increase environmental adaptability as can the selection of additional properties other than just fertility. In the scope of several cycles of clone selection of apricots which Professor Vachůn initiated in the 60s, variability of the blooming period was also detected (a 1–2-day delay of full bloom can in some years positively influence the harvest), as was the number of blossoms with two pistils making the clones with higher number of pistils more efficient. Of other variable properties, clone selection can be used to improve the health of trees (early death or economically significant viral diseases) when even in these

In Hungary, Nyujtó chose large fruits of local cultivars and saw the European group of apricots as being suitable enough to achieve a faster and significant improvement in cultivar adaptability using clone selection as opposed to the creation of new cultivars through crossing. His program saw the creation of varieties such as Cegledi Biborkajzsi C.244, Kéczkei rozsa barack C.171, Cegledi oriás and Mandula kajzsi C.712. When selecting 532 items of apricot, it was discovered that the size of fruits and excellent taste are rarely related to high productivity and even less to frost hardiness and drought tolerance [20]. Later, Szabó showed that Keczkei rozsa and Mandula kajzsi varieties introduced frost hardiness into its progeny

The degree of apricot adaptability is, to a certain extent, conditioned by the level of frost hardiness in buds, blossoms and juvenile fruits. Therefore, the period of vegetative dormancy

dominantly and irrespective of them being used as female or male parents [21].

Amic and Grossa del Giardino) [17].

74 Breeding and Health Benefits of Fruit and Nut Crops

Carpo, Joubert, Toulon and Precoce de Italia [18].

**2.3. Achieved findings in apricot breeding**

traits conclusive differences were found [19].

Development of blossom buds at the time of dormancy is assured by individual changes and intensity of metabolism. Jablonskij and Elmanova found a relation between accumulated hydrocarbons and the pace of morphogenesis as well as a relation between morphological and anatomical changes and the dynamics of phenol contents in buds (r = 0,83). This occurrence relates to the fact that most phenols inhibit the growing process. They found that the highest contents of phenols in blossom buds were present (as detected in the beginning of April) in the following varieties: Narjadniy, Vynosliviy and Orfyand Amur [23].

Kostina et al. compared the inheritance of dormancy length in reciprocal crossing of varieties Zard and Shalah, which belong to two different eco-geographical groups with different dormancy lengths. In this cross, 34.8% of progeny inherited the character of slow development of Zard and 21.9% of progeny inherited the character of its father variety Zard [24].

The study on two progenies, where the late terminating dormancy variety Stark Early Orange (SEO) was the male variety for both of them and female varieties from early to mid-earlybreaking dormancy of Vestar and Velkopavlovická LE-19/2 were utilized, observed that in both the combinations 82.1% of hybrids terminated their deep vegetative dormancy after their parents and intermediates. New traits appeared in 17.9% of progeny, of which being positive (late dormancy termination) appeared in 9.4% [25].

In her breeding work, Kostina established a dominant inheritance character in apricots from Central Asia regarding several biological traits including late blooming and increased frost hardiness if apricots in this group had been crossed with apricots in the Irano-Caucasian group or European group [8]. Crossa-Raynaud (when crossing Canino and Amerleuch showed the intermediate inheritance character of development rhythm in flower buds of blooming period [26]. Zagorodnaya also found that more than a half of observed progeny possessed frosthardiness properties of blossom buds among parents and quite a considerable number of progeny were drawing nearer to their frost-hardy parent, particularly if this parent was used as a mother cultivar [27].

Oranževokrasniy is also recommended as a frost-hardy donor for crossing by Kostina together with Churmai variety (long dormancy period and late blooming) [24]. Hough also recommended (written statement, 1989) the crossing of Oranževokrasnyi × Orange Red with the aim of obtaining frost-hardy individuals with firm fruits and early ripening [28].

