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Jameel M. Al-Khayri · Shri Mohan Jain
Dennis V. Johnson Editors
Advances in
Plant Breeding
Strategies: Nut and
Beverage Crops
Volume 4
Advances in Plant Breeding Strategies:
Nut and Beverage Crops
Jameel M. Al-Khayri • Shri Mohan Jain
Dennis V. Johnson
Editors
Advances in Plant Breeding

Strategies: Nut and Beverage
Crops
Volume 4
Editors
Jameel M. Al-Khayri Shri Mohan Jain
Agricultural Biotechnology Agricultural Sciences
King Faisal University University of Helsinki
Al-Hassa, Saudi Arabia Helsinki, Finland
Dennis V. Johnson
Cincinnati, OH, USA
ISBN 978-3-030-23111-8 ISBN 978-3-030-23112-5 (eBook)

https://doi.org/10.1007/978-3-030-23112-5
© Springer Nature Switzerland AG 2019

This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of
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Preface
Contemporary plant breeders no longer need to rely solely on traditional method-

ologies in their work of assuring a sustainable and elastic level of world food pro-
duction. However, human population is increasing at an alarming rate in developing
countries, and food availability could gradually become a serious problem.
Agriculture production is severely affected because of environmental pollution,
rapid industrialization, water scarcity and quality, erosion of fertile topsoil, limited
remaining arable land to expand production area, lack of improvement of local plant
types, erosion of genetic diversity, and dependence on only few crop species for
food supply worldwide. According to the FAO, 70% more food must be produced
over the next four decades to feed a projected population of 9 billion people by the
year 2050. Currently, only 30 plant species are used to meet 95% of the world’s food
requirements, which are considered as the major crops. The breeding programs of
these crops have been very much dependent on the ready availability of genetic
variation, either spontaneous or induced. Plant breeders and geneticists are under
constant pressure to sustain and increase food production by using innovative breed-
ing strategies and introducing minor crops that are well adapted to marginal lands
and can provide source of nutrition through tolerance of abiotic and biotic stresses.
In traditional breeding, introgression of one or a few genes into a cultivar is carried
out via backcrossing over several plant life cycles.
With the development of new molecular tools, molecular marker-assisted back-
crossing has facilitated rapid introgression of a transgene into a plant and reduced
linkage drag. Continued development and adaptation of plant biotechnology,
molecular markers, and genomics have established ingenious new tools for the cre-
ation, analysis, and manipulation of genetic variation for the development of
improved cultivars. For example, molecular breeding has great potential to become
the standard practice in the improvement of several fruit crops. Adopting a multidis-
ciplinary approach comprised of traditional plant breeding, mutation breeding,
plant biotechnology, and molecular biology would be strategically ideal for devel-
oping new improved crop varieties. This book highlights the recent progress in the
development of plant biotechnology, associated molecular tools, and their usage in
plant breeding.
v
vi Preface
The basic concept of this book is to examine the best use of both innovative and
traditional methods of plant breeding to develop new crop varieties suited to differ-
ent environmental conditions to achieve sustainable food production and enhanced
food security in a changing global climate, in addition to the development of crops
for enhanced production of pharmaceuticals and innovative industrial uses. Three
volumes of this book series were published in 2015, 2016, and 2018, respectively:
Volume 1, Breeding, Biotechnology and Molecular Tools; Volume 2, Agronomic,
Abiotic and Biotic Stress Traits; and Volume 3, Fruits. In 2019, the following four
volumes are concurrently being published: Volume 4, Nut and Beverage Crops;
Volume 5, Cereals; Volume 6, Industrial and Food Crops; and Volume 7, Legumes.
This Volume 4, subtitled Nut and Beverage Crops, focuses on the advances in
breeding strategies using both traditional and modern approaches for the improve-
ment of individual plantation crops. Included in Part I are 11 important nut species
recognized for their economical and nutritional importance including almond,
argan, Brazil nut, cashew nut, chestnut, hazelnut, macadamia, peanut, pine nut, pis-
tachio, and walnut. Part II covers two popular beverage species, coffee and tea.
Chapters are written by internationally reputable scientists and subjected to a
review process to assure quality presentation and scientific accuracy. Each chapter
begins with an introduction covering related backgrounds and provides in-depth
discussion of the subject supported with high-quality color photos, illustrations, and
relevant data. This volume contains a total of 108 figures and 55 tables to illustrate
presented concepts. The chapter concludes with an overview of the current status of
breeding and recommendations for future research directions as well as appendixes
listing research institutes and genetic resources relevant to the topic crop. A compre-
hensive list of pertinent references is provided to facilitate further reading.
The book is an excellent reference source for plant breeders and geneticists
engaged in breeding programs involving biotechnology and molecular tools together
with traditional breeding. It is suitable for both advanced undergraduate and post-
graduate students specializing in agriculture, biotechnology, and molecular breed-
ing as well as for seed companies and policy-makers.
We are greatly appreciative of all the chapter authors for their contributions
towards the success and quality of this book. We are proud of this diverse collabora-
tive undertaking, especially since this volume represents the efforts of 53 scientists
from 13 countries. We are also grateful to Springer for giving us an opportunity to
compile this book.
Al-Hassa, Saudi Arabia Jameel M. Al-Khayri

Helsinki, Finland Shri Mohan Jain
Cincinnati, OH, USA Dennis V. Johnson
Contents
Part I Nut Crops

1 Almond [Prunus dulcis (Miller) D.A. Webb] Breeding�� 3
Pedro J. Martínez-García, Angela S. Prudencio,
Raquel Sánchez-Pérez, Felipe Pérez De Los Cobos, Ossama Kodad,
Hassouna Gouta, Manuel Rubio, and Pedro Martínez-Gómez
2 Genetic Diversity and Breeding of Argan Tree
(Argania spinosa L. Skeels)�� 31
Naima Ait Aabd, Rachid Bouharroud, Abdelghani Tahiri,
Ahmed Wifaya, Abdelaziz Mimouni, and Abdelhamid El Mousadik
3 Brazil Nut (Bertholletia excelsa Bonpl.) Breeding�� 57
Aisy Botega Baldoni, Lúcia Helena de Oliveira Wadt,
and Cássia Ângela Pedrozo
4 Cashew Nut (Anacardium occidentale L.) Breeding Strategies�� 77
Suvalaxmi Palei, Rasheswari Dasmohapatra, Sunil Samal,
and Gyana Ranjan Rout
5 Chestnut (Castanea spp. Miller) Breeding�� 105
J. Hill Craddock and M. Taylor Perkins
6 Hazelnut (Corylus spp.) Breeding �� 157
Roberto Botta, Thomas J. Molnar, Veli Erdogan, Nadia Valentini,
Daniela Torello Marinoni, and Shawn A. Mehlenbacher
7 Macadamia (Macadamia spp.) Breeding�� 221
Bruce L. Topp, Catherine J. Nock, Craig M. Hardner,
Mobashwer Alam, and Katie M. O’Connor
8 Peanut (Arachis hypogaea L.) Breeding �� 253
Saikat Gantait, Jitendriya Panigrahi, Illa C. Patel,
Catherine Labrooy, Arulthambi Luke Rathnakumar,
and Jeshima Khan Yasin
vii
viii Contents
9 Advances in Edible Pine Nut Trees (Pinus spp.)

Breeding Strategies�� 301
Hanguo Zhang and Zhen Zhang
10 Pistachio (Pistacia spp.) Breeding�� 353
Abdollatif Sheikhi, Mohammad M. Arab, Patrick J. Brown,
Louise Ferguson, and Mohammad Akbari
11 Advances in Persian Walnut (Juglans regia L.) Breeding Strategies�� 401
Kourosh Vahdati, Mohammad M. Arab,
Saadat Sarikhani, Mohammad Sadat-Hosseini,
Charles A. Leslie, and Patrick J. Brown
Part II Beverage Crops

12 Genetic Resources and Breeding of Coffee (Coffea spp.) �� 475
Manoj K. Mishra
13 Advances in Tea [Camellia sinensis (L.) O. Kuntze] Breeding�� 517
Mahasen A. B. Ranatunga
Index�� 567
About the Editors and Contributors
Editors
Jameel M. Al-Khayri is a Professor of Plant