*Plum pox virus* (PPV) resistance presents one of the most discussed topics in European breeding programs. This topic has been reviewed by Zhebentyayeva et al. [29]. All apricot cultivars of European origin are susceptible to PPV. Currently, most apricot breeding programs in Europe use the PPV-resistant North American cultivars to introduce this trait into European germplasm. Resistance among apricots has been found only in some North American cultivars such as "Goldrich," "Harlayne," "Stark Early Orange," (SEO) "Stella" and "Harcot" [30, 31]. Therefore, most conventional breeding programs very often use one of these as a source of resistance in the development of new varieties. Badenes et al. were the first to suggest the role of Eastern Asiatic species, particularly *P. mandshurica,* as a potential source of PPV resistance into North American germ-plasm [32]. The results from Karayiannis et al. [31, 33] gave more support to this idea, even if not all the progenies of *P. mandshurica* were PPV resistant [34]. North American selections derived from *P. mandshurica* were introduced for their cold hardiness in midwinter and spring, late blooming and the ability to set fruit under adverse conditions for pollination [35]. Besides *P. mandshurica,* other East Asian species such as *P. sibirica* var. *davidiana* and *P. mume* may also have been involved in the pedigree of PPV-resistant North American apricots. A likely scenario for introgression of resistance into North American germplasm might include hybridization of European apricots with Northern Chinese varieties cultivated in overlapping areas of *P. armeniaca* and East Asian apricot species [28]. The histories of apricot domestication and of its resistance to Sharka are however still poorly understood. In another piece of work, Decroocq et al. used 18 microsatellite markers to genotype a collection of 230 wild trees from Central Asia and 142 cultivated apricots as representatives of the worldwide cultivated apricot germ-plasm. The genetic markers confirmed highest levels of diversity in both wild and cultivated apricots in their original areas (Central Asia, China). Furthermore, high frequency of resistance to Sharka was detected in apricots native to Central Asia [36].

resistance of Harlayne cultivar in a large F1 population [44]. Dondini et al. established that a major QTL for resistance to both PPV-M and PPV-D strains was found in the top half of "Lito" LG1 (just like in "Stark Early Orange") and when the resistant, tolerant and recovered seedlings were pooled together, the ratio of these to susceptible plants was fixed as 1:1, explaining why such a large part of the phenotypic variability is accounted for by a single QTL in the LG1 [45]. A first determinant was mapped on linkage group 1 (LG1) by using an F1 progeny of Goldrich 9 Valenciano [46]. These studies were also verified by a quantitative trait locus (QTL). Goldrich is known to be tolerant to the pathogen while Valenciano was highlighted as susceptible [39, 47]. This preliminary result was recently confirmed by a quantitative trait locus(QTL) analysis carried out on another F1 progeny, Goldrich 9 Currot [47]. A major QTL was also found in LG1 by the analysis of F1 and F2 progenies of Stark Early Orange (SEO) [48] and its progeny Lito [46, 49]. Minor QTL were discovered in the Polonais 9 SEO progeny in LG3 and LG5 of both SEO and Polonais [50]. The main QTL on LG1 was upheld by Sicard et al. [51] establishing new microsatellite (SSR) markers flanking the QTL and by Lalli et al. [52] in a backcross population of SEO 9 Vestar and was again confirmed by Soriano et al. [46] in the extended F2 Lito selfed progeny. Most of the resistance determinants shown above were characterized using the PPV-D strain as a source of inoculum and, at present, the only resistances specific to the PPV-M strain are those endorsed in the BC1 SEO xVestar [52] and in *P. davidiana* [44]. Dodini et al. [45] introduced a potentially different QTL which either explained a very small part of the variance or was below the LOD threshold. The markers closer to the QTL peaks would already be suitable for marker-assisted breeding. Those plants that recovered should

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be subjected to another observation to evaluate their tolerance or resistance.

apricot breeding program at the Mendel University in Brno, Czech Republic [53].

1. The apricot is a species with lower adaptability to the environment, which has, in the scope of its species, many genotypes—donors that can be used in breeding against biotic factors.

**2.** At present, there are substantial gene resources for apricot breeding and scientific activity relating to this species. These are used intensively to achieve mainly higher quality of fruits

**3. Conclusion**

and resistance to Sharka.

Krška et al. studied the inheritance of resistance to PPV in F1 progeny of cross "Harlayne" × "Betinka" obtained within gene pyramiding. The observed segregation ratio (153,30,16) for the progeny of "Harlayne" × "Betinka" was not significantly different from the predicted 12:3:1 segregation ratio (χ<sup>2</sup> = 2.551, P = 27.9%). These findings showed that PPV resistance in apricot is controlled by two independent dominant genes with epistatic interaction, where resistance would be a dominant trait and the resistant parents ("Harlayne" and "Betinka") would be heterozygous for both loci. Tolerant plants would be heterozygous for the hypostatic (masked) locus, and susceptible plants would be homozygous recessive for both loci. They established that it is possible to pyramidize genes for resistance to PPV in apricot and gain a cultivar with durable resistance to PPV. This principle was upheld in the

The PPV resistance trait in apricots was first mentioned by Dosba et al. [37]. In the population of Stark Early Orange × Screara, 64 inoculated (by both methods, chip budding and aphids) hybrids were tested for PPV and a polygynous character of heredity for PPV was found. Dosba et al. mentioned 30% resistant hybrids in the same hybrid combination [37]. Moustafa et al. agreed with Dosba [37]. A ratio of 3:1 sensitive/resistant genotypes in hybrid populations of North American PPV resistant varieties and local Spanish sensitive cultivars corresponds with the hypothesis of heredity to PPV resistance by two independent dominant genes. Resistance donors who used incrossing could be heterozygous for both loci. Only those heterozygous seedlings in both loci as the parental donors could be resistant [38].

On the contrary, Dicenta et al. established the ratio of 1:1 in 291of seedlings of 20 different crossing activities between resistant and sensitive parents. Based on these results, the authors conclude that PPV resistance in apricots is controlled by one dominant gene and that resistant parents could be heterozygous even in this trait [39].

Faculty of Horticulture in Lednice has got a long tradition of apricot breeding which started in 1990 and focuses on resistant breeding against Sharka (PPV). One of the parents used in the breeding program in Lednice (Harlayne) was assessed as resistant by Martínez-Gómez et al. [40], while both Dosba et al. [37] and Fuchs et al. [41] classified it as immune.

When crossing the parents of resistant to susceptible, using Harlayne species, a hypothesis was reached on heredity of three dominant genes which are responsible for PPV resistance [42]. In the progeny derived from the four apricot crosses (resistant to susceptible), the segregation ratios were compatible with the hypothesis of three dominant genes being responsible for PPV resistance, with "Harlayne" being heterozygous for all three genes [43]. Four quantitative trait loci (QTLs) were identified of which three mapped on linkage group 1 (LG1) which explained between 5 and 39% of the phenotypic variance. This happened when analyzing quantitative resistance of Harlayne cultivar in a large F1 population [44]. Dondini et al. established that a major QTL for resistance to both PPV-M and PPV-D strains was found in the top half of "Lito" LG1 (just like in "Stark Early Orange") and when the resistant, tolerant and recovered seedlings were pooled together, the ratio of these to susceptible plants was fixed as 1:1, explaining why such a large part of the phenotypic variability is accounted for by a single QTL in the LG1 [45].