Biotechnology affiliated with the Department of
Agricultural Biotechnology, King Faisal University,
Saudi Arabia. He received his B.S. in Biology in 1984
from the University of Toledo and his M.S. in
Agronomy in 1988 and Ph.D. in Plant Science in 1991
both from the University of Arkansas. He is a Member
of the International Society for Horticultural Science
and Society for In Vitro Biology as well as the National
Correspondent of the International Association of
Plant Tissue Culture and Biotechnology. His graduate
work resulted in the establishment of in vitro regenera-
tion protocols for spinach and zoysiagrass. For the last
two decades, he dedicated his research efforts to date
palm. He has authored over 60 research articles in ref-
ereed international journals and 25 review chapters and
edited 7 journal special issues. In addition, he edited 5
reference books on date palm biotechnology and utili-
zation of genetic resources and 7 volumes of the book
series Advances in Plant Breeding Strategies. He has
been involved in organizing international scientific
conferences and contributed numerous research pre-
sentations. In addition to teaching, student advising,
and research, he held administrative responsibilities as
the Assistant Director of Date Palm Research Center,
Head of the Department of Plant Biotechnology, and
Vice Dean for the Development and Quality Assurance.
He served as a Member of Majlis-ash-Shura (Saudi
ix
x About the Editors and Contributors
Legislative Council) for the 2009–2012 term. Currently,

he is maintaining an active research program on date
palm focusing on genetic transformation, secondary
metabolites, and in vitro mutagenesis to enhance toler-
ance to abiotic and biotic stress.
Shri Mohan Jain is a Consultant and Plant

Biotechnologist, Department of Agricultural Sciences,
University of Helsinki, Helsinki, Finland; he received
his M.Phil. in 1973 and Ph.D. in 1978 at Jawaharlal
Nehru University, New Delhi, India. He was a
Postdoctoral Fellow in Israel and the USA; a Visiting
Scientist/Professor in Japan, Malaysia, Germany, and
Italy; and a Technical Officer, Plant Breeding and
Genetics, International Atomic Energy Agency (IAEA),
in Vienna, Austria, 1999–2005. He is a Member of the
International Association for Plant Tissue Culture and
Biotechnology and Editorial Board Member of
Euphytica, In Vivo, Propagation of Ornamental Plants,
Emirates J. Food and Agriculture, and the series Forest
Health and Biotechnology. His publications are more
than 160 in peer reviewed journals, book chapters, and
conference proceedings. He edited 55 books and acted
as a Chair Person in several international conferences
worldwide and has been an Invited Speaker. He was
awarded Nobel Peace Prize in 2005 in commemoration
of the awarding to the IAEA of the Nobel Peace Prize
for 2005. In addition, he was Former Consultant to the
IAEA, the European Union, the Government of
Grenada, Iranian Private Company, and the Egyptian
Government. Currently, his research interests are on
somatic embryogenesis, organogenesis, haploidy,
somatic cell hybridization, somaclonal variation, and
mutagenesis mainly in medicinal plants, date palm, and
banana genetic improvement, genetic diversity, ero-
sion, conservation, and utilization in the context of cli-
mate change and food and nutritional security.
About the Editors and Contributors xi
Dennis V. Johnson is a Consultant and Former

University Professor. He is a graduate of the University
of California, Los Angeles, where he completed his
B.A. (1966), M.A. (1970), and Ph.D. (1972) degrees in
Geography, with specialization in agriculture and bio-
geography. He has taught at several colleges and uni-
versities, including the University of Houston, and was
a Visiting Professor for 2 years at the University of
Ceará, Fortaleza, Brazil. He also has worked exten-
sively with the international development agencies pro-
viding technical assistance to agriculture and forestry
on projects and programs in Africa, Asia, Europe, and
Latin America. He has published numerous articles on
palm utilization and conservation and has edited or
written books for the FAO, IUCN, and UNEP. He has
also translated into English plant science books from
Portuguese and Spanish. A decade ago, he began to
focus his research on date palm, in particular its intro-
duction to nontraditional areas such as Spain, North
and South America, and Australia. Moreover, he coau-
thored a book on date growing in the USA, has made
presentations at five international date palm confer-
ences, and coedited books on date palm, sago palm,
and plant breeding.
Contributors
Naima Ait Aabd National Institute for Agricultural Research (INRA)-CRRA,

Agadir, Morocco
Mohammad Akbari Department of Horticultural Sciences, Faculty of Agriculture,
University of Tabriz, Tabriz, Iran
Mobashwer Alam Queensland Alliance for Agriculture and Food Innovation,
University of Queensland, Brisbane, QLD, Australia
Mohammad M. Arab Department of Horticulture, Aburaihan Campus, University
of Tehran, Pakdasht, Tehran, Iran
Aisy Botega Baldoni Brazilian Agricultural Research Corporation, Embrapa
Agrossilvipastoril, Sinop, Mato Grosso, Brazil
Roberto Botta Dipartimento di Scienze Agrarie, Forestali e Alimentari, Università
degli Studi Torino, Turin, Italy
xii About the Editors and Contributors
Rachid Bouharroud National Institute for Agricultural Research (INRA)-CRRA,

Agadir, Morocco
Patrick J. Brown Department of Plant Sciences, University of California, Davis,
CA, USA
J. Hill Craddock Department of Biology, Geology, and Environmental Science,
University of Tennessee at Chattanooga, Chattanooga, TN, USA
Rasheswari Dasmohapatra Department of Fruit Science & Horticulture
Technology, College of Agriculture, Odisha University of Agriculture & Technology,
Bhubaneswar, Odisha, India
Felipe Pérez De Los Cobos CEBAS-CSIC, Murcia, Spain
Lúcia Helena de Oliveira Wadt Brazilian Agricultural Research Corporation,
Embrapa Rondônia, Porto Velho, Rondônia, Brazil
Abdelhamid El Mousadik Laboratory of Biotechnology and Valorization of
Natural Resources, Ibn Zohr University, Agadir, Morocco
Veli Erdogan Department of Horticulture, Faculty of Agriculture, Ankara
University, Ankara, Turkey
Louise Ferguson Department of Plant Sciences, University of California, Davis,
USA
Saikat Gantait All India Coordinated Research Project on Groundnut, Directorate
of Research, Bidhan Chandra Krishi Viswavidyalaya, Kalyani, West Bengal, India
Crop Research Unit (Genetics and Plant Breeding), Bidhan Chandra Krishi
Viswavidyalaya, Mohanpur, West Bengal, India
Hassouna Gouta Olive Tree Institute, Sfax, Tunisia
Craig M. Hardner Queensland Alliance for Agriculture and Food Innovation,
Saadat Sarikhani Department of Horticulture, College of Aburaihan, University
of Tehran, Tehran, Iran
Ossama Kodad Ecole Nationale d’Agriculture de Meknes, Meknes, Morocco
Catherine Labrooy Department of Crop Science, Faculty of Agriculture,
Universiti Putra Malaysia, Serdang, Selangore, Malaysia
Charles A. Leslie Department of Plant Sciences, University of California, Davis,
CA, USA
Daniela Torello Marinoni Dipartimento di Scienze Agrarie, Forestali e Alimentari,
Università degli Studi Torino, Turin, Italy
Pedro J. Martínez-García CEBAS-CSIC, Murcia, Spain
About the Editors and Contributors xiii
Pedro Martínez-Gómez CEBAS-CSIC, Murcia, Spain

Shawn A. Mehlenbacher Department of Horticulture, Oregon State University,
Corvallis, OR, USA
Abdelaziz Mimouni National Institute for Agricultural Research (INRA)-CRRA,
Agadir, Morocco
Manoj K. Mishra Tissue Culture and Plant Biotechnology Division, Unit of
Central Coffee Research Institute, Coffee Board, Dr. S. Radhakrishnan Road,
Manasagangothri, Mysore, Karnataka, India
Thomas J. Molnar Plant Biology Department, Rutgers University, New
Brunswick, NJ, USA
Catherine J. Nock Southern Cross Plant Science, Southern Cross University,
Lismore, NSW, Australia
Katie M. O’Connor Queensland Alliance for Agriculture and Food Innovation,
Suvalaxmi Palei Department of Fruit Science & Horticulture Technology, College
of Agriculture, Odisha University of Agriculture & Technology, Bhubaneswar,
Odisha, India
Jitendriya Panigrahi Department of Biotechnology, Shri A.N. Patel Post Graduate
Institute of Science and Research, Anand, Gujarat, India
Illa C. Patel Department of Life Sciences, Hemchandracharya North Gujarat
University, Patan, Gujarat, India
Cássia Ângela Pedrozo Brazilian Agricultural Research Corporation, Embrapa
Roraima, Boa Vista, Roraima, Brazil
M. Taylor Perkins Department of Biology, Geology, and Environmental Science,
University of Tennessee at Chattanooga, Chattanooga, TN, USA
Angela S. Prudencio CEBAS-CSIC, Murcia, Spain
Mahasen A. B. Ranatunga Plant Breeding Division, Tea Research Institute of Sri
Lanka, Talawakelle, Sri Lanka
Arulthambi Luke Rathnakumar ICAR-Directorate of Groundnut Research,
Junagadh, Gujarat, India
Gyana Ranjan Rout Department of Agricultural Biotechnology, College of
Agriculture, Odisha University of Agriculture & Technology, Bhubaneswar, Odisha,
India
Manuel Rubio CEBAS-CSIC, Murcia, Spain
Mohammad Sadat-Hosseini Department of Horticulture, College of Aburaihan,
University of Tehran, Tehran, Iran
Department of Horticulture, Faculty of Agriculture, University of Jiroft, Jiroft, Iran
xiv About the Editors and Contributors
Sunil Samal College of Horticulture, Chiplima, Odisha University of Agriculture

& Technology, Chiplima, Burla, Sambalpur, India
Raquel Sánchez-Pérez CEBAS-CSIC, Murcia, Spain
Abdollatif Sheikhi Department of Horticulture, Faculty of agriculture, Vali-e-Asr
University of Rafsanjan, Rafsanjan, Kerman, Iran
Abdelghani Tahiri National Institute for Agricultural Research (INRA)-CRRA,
Agadir, Morocco
Bruce L. Topp Queensland Alliance for Agriculture and Food Innovation,
Kourosh Vahdati Department of Horticulture, College of Aburaihan, University
of Tehran, Tehran, Iran
Nadia Valentini Dipartimento di Scienze Agrarie, Forestali e Alimentari,
Università degli Studi Torino, Turin, Italy
Ahmed Wifaya National Institute for Agricultural Research (INRA)-CRRA,
Agadir, Morocco
Jeshima Khan Yasin Division of Genomic Resources, ICAR-National Bureau of
Plant Genetic Resources, New Delhi, Delhi, India
Hanguo Zhang State Key Laboratory of Tree Genetics and Breeding, Northeast
Forestry University, Harbin, China
Zhen Zhang Research Institute of Subtropical Forestry, Chinese Academy of
Forestry, Hangzhou, China
Part I
Nut Crops
Chapter 1
Almond [Prunus dulcis (Miller)
D.A. Webb] Breeding
Pedro J. Martínez-García, Angela S. Prudencio, Raquel Sánchez-Pérez,

Felipe Pérez De Los Cobos, Ossama Kodad, Hassouna Gouta, Manuel Rubio,
and Pedro Martínez-Gómez
Abstract Almond [Prunus dulcis (Miller) D.A. Webb] presents clear challenges
within plant breeding given its plurennial woody character, long juvenile period and
multiplication by grafting. These challenges make the improvement process gener-
ally long and tedious. Therefore, it is necessary to have the most current information
on developing new cultivars that currently take 12 years. Additionally, the breeder
has to consider diverse internal (genetic background of the existing material, actual
and the new methodologies) and external factors (consumer preferences, biotic and
abiotic factors) to ensure the success of a new cultivar. The degree of knowledge of
these aspects determines the quality of the prediction and success in the design of
new almond cultivars. Although the size of the breeding population can be unlim-
ited, the management, phenotyping and selection of these seedlings are major limit-
ing factors. High-throughput phenotyping methods, genomic (DNA), transcriptomic
(RNA) and epigenetic studies can help to develop better selection strategies particu-
larly useful to deal with complex target traits in tree crops such as almond, with a
long juvenile periods and a high degree of heterozygosity. Recently, epigenetic
marks have been developed associated to dormancy in flower buds.
Keywords Almond · Breeding · Epigenetic marks · Genomics · Molecular

markers · Prunus · Transcriptomics
P. J. Martínez-García (*) · A. S. Prudencio · R. Sánchez-Pérez · F. P. De Los Cobos

M. Rubio · P. Martínez-Gómez
CEBAS-CSIC, Murcia, Spain
e-mail: pjmartinez@cebas.csic.es; asanchez@cebas.csic.es; rsanchez@cebas.csic.es;
mrubio@cebas.csci.es; pmartinez@cebas.csic.es
O. Kodad
Ecole Nationale d’Agriculture de Meknes, Meknes, Morocco
H. Gouta
Olive Tree Institute, Sfax, Tunisia
© Springer Nature Switzerland AG 2019 3

J. M. Al-Khayri et al. (eds.), Advances in Plant Breeding Strategies: Nut
and Beverage Crops, https://doi.org/10.1007/978-3-030-23112-5_1
4 P. J. Martínez-García et al.
1.1 Introduction
The cultivated almond [Prunus dulcis (Miller) Webb, syn. P. amygdalus (Batsch)],
of the Rosaceae family, is an ancient domesticated nut crop that has coevolved with
human civilization. The edible and tasty kernel of this nut crop species is the main
reason for its production and use (Gradziel and Martínez-Gómez 2013).
Almond is the only Prunus species cultivated exclusively for its kernel (Fig. 1.1).
Today, almonds are cultivated in more than 50 countries (http://faostat.fao.org),
with 95% grown in California, Australia and the Mediterranean Basin. Historically,
almonds were consumed as fresh and processed food, and are actually considered as
a functional food with both nutritional and purported medical properties including
purported anti-inflammatory and hypocholesterolemia properties (Musa-Velasco
et al. 2016; Poonan et al. 2011).
The objective of this review is to assess new breeding perspectives including
nutraceutical values and drought resistance together with the development of the
more recent technologies applied to the breeding programs including DNA, RNA
and epigenetic marks.
Fig. 1.1 (a) Cultivated almond tree, (b) Almond flowers, (c) Fruit and kernel morphology.
(Source: Figure created by Raquel Sánchez-Pérez)
1 Almond [Prunus dulcis (Miller) D.A. Webb] Breeding 5
1.2 Origin of Cultivation
A wide dissemination of cultivated almond is notable today; several authors have

distinguished four different stages: (a) Asiatic, (b) Mediterranean, (c) Californian
and (d) Southern Hemisphere stages. Gradziel (2011) described each stage in detail.
In each of the different stages, there was particular reference to almonds in several
aspects of human civilization including in literature, medicine and food. However,
although the ancestry and domestication history of cultivated almond has been stud-
ied extensively (Gradziel 2011; Grasselly 1976; Kester et al. 1991; Ladizinky 1999;
Velasco et al. 2016; Watkins 1979), many facts still remain unknown.
Central and western Asia are believed to be the origin of the wild almond rela-
tives (Browicz 1972; Browicz and Zohary 1996; Denisov 1988; Grasselly and
Crossa-Reynaud 1980; Grasselly 1976; Kester et al. 1991; Komorov et al. 1941).
Primarily, Amygdalus communis L. (syn. Prunus communis Archang.) was pro-
posed as the origin of cultivated almond (Kovalyov and Kostina 1935). A later study
by Ladizinsky (1999) proposed A. fenziliana Fiitsch as the ancestral species of the
cultivated almond. This origin propositions have led to at least three different theo-
ries being suggested, which seem to have some degree of communality. The first
states that the cultivated almond derived from wild Near East P. dulcis populations
(Levant theory), the second hypothesizes that cultivated almonds are stabilized
hybrids among south-western Asian species (hybrid theory) and the third theory is
based on coalescence-based models, which partly supports both scenarios and sug-
gests that most almond lineages differentiated during the Holocene (within the past
10,000 years), concomitant with the origins and expansion of agricultural practices
in the Mediterranean Basin (Delplancke et al. 2012).
Another unknown in the history of almond cultivation, is the place of origin of
domestication itself. Some evidence suggests that the most plausible spatio-temporal
origin of Prunus dulcis domestication occurred in the Fertile Crescent, and dis-
persed across present day Israel, Palestine and Lebanon up through Syria and east-
ern Turkey and across to Iraq (Albala 2009), during the first half of the Holocene,
~5000 years ago (Browicz and Zohary 1996; Delplancke 2011; Velasco et al. 2016;
Willcox et al. 2009; Zohary et al. 2012).
1.3 Almond Production
With a world value approaching USD 2.30 billion in 2016, almond has become the
largest specialty crop export in the USA and the largest agricultural export by value
(USD 4532 million) of California (ABC 2016).
California accounted for over 80% of the global production of 929,864 mt in
2017 on over 380,405 ha. Australia is now the second largest producer of almond
kernels with 79,461 mt and only 39,662 ha (29,358 ha bearing, 6758 ha non-
bearing, 3546 ha new plantings). Spain, is the third largest producer but with the
largest area, at over 544,518 ha, under cultivation. The rest of the world’s almond
production comes from about 20 countries, mainly Tunisia, Morocco, Italy,
Turkey, Chile and Iran. The Balkan Peninsula including areas of Bulgaria and
Romania; Mexico and Hungary also have limited almond production. Additional
areas exist in central and southwestern Asia including Syria, Iraq, Israel, Ukraine,
Tajikistan, Uzbekistan, Afghanistan and Pakistan, extending into western China
(ABA 2017) (Table 1.1).
The main grown cultivar in California is Nonpareil, with a cultivation area of
131,622 ha, representing 38% of the total area cultivated and producing 40.7% of
the total production. Other important cultivars are Monterey, Butte and Carmel with
cultivation areas of 46,085 ha, 32,810 ha and 30,495 ha, respectively. In addition,
the Spanish cultivar Marcona is cultivated in only 60 ha in California. This cultivar
is the best-evaluated traditional cultivar in Spain recognized by growers and market
for its excellent kernel quality.
Almond kernels can be eaten in their natural state or processed. They have an
excellent taste, crunchy texture and good visual appeal and support diverse food
uses. Kernels may be roasted dry or in oil followed by salting with various season-
ings and used as an ingredient in many manufactured food products. The processed
kernel is usually blanched, which removes the pellicle (skin) using hot water or
steam. For pastry, ice cream, breakfast cereals and vegetable mixtures use, almond
kernels can be sliced or diced. The kernels are important ingredients in bakery
products and, ground into a paste, in the production of marzipan. Several factors
such as cultivar, moisture content, processing and handling procedures can impact
the flavor and texture of almonds (Kester and Gradziel 1996).
Almond cultivation in the Mediterranean countries is mostly non-irrigated, with
kernel yields around 400–500 kg/ha, much lower than the 2300 kg/ha of the USA,
where the production is much more intensive and exclusively irrigated (http://fao-
stat.fao.org). The Mediterranean almond-producing areas play an important role in
social strengthening and retention of families in the area, thus contributing to reduce
Table 1.1 Global almond Country Production (mt)