A first determinant was mapped on linkage group 1 (LG1) by using an F1 progeny of Goldrich 9 Valenciano [46]. These studies were also verified by a quantitative trait locus (QTL). Goldrich is known to be tolerant to the pathogen while Valenciano was highlighted as susceptible [39, 47]. This preliminary result was recently confirmed by a quantitative trait locus(QTL) analysis carried out on another F1 progeny, Goldrich 9 Currot [47]. A major QTL was also found in LG1 by the analysis of F1 and F2 progenies of Stark Early Orange (SEO) [48] and its progeny Lito [46, 49]. Minor QTL were discovered in the Polonais 9 SEO progeny in LG3 and LG5 of both SEO and Polonais [50]. The main QTL on LG1 was upheld by Sicard et al. [51] establishing new microsatellite (SSR) markers flanking the QTL and by Lalli et al. [52] in a backcross population of SEO 9 Vestar and was again confirmed by Soriano et al. [46] in the extended F2 Lito selfed progeny. Most of the resistance determinants shown above were characterized using the PPV-D strain as a source of inoculum and, at present, the only resistances specific to the PPV-M strain are those endorsed in the BC1 SEO xVestar [52] and in *P. davidiana* [44]. Dodini et al. [45] introduced a potentially different QTL which either explained a very small part of the variance or was below the LOD threshold. The markers closer to the QTL peaks would already be suitable for marker-assisted breeding. Those plants that recovered should be subjected to another observation to evaluate their tolerance or resistance.

Krška et al. studied the inheritance of resistance to PPV in F1 progeny of cross "Harlayne" × "Betinka" obtained within gene pyramiding. The observed segregation ratio (153,30,16) for the progeny of "Harlayne" × "Betinka" was not significantly different from the predicted 12:3:1 segregation ratio (χ<sup>2</sup> = 2.551, P = 27.9%). These findings showed that PPV resistance in apricot is controlled by two independent dominant genes with epistatic interaction, where resistance would be a dominant trait and the resistant parents ("Harlayne" and "Betinka") would be heterozygous for both loci. Tolerant plants would be heterozygous for the hypostatic (masked) locus, and susceptible plants would be homozygous recessive for both loci. They established that it is possible to pyramidize genes for resistance to PPV in apricot and gain a cultivar with durable resistance to PPV. This principle was upheld in the apricot breeding program at the Mendel University in Brno, Czech Republic [53].

#### **3. Conclusion**

Therefore, most conventional breeding programs very often use one of these as a source of resistance in the development of new varieties. Badenes et al. were the first to suggest the role of Eastern Asiatic species, particularly *P. mandshurica,* as a potential source of PPV resistance into North American germ-plasm [32]. The results from Karayiannis et al. [31, 33] gave more support to this idea, even if not all the progenies of *P. mandshurica* were PPV resistant [34]. North American selections derived from *P. mandshurica* were introduced for their cold hardiness in midwinter and spring, late blooming and the ability to set fruit under adverse conditions for pollination [35]. Besides *P. mandshurica,* other East Asian species such as *P. sibirica* var. *davidiana* and *P. mume* may also have been involved in the pedigree of PPV-resistant North American apricots. A likely scenario for introgression of resistance into North American germplasm might include hybridization of European apricots with Northern Chinese varieties cultivated in overlapping areas of *P. armeniaca* and East Asian apricot species [28]. The histories of apricot domestication and of its resistance to Sharka are however still poorly understood. In another piece of work, Decroocq et al. used 18 microsatellite markers to genotype a collection of 230 wild trees from Central Asia and 142 cultivated apricots as representatives of the worldwide cultivated apricot germ-plasm. The genetic markers confirmed highest levels of diversity in both wild and cultivated apricots in their original areas (Central Asia, China). Furthermore, high frequency of resistance to Sharka was detected in apricots native to Central Asia [36].