kernel production in 2017
California (USA) 929,864
Australia 79,461
Spain 50,954
Tunisia 15,000
Iran 15,000
Turkey 13,000
Chile 14,000
Morocco 11,000
Italy 7500
Greece 3000
Others 30,000
Source: ABA (2017)
emigration. However, in spite of the higher oil quality and nutraceutical content
found in the cultivars grown in the Mediterranean area, almond culture is affected
by the low rainfall and drought-limited production.
The Spanish Agriculture Services reported that the summer 2014 drought
affected 72,829 ha (18% of cultivated area) in non-irrigated areas, considering only
the Murcia Region. Damage was estimated by agricultural organizations at around
EUR 123.5 million and more than 2 million trees died. The situation was even more
severe in Morocco and Tunisia. Data reported from Tunisia indicated that during the
last major drought period, 2000 to 2003, more than 20 million trees and 200,000 ha
(80% of the cultivated area) were seriously affected. In these areas of the
Mediterranean Basin, water has become a scarce resource due to high consumption
and the high level of overall pollution. Additionally, the increasing vulnerability to
inclement weather and to long periods of drought has led to depopulation of many
rural areas. Therefore, the implementation of appropriate measures is absolutely
necessary to protect affected farms, their trade and income, as well as that of asso-
ciations, to ensure their independence through collective management. Beyond the
ethical imperative of doing research on how to increase production and to strengthen
the adaptation of plants to drought on a global basis, there is also a need to provide
a beneficial solution to the Mediterranean countries affected by spatial and temporal
complexity of regional droughts (Gouveia et al. 2017). Moreover, climate change
conditions accentuate drought since a reduction of 10% in precipitation translates
into 25% loss of soil water. In almond growing, water availability is a major con-
straint in the cultivation of the species.
1.4 Genetic Diversity and Conservation
Although, the wild almond progenitor is unknown, a significant reduction in genetic