The PPV resistance trait in apricots was first mentioned by Dosba et al. [37]. In the population of Stark Early Orange × Screara, 64 inoculated (by both methods, chip budding and aphids) hybrids were tested for PPV and a polygynous character of heredity for PPV was found. Dosba et al. mentioned 30% resistant hybrids in the same hybrid combination [37]. Moustafa et al. agreed with Dosba [37]. A ratio of 3:1 sensitive/resistant genotypes in hybrid populations of North American PPV resistant varieties and local Spanish sensitive cultivars corresponds with the hypothesis of heredity to PPV resistance by two independent dominant genes. Resistance donors who used incrossing could be heterozygous for both loci. Only those

On the contrary, Dicenta et al. established the ratio of 1:1 in 291of seedlings of 20 different crossing activities between resistant and sensitive parents. Based on these results, the authors conclude that PPV resistance in apricots is controlled by one dominant gene and that resistant

Faculty of Horticulture in Lednice has got a long tradition of apricot breeding which started in 1990 and focuses on resistant breeding against Sharka (PPV). One of the parents used in the breeding program in Lednice (Harlayne) was assessed as resistant by Martínez-Gómez et al.

When crossing the parents of resistant to susceptible, using Harlayne species, a hypothesis was reached on heredity of three dominant genes which are responsible for PPV resistance [42]. In the progeny derived from the four apricot crosses (resistant to susceptible), the segregation ratios were compatible with the hypothesis of three dominant genes being responsible for PPV resistance, with "Harlayne" being heterozygous for all three genes [43]. Four quantitative trait loci (QTLs) were identified of which three mapped on linkage group 1 (LG1) which explained between 5 and 39% of the phenotypic variance. This happened when analyzing quantitative

heterozygous seedlings in both loci as the parental donors could be resistant [38].

[40], while both Dosba et al. [37] and Fuchs et al. [41] classified it as immune.

parents could be heterozygous even in this trait [39].

76 Breeding and Health Benefits of Fruit and Nut Crops


**3.** In chosen and traditionally most significant apricot cultivars, it is essential to carry out clone selection and it is important to maintain this process so that the biological quality of clones is maintained. The use of local genotypes and clones of genetically similar cultivars should not just be the nostalgia of "good old days" but the reality of economically justified growing and agri-tourist utilization.

[5] Blaha J, Luža J, Kalášek J. Broskvoně, meruňky, mandloně. Ovocnickáedice. Academia,

Genetic Apricot Resources and their Utilisation in Breeding

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[6] Kostina KF. Introdukcija i selekcija abrikosa. Selskochozjajstvennaja biologija, Tom VII,

[7] Bourguiba H, Audergon JM, Krichen L, Trifi-Farah N, Mamouni A, Trabelsi S, D'Onofrio C, Asma B, Santoni S, Khadari B. Loss of genetic diversity as a signature of apricot domestication and diffusion into the Mediterranean Basin. BMC Plant Biology. 2012;**12**:49.

[8] Kostina KF. The use of varietal resources of apricots for breeding. TrudNikit Bot Sad.

[9] Michurin IV. Itogi šestidesjatiletnich rabot. Ogizselchozgiz, Moskva; 1949. (in Russian) [10] Whitson J, John R, Williams HS. Luther Burbank: His Methods and Discoveries and their Practical Application. Vol. I. New York and London: Luther Burbank Press; 1914 [11] Hummel R. A classification of hardy north American Prunus cultivars and native species

[12] Krška B. Genetic resources of apricot for adaptability improvement and breeding. In: Proceedings of the XIV International Symposium on Apricot Breeding and Culture, Matera, Italy. International Society for Horticultural Science (ISHS); 2010. Vol. 862 [13] Djurič B. Prilogproučavanjuosetljivostimladihplodovasortikajsijepremapoznomproléčn

[14] Bassi D et al. Tolerance of apricot to winter temperature fluctuation and spring frost in

[15] Krška B. Hodnocení vnitrodruhových hybridů *Prunus armeniaca* L. z hlediska jejich využití v pěstitelské praxi a ve šlechtění [PhD thesis]. Mendel University in Brno; 1996

[16] Chloupek O. Genetická diverzita, šlechtění a semenářství. Praha: Academia; 1995 (in

[17] Syrgiannidis GD. Observations on the sensitivity of certain apricot varieties to Sarka

[18] Murg S, Minoiu N, Isac M, Stegerean P. Sensibilita teadiferitelorsoiuri de cais la virusul

[19] Vachůn Z. Clonal selection on apricot cultivar Velkopavlovická. [Habitant thesis] Mendel

[20] Nyujtó F. Significance of clone selection in stone fruits. Agrártudományii Közlemények.