diversity relative to wild progenitor species (once the wild progenitor is confirmed)
should be expected as a consequence of the process of domestication, which is attrib-
uted to a genetic bottleneck. However, in almond this domestication bottleneck and
the following expansion in population size seems unlikely (Meyer and Purugganan
2013; Velasco et al. 2016). With the use of population genetics approaches and after
the evaluation of genome-wide diversity patterns, results have shown no evidence of
a bottleneck during the domestication of almond (Velasco et al. 2016). Additionally,
the authors observed that some regions showed signatures of selection during domes-
tication and also candidate regions presented some overlap between almond and
peach (having a divergence time ~8 million years ago). The lack of a genetic bottle-
neck has also been observed in other perennial crops such as apple and grape (Gross
et al. 2014; Zhou et al. 2017), and was associated with the life history of perennials
exploited for domestication (Gaut et al. 2015). Interestingly, Velasco et al. (2016)
observed that fruit traits were not preferentially targeted during domestication but
likely selected much earlier during species (peach and almond) divergence.
The outcrossing nature of Prunus dulcis, due to its self-incompatible origin, has
contributed to maintain a high level of genetic variability in cultivated almond.
Early studies observed a limited genetic base in commercial cultivars; the authors
considered the use of a few founder genotypes selected for their desirable regional
value could be the main reason of the low variability (Socias i Company and Felipe
1992). This can be the case of most commercially-important California cultivars
originated from crosses between only two parents: cvs. Nonpareil and Mission
(Bartolozzi et al. 1998; Hauagge et al. 1987; Kester and Gradziel 1996). Also, in
Europe the main source of self-compatibility used by breeders has been the almond
cv. Tuono, which could limit the genetic base present in new commercial cultivars.
In addition, only a limited number of accessions are currently maintained
(Appendix II). However, more recently, Szikriszt et al. (2011) found no correla-
tions between the origin of the germplasm and the level of diversity. According to
these authors, the actual germplasm has not suffered any genetic erosion after
inbreeding, since the self-compatibility is a new trait only present in almond
breeding for the last few decades.
A complete genetic characterization of the whole almond germplasm should be
considered by the research community to increase information about the historical
gene pool and to assure its useful protection and utilization for future breeding.
Advances in DNA sequencing technology now permits a more precise analysis of
large collections. Traditionally, markers such as simple-sequence repeats (SSRs)
have been used to address all these questions (Fernández í Martí et al. 2009;
Martínez-Gómez et al. 2003; Xie et al. 2006). More recently, different markers such
as inter-retrotransposon amplified polymorphism (IRAP), sequence-specific ampli-
fication polymorphism (S-SAP), retrotransposon-microsatellite amplified polymor-
phism (REMAP) and inter simple sequence repeats (ISSR), have been used to
genotype large number of accessions of several species. In addition, high resolution
melting (HRM) analysis was developed to detect SNPs in wild almond species
based on publicly available Prunus ESTs and next-generation sequencing (NGS)
data (Sorkheh et al. 2017). An increment of the deployment of genome-wide genetic
diversity using high-throughput technologies and NGS data will provide new tools
to add significant value to germplasm collections.
1.5 Classical Almond Breeding
Classical almond breeding can be described as a long and tedious process, similar
to the improvement processes of the majority of fruit trees. Specifically, its pluren-
nial woody character, long juvenile period and multiplication by grafting as a stan-
dard has clearly controlled the selection process. In this sense, breeders must have a
rigorous and extensive set of data for the specific design of new cultivars (Fig. 1.2).
New cultivars will have an impact in the agricultural system around 12 years
from the time they were first selected, which is the average time to release a new
almond variety (Fig. 1.2). One of the fundamental keys of this process is the
Fig. 1.2 Scheme of an almond breeding program based on the realization of crosses and later
selection. (Source: Figure created by Pedro Martínez-Gómez)
s election of the genotypes that should be used as genitors. The basis of the design is
the realization of crosses and, therefore, it is necessary to choose well which culti-
vars must be crossed to generate new cultivars. These crosses can be of the comple-
mentary type, when two cultivars are crossed with complementary characteristics to
obtain a new cultivar that assembles the interesting phenotypic traits of both geni-
tors, or of the transgressive type, where two cultivars with good phenotypic traits are
crossed to obtain offspring exhibiting a more extreme phenotype than either parent
(Martínez-Gómez et al. 2003).
One important issue, from the economic point of view, is the cost of maintaining
a large number of offspring populations, something that must be considered by the
breeder at each crossing period. This population number is directly related to the
cost in the process of evaluation of the hybrids.
In addition, to develop suitable new cultivars, we need to know the relevant vari-
ables that influence our knowledge of the possible future in the success of new
cultivars. It is thus necessary to examine the endogenous (internal) or exogenous
(external) variables influencing the phenomenon in question. Both types of vari-
ables must be subjected to scientific evaluation (Fig. 1.3).
Fig. 1.3 Internal and external variables affecting the viability and establishment of a new almond
cultivar obtained through classical breeding. (Source: Figure created by Pedro Martínez-Gómez)
The first internal variables to consider in any almond breeding program are
derived from plant traits that are considered as objectives. The most important char-
acteristics in these breeding programs are as follows (Martínez-Gómez et al. 2003):
(a) Tree: Floral self-compatibility, time of flowering and maturation, productivity
and resistance to pests and diseases (Ortega and Dicenta 2003; Sánchez-Pérez
et al. 2004).
(b) Seed (kernel) quality: Organoleptic quality of the seed size, shape, flavor, or
hardness of the shell (Socias i Company et al. 2007).
The knowledge of these internal variables of genetic type will direct their suit-
ability with respect to the proposed design objectives. Prunus species, such as
almond, are characterized by great interspecific compatibility, meaning it is rela-
tively easy to perform interspecific crosses. In addition, one of the main handicaps
of Prunus breeding is the reduced variability of most species derived from their
mating system, being less polymorphic self-compatible species such as peach or
apricot in comparison with self-incompatible such as almond and cherry (Graziel
and Martínez-Gómez 2013). In this context of reduced genetic variability, the real-
ization of interspecific crosses is very useful for the development of new cultivars
(Gradziel et al. 2001; Martínez-Gómez et al. 2004; Sorkheh et al. 2009).
Secondly, internal variables also include the current methodologies available for
use in the selection of individuals. These methodologies are closely related to the
level of knowledge available at the meso- and micro-levels, especially in relation to
the development of molecular markers for the selection of individuals (Gradziel et al.
2001; Martínez-Gómez et al. 2003) and will provide an idea of the efficacy and fea-
sibility of new designs or cultivars. Finally, we can include various economic factors
within the production framework, i.e. the goals, processes and outcomes. The effec-
tiveness and viability of the improvement program will depend on these methodologi-
cal developments as applied to the evaluation. These two terms (effectiveness and
viability) are related in the economic context to the relationship between science and
economics. The evaluation of this feasibility, effectiveness and feasibility can lead to
the abandonment of the new design if it is not feasible or to its production. We would
have to proceed with the design of a new variety with new crosses and new variables
to evaluate within the program. In any case, the evaluation process is continuous.
On the other hand, in almond breeding, it is necessary to consider a number of
external variables including interactions with the environment over a period of sev-
eral years. Many characteristics such as flowering time and floral compatibility
must be evaluated for at least 3 years, when trees produce the first flowers. Due to
drawbacks, the use of molecular marker-assisted selection methods is of particular
interest in breeding programs. Environmental conditions will determine the feasi-
bility of the objectives and the new cultivar in a given location. The adaptation of
new designs or new cultivars to specific climatic conditions is fundamentally condi-
tioned by their winter cold needs for dormancy (Campoy et al. 2011) and by their
adaptation to different soil conditions in the case of rootstocks. Their capacity for
production depends largely on this character and on their interaction with the envi-
ronment considered as an external variable.
Almond bloom in response to an established pattern of low (cold needs) and high
(heat needs) temperatures after the breaking of winter dormancy (Sánchez-Pérez
et al. 2014). These requirements of cold and heat temperatures guarantee that in each
zone the flowering will take place at a favorable time for pollination. Pollination will
be viable in a context of floral self-compatibility (fertilization of the ovule by pollen
from the same variety occurs) (Gradziel et al. 2002; Gradziel and Martínez-Gómez
2013), whereas in the context of floral self-incompatibility flowering will require
pollen of another variety and an isolated variety will not produce fruit (Sánchez-
Pérez et al. 2004). Therefore, late flowering cultivars are sought to avoid frost but are
self-compatible in the case of the almond tree (Sánchez-Pérez et al. 2014). In other
fruit trees such as peach or apricot trees where flowering is not as early, what is
sought is the precocity in flowering and fruiting to obtain better market prices.
The knowledge of these external variables related to possible stresses will indi-
cate their feasibility with respect to the proposed design objectives. The feasibility
of developments is another important issue to consider. The technological feasibil-
ity will also depend on the existence of germplasm with the desired genes for resis-
tance to different biotic or abiotic stresses.
Satisfying consumer demands is also an important external variable. New designs
must have the appropriate characteristics to meet such demands. In addition, the
availability of different techniques for analysis and selection is another important
external variable. In addition, the social acceptance of these new cultivars will
depend on criteria of environmental sustainability. On the other hand, we can
include various factors of an economic nature, provided that they are considered
within the framework of production: the phase of objectives, processes and results.
This affects external economic variables, which are those that affect their market
value. Because the result of a new genetic cultivar presents a durability (a life cycle)
and involves a market price, which conditions the creative transformation carried
out by technological innovation.
The degree of knowledge of the variables determines the quality of the prediction in
the design of new almond cultivars, in addition to their effectiveness, feasibility, suit-
ability and feasibility. New designs must provide the appropriate characteristics to meet
those requirements. External variables include the availability of the different analysis
and the selection contemporary techniques, since they have varied throughout history.
Making predictions in Prunus breeding also involves human activity that is geared
towards reaching feasible goals in the field. The processes to be carried out must have
a reasonable cost, in terms of the level of effort for the realization of the design and
the means to be used. The result is to create a new cultivar, which requires scientific
knowledge and scientific processes. However, the real impact for society comes with
the technology that fosters this innovation through objectives and a task, which origi-
nate in a new cultivar that arises from transforming one’s own reality and making it
have a value in the market. Thus, the scientific, technological and economic rationali-
ties interact. Economic mediation directly affects the human activity deployed.
Under new climatic and consumer conditions, an integrated approach to evaluat-
ing more drought resistant and productive cultivars with enhanced nutraceutical
quality, is of great interest in almond breeding.
1.6 Mutation Breeding
Almond mutation breeding is not common as far as we know. However the use of
spontaneous mutants in almond breeding has been performed. One example is the
development of a late-blooming almond through classical breeding. Martínez-
Gómez et al. (2017) concluded that the adaptation of almonds from the Mediterranean
Basin to colder regions in northern Europe and North America has been mainly
achieved through delayed flowering. These adapted late-flowering cultivars have
usually been developed by selecting desired quantitative genes within each region.
The use of molecular markers for the early selection of genes conferring late flower-
ing has also facilitated the development of additional late cultivars including ultra-
late cultivars, flowering as late as April, together with the combination of material
from different gene pools. These ultra-late flowering cultivars also include genetic
material from spontaneous late-flowering mutations.
1.7 Drought Resistance
Climate variability and water availability require the rapid development of produc-
tion systems able to cope with risk and uncertainty. In this sense, drought resistance
in almond, linked to the efficient water use (Yadollahi et al. 2011), together with the
ability of the root system to access water, are important breeding targets.
Rusticity and flexibility of the different components of the production systems
(including cultivars and rootstocks) should be improved. Clearly, the plantation and
management of new sustainable agro-systems in the Mediterranean Basin for
almond production, designed to last for long periods, must consider the impact of
climatic conditions when selecting new cultivars and rootstocks. In this sense,
drought is one of the biggest problems for non-irrigated culture across the
Mediterranean countries. Studies about drought resistance have been conducted on
wild and cultivated almonds (Camposeo et al. 2011; Palasciano et al. 2014) but these
are scarce in the case of native germplasm from other areas. Some of these studies
conducted by different research groups in Morocco, Tunisia and Spain provided
preliminary results on drought resistance of almond cultivars and hybrid rootstocks
(Alarcón et al. 2002; Esmaeli et al. 2017) and also the discovery of new genetic
diversity from these regions (Gouta et al. 2012; Kodad and Socias i Company 2008).
Under drought conditions, plant develop strategies to cope with stress (accelerat-
ing the life cycle) or to avoid it (controlling stomatal conductance, investing in the
development of the root system, reducing canopy, etc.) or activate osmotic adjust-
ments to increase tolerance to low tissue water potential (for instance accumulating
compatible solutes). The efficiency of photosynthetic carbon gain relative to the rate
of water loss can be used as an indicator (Ennaheli and Earl 2005; Jiménez et al.
2013). In the seasonally dry and variable environment of the Mediterranean region,
the ability of species like almond to cope with water scarcity is not only dependent
on the cultivar, but also very dependent on the rootstock on which it is grafted.
The characterization of drought resistance in almond cultivars and other fruit tree
crops is linked to efficient water use (Yadollahi et al. 2011), together with the ability
of the root system to access water. Collecting breeding material for future research
provides helpful tools to reduce drought losses, but this is a lengthy process (Neale
et al. 2017). However, development of improved production systems using drought
resistant almonds may be possible utilizing native germplasm. These materials
allow a more sustainable production, particularly in the marginal areas of harsh
climate conditions found around the Mediterranean Basin (Gouta et al. 2010, 2011,
2019). This preliminary material has proven to be more efficient and resilient than
wild species that, moreover, show poorer agronomical behavior (Sorkheh et al.
2009). However, an integrated approach evaluating new materials, having good
yield, better nutraceutical quality and highly resistant to drought has not yet been
developed as far as we know.
1.8 Nutraceutical Values
Among cultivated nuts, almonds are one of the most nutritive (Kendall et al. 2003).
They are considered a good source of essential fatty acids, vitamins and minerals
(Table 1.2) (Saura-Calixto et al. 1981). Also, one of the best natural sources of vita-
min E (Sabate and Haddad 2001) which is purported to play a role in preventing
heart disease, certain kinds of cancer and cataract formation (Kodad et al. 2006).
One ounce of almonds (20–25 kernels) contains 15% of the recommended
daily values of phosphorus, 37% of vitamin E and 21% of magnesium. Almonds
also represent a convenient source of fiber and folic acid. Historical uses of sweet
and/or bitter almond ointment include the treatment of asthma, male-pattern bald-
ness and as a soothing salve for burns (Gradziel and Martínez-Gómez 2013).
Almond kernels are also a source of high-quality oil, representing over 50% of
the kernel dry weight. This oil is primarily composed of the more stable oleic acid,
making it desirable from ancient times to the present for use as a base for various
ointments and pharmaceuticals. The high level of this monounsaturated fat in
almond kernels may be partly responsible for the observed association between
frequent nut consumption and purported reduced risk of coronary heart disease
(Fulgoni et al. 2002; Lovejoy et al. 2002).
The almond is appreciated in the processed food industry and also as a func-
tional food with both nutritional and medical (nutraceutical) properties including
nutrients, vitamins, healthy blood lipids and purported anti-inflammatory and
hypocholesterolemic properties (Kodad and Socias i Company 2008; Kodad
Table 1.2 Nutrient Nutrient Value

composition of the almond
Energy 578 kcal
kernel per 100 g fresh weight
of edible portion Protein 21.26 g
Carbohydrate 19.74 g
Fiber, total dietary 11.8 g
Glucose 4.54 g
Starch 0.73 g
Calcium 248 mg
Magnesium 275 mg
Phosphorus 474 mg
Potassium 728 mg
Sodium 1 mg
Folate, total 29 mcg
Vitamin E 25.87 mg
Saturated fatty acids 3.88 g
Monounsaturated fatty acids 32.16 g
Polyunsaturated fatty acid 12.21 g
Source: Adapted from Socias i Company
et al. (2007)
et al. 2011; Musa-Velasco et al. 2016; Poonam et al. 2011). The implementation
of methodologies to phenotype the nutraceutical properties of traditional almond
cultivars, new cultivars and breeding populations are new goals in almond breed-
ing programs. Understanding the influence of diverse stresses such as drought
and heat that can modify the composition of the beneficial components in kernels
is an important goal in almond breeding. New strategies combining food science
knowledge and breeding approaches must be incorporated into the actual breed-
ing programs to obtain better food for future generations (Kodad 2017).
1.9 Development of DNA Markers
The genetic mapping of populations to segregate characters of interest has been the
principal approach to develop marker-assisted selection (MAS) strategies in almond.
MAS allows early selection of a large number of plants, which then undergo final
evaluation to certify the desirable traits using traditional methods. For instance, the
accurate phenotyping of the parents and offspring is key to identifying the molecu-
lar markers linked to disease resistance.
The analysis of cosegregation among markers and characters allows isolating the
map position of major genes and quantitative trait loci (QTL) responsible for their
expression. Genetic linkage analysis was initially performed in almond by combin-
ing different molecular makers including RFLPs, CGs (candidate genes), SSRs and
SNPs. Regarding the genetic structure of almond mapping populations, the typi-
cally assayed populations were of type F1 or F2. These different population types
have advantages and disadvantages. The work developed in F1 populations is more
extensive because the populations are easier to develop than F2, given the longer
period of juvenile growth. F2 populations are common in interspecific crosses in the
case of peach and almond species (Donoso et al. 2016).
During the last two decades, many QTL mapping studies have been carried out.
The genetic dissection of the most important agronomic traits has resulted in the
identification of several different QTLs across the almond genome (Ballester et al.
1998, 2001; Donoso et al. 2016; Fernández í Martí et al. 2011, 2013; Font i Forcada
et al. 2012; Rasouli et al. 2014; Sánchez-Pérez et al. 2007, 2010, 2012) (Fig. 1.4;
Table 1.3). Although, in general, the QTL are distributed in all the linkage groups
(LGs), a high number of QTL can be observed in LG1 of almond, with LG8 having
the lowest number of QTL.
The application of these results in marker-assisted selection has been developed
for blooming time and kernel flavor (Ricciardi et al. 2018; Sánchez-Pérez et al.
2010). In the case of blooming date, different publications report the use of SSR
markers in a F1 population between a seedling of cvs. Tardy Nonpareil (R1000) ×
Desmayo Largueta (R×D), and also confirmed the location of late blooming gene
(Lb) in G4 while identifying other QTLs for flowering time in G1 and G6 (Rasouli
et al. 2014; Sánchez-Pérez et al. 2007) (Fig. 1.4; Table 1.3). In these studies, the SSR
UDP-96003 was located very close to the Lb gene in G4 linkage group. When QTL
Fig. 1.4 Almond QTLs linked to traits of interest: blooming date (Bd), chilling requirements
(CR), heat requirements (HR), productivity (P), self-incompatibility (S), double kernels (Dk), ker-
nel length (Ln), kernel length/width (Ln/Wn), shell hardness (D), kernel size (Sz), spherical index
(Sin), kernel thickness (Tn), kernel thickness/length (Tn/Ln), kernel weight (Wgn), kernel width
(Wn), leafing date (Lf), ripening time (rp), kernel geometric diameter (GDn), amygdalin hydrolase
(AH), Glucosyl transferase (GT), kernel taste (bitterness/sweet) (Sk), linoleic acid (Linoleic),
mandelonitrile lyase (MDL), palmitic acid (palmitic), palmitoleic acid (palmitoleic), oil seed con-
tent (Oil), oleic acid (Oleic), prunasin hydrolase (PH), stearic acid (Stearic), tocopherol homologes
(T-), total seed protein (Protein), germination date (GD), juiciness (jui) and almond fruit type (alf).
A tentative scale of the map is performed in cM using as framework bin map of reference in Prunus
indicating each bin in red. Between branches the references, 1: Ballester et al. (1998); 2: Ballester
et al. (2001); 3: Silva et al. (2005); 4: Sánchez-Pérez et al. (2007); 5: Sánchez-Pérez et al. (2010);
6: Fernández í Martí et al. (2011); 7: Font i Forcada et al. (2012); 8: Sánchez-Pérez et al. (2012);
9: Fernández í Martí et al. (2013); 10: Rasouli et al. (2014); 11: Donoso et al. (2016). (Source:
Figure created by Pedro Martínez-Gómez)
analysis was performed, this major QTL (Lb) in G4 was able to explain 50.0–86.3%
of the variance. In addition, SSR markers PCeGA025, BPPCT037 and EPDCU2584
explained most of the observed variability in kernel flavor, being highly valuable for
marker-assisted selection (Sánchez-Pérez et al. 2010). In addition, the markers
linked to juiciness and almond-peach fruit type are also starting to be applied in
breeding programs (Donoso et al. 2016). In addition, MAS for self-(in)compatibility
is being performed in the routine basis in almond breeding programs using specific
markers derived from transcriptomic studies, by sequencing RNAse genes associ-
ated to this trait (Sánchez-Pérez et al. 2004; Ushijima et al. 1998, 2003).
On the other hand, other traits including in-shell weight, leafing time, shell
hardness, kernel length, nut width, nut length, kernel length, palmitoleic acid or
tocopherol homologues were described as linked to specific molecular markers
(Fig. 1.4), although until now the application of these results is still limited.
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sense or memory, but these can never be the objects of action or of
voluntary pursuit, which must, by the supposition, have an eye to
future events. But with respect to these the chain of self-interest is
dissolved and falls in pieces by the very necessity of our nature, and
our obligations to self as a blind, mechanical, unsociable principle
are lost in the general law which binds us to the pursuit of good as it
comes within our reach and knowledge.
A. A most lame and impotent conclusion, I must say. Do you mean
to affirm that you have really the same interest in another’s welfare
that you have in your own?
B. I do not wish to assert any thing without proof. Will you tell me
if you have this particular interest in yourself what faculty is it that
gives it you—to what conjuration and what mighty magic it is owing
—or whether it is merely the name of self that is to be considered as a
proof of all the absurdities and impossibilities that can be drawn
from it?
A. I do not see that you have hitherto pointed out any.
B. What! not the impossibility that you should be another being,
with whom you have not a particle of fellow-feeling?
A. Another being! Yes, I know it is always impossible for me to be
another being.
B. Ay, or yourself either, without such a fellow-feeling, for it is that
which constitutes self. If not, explain to me what you mean by self.
But it is more convenient for you to let that magical sound lie
involved in the obscurity of prejudice and language. You will please
to take notice that it is not I who commence these hairbreadth
distinctions and special-pleading. I take the old ground of common
sense and natural feeling, and maintain that though in a popular,
practical sense mankind are strongly swayed by self-interest, yet in
the same ordinary sense they are also governed by motives of good-
nature, compassion, friendship, virtue, honour, &c. Now all this is
denied by your modern metaphysicians, who would reduce every
thing to abstract self-interest, and exclude every other mixed motive
or social tie in a strict, philosophical sense. They would drive me
from my ground by scholastic subtleties and newfangled phrases; am
I to blame then if I take them at their word, and try to foil them at
their own weapons? Either stick to the unpretending jog-trot notions
on the subject, or if you are determined to refine in analysing words
and arguments, do not be angry if I follow the example set me, or
even go a little farther to arrive at the truth. Shall we proceed on this
understanding?
A. As you please.
B. We have got so far then (if I mistake not, and if there is not
some flaw in the argument which I am unable to detect) that the past
and present (which alone can appeal to our selfish faculties) are not
the objects of action, and that the future (which can alone be the
object of practical pursuit) has no particular claim or hold upon self.
All action, all passion, all morality and self-interest, is prospective.
A. You have not made that point quite clear. What then is meant
by a present interest, by the gratification of the present moment, as
opposed to a future one?
B. Nothing, in a strict sense; or rather in common speech, you
mean a near one, the interest of the next moment, the next hour, the