[21] Szabó Z, Nyéki J. Blossoming, fructification and combination of apricot varieties. In: Book of Abstracts, IX. International Symposium on Apricot Culture; 9-15 July 1989;

(plum pox) virus. Geoponika, No. 218, May–June 1974, 129-132

University in Brno. Lednice; 1992. pp. 193-194. (in Czech)

plum pox. Protectia Plantelor; 1999. Anul IX. Nr. 35 a. 44. (in Romanian)

Published online 2012 Apr 17. DOI: 10.1186/1471-2229-12-49

based on hardiness zones. Fruit Varieties Journal. 1976;**29**(4):62-70

ommrazu. Jug. Vočarstvo br. 1982;**2**:41-47 (in Serbian)

northern Italy. Acta Horticulturae. 1995;**384**:315-322

Praha; 1966 (in Czech)

1969;**41**:45-63

(in Czech)

1970;**29**:445-457

Caserta, Italy. pp. 64

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No. 1, 1972, 86-91. (in Russian)


### **Author details**

Boris Krška

Address all correspondence to: borokrska@gmail.com

Independent Researcher, Lednice, Czech Republic

#### **References**


[5] Blaha J, Luža J, Kalášek J. Broskvoně, meruňky, mandloně. Ovocnickáedice. Academia, Praha; 1966 (in Czech)

**3.** In chosen and traditionally most significant apricot cultivars, it is essential to carry out clone selection and it is important to maintain this process so that the biological quality of clones is maintained. The use of local genotypes and clones of genetically similar cultivars should not just be the nostalgia of "good old days" but the reality of economically justified

**4.** The study of apricot germ-plasm so far has enabled not only the selection of suitable genotypes to be tested in growing practice but also the selection of suitable genotypes for targeted breeding while aiming to use gene variability. Results obtained using the analysis of polymorphic markers have enabled breeders to carry out their work. It was possible to use them in the context of evaluated collection of apricot genotypes to identify the difference to assess the genetic similarity and more detailed characteristics of varieties, genetic resources and breeding material as well as for methods of early selection of progenies with chosen traits. Developed molecular markers for selection in breeding programs (MAS) represent almost a routine method today, particularly in the area of

**5.** New methods of selection (MAS) have so far aided in early selection of traits of PPV resist-

**6.** Currently, newly created cultivars present the opportunity of genotype selection with appropriately combined traits of resistance to abiotic or biotic pathogens but also in terms of requirement for an ideotype of fruits with increased adaptability, so that the apricot

[1] Top Ten Countries by Apricot Production. "Ranking America". [Internet] 2013. Available

[2] Kostina KF. Proischozzdeniye i evoluciya kulturnogo abrikosa. Trudy GNBS, Tom

[3] Vavilov IN. Bailey CH, Hough LF. (1975). Apricots. In: Janick J, Moore JN, editors. Advances in Fruit Breeding. West Lafayette, Indiana: Purduee University Press; 1926 [4] Vavilov IN. Phytogeographic basis of plant breeding. The origin, variation, immunity and breeding of cultivated plants (tr. K.S. Chester). Chronica Botanica. 1951:13-54

from: http://faostat.fao.org/site/567. [Accessed: October 8, 2013]

XXIV, vyp. 1, Krymizdat. 1946. pp. 25-39. (In Russian)

could again, as stated by Hough and Bailey, descend from the mountains.

growing and agri-tourist utilization.

78 Breeding and Health Benefits of Fruit and Nut Crops

ance and determination of self-compatibility.

Address all correspondence to: borokrska@gmail.com

Independent Researcher, Lednice, Czech Republic

resistant breeding.

**Author details**

Boris Krška

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**Section 2**

**Health Benefits**


**Section 2**