next day, the next year, as it happens.
A. What! would you have me believe that I snatch my hand out of
the flame of a candle from a calculation of future consequences?
D. (laughing.) A. had better not meddle with that question. B. is in
his element there. It is his old and favourite illustration.
B. Do you not snatch your hand out of the fire to procure ease from
pain?
A. No doubt, I do.
B. And is not this case subsequent to the act, and the act itself to
the feeling of pain, which caused it?
A. It may be so; but the interval is so slight that we are not sensible
of it.
B. Nature is nicer in her distinctions than we. Thus you could not
lift the food to your mouth, but upon the same principle. The viands
are indeed tempting, but if it were the sight or smell of these alone
that attracted you, you would remain satisfied with them. But you
use means to ends, neither of which exist till you employ or produce
them, and which would never exist if the understanding which
foresees them did not run on before the actual objects and purvey to
appetite. If you say it is habit, it is partly so; but that habit would
never have been formed, were it not for the connection between
cause and effect, which always takes place in the order of time, or of
what Hume calls antecedents and consequents.
A. I confess I think this a mighty microscopic way of looking at the
subject.
B. Yet you object equally to more vague and sweeping generalities.
Let me, however, endeavour to draw the knot a little tighter, as it has
a considerable weight to bear—no less, in my opinion, than the whole
world of moral sentiments. All voluntary action must relate to the
future: but the future can only exist or influence the mind as an
object of imagination and forethought; therefore the motive to
voluntary action, to all that we seek or shun, must be in all cases
ideal and problematical. The thing itself which is an object of pursuit
can never co-exist with the motives which make it an object of
pursuit. No one will say that the past can be an object either of
prevention or pursuit. It may be a subject of involuntary regrets, or
may give rise to the starts and flaws of passion; but we cannot set
about seriously recalling or altering it. Neither can that which at
present exists, or is an object of sensation, be at the same time an
object of action or of volition, since if it is, no volition or exertion of
mine can for the instant make it to be other than it is. I can make it
cease to be indeed, but this relates to the future, to the supposed
non-existence of the object, and not to its actual impression on me.
For a thing to be willed, it must necessarily not be. Over my past and
present impressions my will has no control: they are placed,
according to the poet, beyond the reach of fate, much more of human
means. In order that I may take an effectual and consistent interest
in any thing, that it may be an object of hope or fear, of desire or
dread, it must be a thing still to come, a thing still in doubt,
depending on circumstances and the means used to bring about or
avert it. It is my will that determines its existence or the contrary
(otherwise there would be no use in troubling oneself about it); it
does not itself lay its peremptory, inexorable mandates on my will.
For it is as yet (and must be in order to be the rational object of a
moment’s deliberation) a nonentity, a possibility merely and it is
plain that nothing can be the cause of nothing. That which is not,
cannot act, much less can it act mechanically, physically, all-
powerfully. So far is it from being true that a real and practical
interest in any thing are convertible terms, that a practical interest
can never by any possible chance be a real one, that is, excited by the
presence of a real object or by mechanical sympathy. I cannot
assuredly be induced by a present object to take means to make it
exist—it can be no more than present to me—or if it is past, it is too
late to think of recovering the occasion or preventing it now. But the
future, the future is all our own; or rather it belongs equally to
others. The world of action then, of business or pleasure, of self-love
or benevolence, is not made up of solid materials, moved by
downright, solid springs; it is essentially a void, an unreal mockery,
both in regard to ourselves and others, except as it is filled up,
animated, and set in motion by human thoughts and purposes. The
ingredients of passion, action, and properly of interest are never
positive, palpable matters-of-fact, concrete existences, but
symbolical representations of events lodged in the bosom of futurity,
and teaching us, by timely anticipation and watchful zeal, to build up
the fabric of our own or others’ future weal.
A. Do we not sometimes plot their woe with at least equal good-
will?
B. Not much oftener than we are accessory to our own.
A. I must say that savours more to me of an antithesis than of an
answer.
B. For once, be it so.
A. But surely there is a difference between a real and an imaginary
interest? A history is not a romance.
B. Yes; but in this sense the feelings and interests of others are in
the end as real, as much matters of fact as mine or yours can be. The
history of the world is not a romance, though you and I have had
only a small share in it. You would turn every thing into
autobiography. The interests of others are no more chimerical,
visionary, fantastic than my own, being founded in truth, and both
are brought home to my bosom in the same way by the force of
imagination and sympathy.
D. But in addition to all this sympathy that you make such a rout
about, it is I who am to feel a real, downright interest in my own
future good, and I shall feel no such interest in another person’s.
Does not this make a wide, nay a total difference in the case? Am I to
have no more affection for my own flesh and blood than for
another’s?
B. This would indeed make an entire difference in the case, if your
interest in your own good were founded in your affection for
yourself, and not your affection for yourself in your attachment to
your own good. If you were attached to your own good merely
because it was yours, I do not see why you should not be equally
attached to your own ill—both are equally yours! Your own person or
that of others would, I take it, be alike indifferent to you, but for the
degree of sympathy you have with the feelings of either. Take away
the sense or apprehension of pleasure and pain, and you would care
no more about yourself than you do about the hair of your head or
the paring of your nails, the parting with which gives you no sensible
uneasiness at the time or on after-reflection.
D. But up to the present moment you allow that I have a particular
interest in my proper self. Where then am I to stop, or how draw the
line between my real and my imaginary identity?
B. The line is drawn for you by the nature of things. Or if the
difference between reality and imagination is so small that you
cannot perceive it, it only shows the strength of the latter. Certain it
is that we can no more anticipate our future being than we change
places with another individual, except in an ideal and figurative
sense. But it is just as impossible that I should have an actual
sensation of and interest in my future feelings as that I should have
an actual sensation of and interest in what another feels at the
present instant. An essential and irreconcileable difference in our
primary faculties forbids it. The future, were it the next moment,
were it an object nearest and dearest to our hearts, is a dull blank,
opaque, impervious to sense as an object close to the eye of the blind,
did not the ray of reason and reflection enlighten it. We can never
say to its fleeting, painted essence, ‘Come, let me clutch thee!’ it is a
thing of air, a phantom that flies before us, and we follow it, and with
respect to all but our past and present sensations, which are no
longer any thing to action, we totter on the brink of nothing. That self
which we project before us into it, that we make our proxy or
representative, and empower to embody, and transmit back to us all
our real, substantial interests before they have had an existence,
except in our imaginations, is but a shadow of ourselves, a bundle of
habits, passions, and prejudices, a body that falls in pieces at the
touch of reason or the approach of inquiry. It is true, we do build up
such an imaginary self, and a proportionable interest in it; we clothe
it with the associations of the past and present, we disguise it in the
drapery of language, we add to it the strength of passion and the
warmth of affection, till we at length come to class our whole
existence under one head, and fancy our future history a solid,
permanent, and actual continuation of our immediate being, but all
this only proves the force of imagination and habit to build up such a
structure on a merely partial foundation, and does not alter the true
nature and distinction of things. On the same foundation are built up
nearly as high natural affection, friendship, the love of country, of
religion, &c. But of this presently. What shows that the doctrine of
self-interest, however high it may rear its head, or however
impregnable it may seem to attack, is a mere ‘contradiction,’
‘In terms a fallacy, in fact a fiction,’
is this single consideration, that we never know what is to happen to

us beforehand, no, not even for a moment, and that we cannot so
much as tell whether we shall be alive a year, a month, or a day
hence. We have no presentiment of what awaits us, making us feel
the future in the instant. Indeed such an insight into futurity would
be inconsistent with itself, or we must become mere passive
instruments in the hands of fate. A house may fall on my head as I go
from this, I may be crushed to pieces by a carriage running over me,
or I may receive a piece of news that is death to my hopes before
another four-and-twenty hours are passed over, and yet I feel
nothing of the blow that is thus to stagger and stun me. I laugh and
am well. I have no warning given me either of the course or the
consequence (in truth if I had, I should, if possible, avoid it). This
continued self-interest that watches over all my concerns alike, past,
present, and future, and concentrates them all in one powerful and
invariable principle of action, is useless here, leaves me at a loss at
my greatest need, is torpid, silent, dead, and I have no more
consciousness of what so nearly affects me, and no more care about
it, (till I find out my danger by other and natural means,) than if no
such thing were ever to happen, or were to happen to the Man in the
Moon.
‘And coming events cast their shadows before.’
This beautiful line is not verified in the ordinary prose of life. That
it is not, is a staggering consideration for your fine, practical,
instinctive, abstracted, comprehensive, uniform principle of self-
interest. Don’t you think so, D——?
D. I shall not answer you. Am I to give up my existence for an idle
sophism? You heap riddle upon riddle; but I am mystery-proof. I still
feel my personal identity as I do the chair I sit on, though I am
enveloped in a cloud of smoke and words. Let me have your answer
to a plain question.—Suppose I were actually to see a coach coming
along and I was in danger of being run over, what I want to know is,
should I not try to save myself sooner than any other person?
B. No, you would first try to save a sister, if she were with you.
A. Surely that would be a very rare instance of self, though I do not
deny it.
B. I do not think so. I believe there is hardly any one who does not
prefer some one to themselves. For example, let us look into
Waverley.
A. Ay, that is the way that you take your ideas of philosophy, from
novels and romances, as if they were sound evidence.
B. If my conclusions are as true to nature as my premises, I shall
be satisfied. Here is the passage I was going to quote: ‘I was only
ganging to say, my lord,’ said Evan, in what he meant to be an
insinuating manner, ‘that if your excellent honour and the
honourable court would let Vich Ian Vohr go free just this once and
let him gae back to France and not trouble King George’s
government again, that any six o’ the very best of his clan will be
willing to be justified in his stead; and if you’ll just let me gae down
to Glennaquoich, I’ll fetch them up to ye myself to head or hang, and
you may begin with me the very first man.’[22]
A. But such instances as this are the effect of habit and strong
prejudice. We can hardly argue from so barbarous a state of society.
B. Excuse me there. I contend that our preference of ourselves is
just as much the effect of habit, and very frequently a more
unaccountable and unreasonable one than any other.
A. I should like to hear how you can possibly make that out.
B. If you will not condemn me before you hear what I have to say, I
will try. You allow that D——, in the case we have been talking of,
would perhaps run a little risk for you or me; but if it were a perfect
stranger, he would get out of the way as fast as his legs would carry
him, and leave the stranger to shift for himself.
A. Yes; and does not that overturn your whole theory?
B. It would if my theory were as devoid of common sense as you
are pleased to suppose; that is, if because I deny an original and
absolute distinction in nature (where there is no such thing,) it
followed that I must deny that circumstances, intimacy, habit,
knowledge, or a variety of incidental causes could have any influence
on our affections and actions. My inference is just the contrary. For
would you not say that D—— cared little about the stranger for this
plain reason, that he knew nothing about him?
A. No doubt.
B. And he would care rather more about you and me, because he
knows more about us?
A. Why yes, it would seem so.
B. And he would care still more about a sister, (according to the
same supposition) because he would be still better acquainted with
her, and had been more constantly with her?
A. I will not deny it.
B. And it is on the same principle (generally speaking) that a man
cares most of all about himself, because he knows more about
himself than about any body else, that he is more in the secret of his
own most intimate thoughts and feelings, and more in the habit of
providing for his own wants and wishes, which he can anticipate with
greater liveliness and certainty than those of others, from being more
nearly ‘made and moulded of things past.’ The poetical fiction is
rendered easier and assisted by my acquaintance with myself, just as
it is by the ties of kindred or habits of friendly intercourse. There is
no farther approach made to the doctrines of self-love and personal
identity.
D. E., here is B. trying to persuade me I am not myself.
E. Sometimes you are not.
D. But he says that I never am.—Or is it only that I am not to be
so?
B. Nay, I hope ‘thou art to continue, thou naughty varlet’—
‘Here and hereafter, if the last may be!’
You have been yourself (nobody like you) for the last forty years of
your life: you would not prematurely stuff the next twenty into the
account, till you have had them fairly out?
D. Not for the world, I have too great an affection for them.
B. Yet I think you would have less if you did not look forward to
pass them among old books, old friends, old haunts. If you were cut
off from all these, you would be less anxious about what was left of
yourself.
D. I would rather be the Wandering Jew than not be at all.
B. Or you would not be the person I always took you for.
D. Does not this willingness to be the Wandering Jew rather than
nobody, seem to indicate that there is an abstract attachment to self,
to the bare idea of existence, independently of circumstances or
habit?
B. It must be a very loose and straggling one. You mix up some of
your old recollections and favourite notions with your self elect, and
indulge them in your new character, or you would trouble yourself
very little about it. If you do not come in in some shape or other, it is
merely saying that you would be sorry if the Wandering Jew were to
disappear from the earth, however strictly he may have hitherto
maintained his incognito.
D. There is something in that; and as well as I remember there is a
curious but exceedingly mystical illustration of this point in an
original Essay of yours which I have read and spoken to you about.
B. I believe there is; but A—— is tired of making objections, and I
of answering them to no purpose.
D. I have the book in the closet, and if you like, we will turn to the
place. It is after that burst of enthusiastic recollection (the only one
in the book) that Southey said at the time was something between
the manner of Milton’s prose-works and Jeremy Taylor.
B. Ah! I as little thought then that I should ever be set down as a
florid prose-writer as that he would become poet-laureat!
J. D. here took the volume from his brother, and read the following
passage from it.
‘I do not think I should illustrate the foregoing reasoning so well
by any thing I could add on the subject, as by relating the manner in
which it first struck me. There are moments in the life of a solitary
thinker which are to him what the evening of some great victory is to
the conqueror and hero—milder triumphs long remembered with
truer and deeper delight. And though the shouts of multitudes do not
hail his success—though gay trophies, though the sounds of music,
the glittering of armour, and the neighing of steeds do not mingle
with his joy, yet shall he not want monuments and witnesses of his
glory—the deep forest, the willowy brook, the gathering clouds of
winter, or the silent gloom of his own chamber, “faithful
remembrancers of his high endeavour, and his glad success,” that, as
time passes by him with unreturning wing, still awaken the
consciousness of a spirit patient, indefatigable in the search of truth,
and the hope of surviving in the thoughts and minds of other men. I
remember I had been reading a speech which Mirabaud (the author
of the “System of Nature”) has put into the mouth of a supposed
Atheist at the last judgment; and was afterwards led on, by some
means or other to consider the question, whether it could properly be
said to be an act of virtue in any one to sacrifice his own final
happiness to that of any other person or number of persons, if it were
possible for the one ever to be made the price of the other? Suppose
it were my own case—that it were in my power to save twenty other
persons by voluntarily consenting to suffer for them: Why should I
not do a generous thing, and never trouble myself about what might
be the consequence to myself the Lord knows when?
‘The reason why a man should prefer his own future welfare to that
of others is, that he has a necessary, absolute interest in the one,
which he cannot have in the other—and this, again, is a consequence
of his being always the same individual, of his continued identity
with himself. The difference, I thought, was this, that however
insensible I may be to my own interest at any future period, yet when
the time comes I shall feel differently about it. I shall then judge of it
from the actual impression of the object, that is, truly and certainly;
and as I shall still be conscious of my past feelings, and shall bitterly
regret my own folly and insensibility, I ought, as a rational agent, to
be determined now by what I shall then wish I had done, when I shall
feel the consequences of my actions most deeply and sensibly. It is
this continued consciousness of my own feelings which gives me an
immediate interest in whatever relates to my future welfare, and
makes me at all times accountable to myself for my own conduct. As,
therefore, this consciousness will be renewed in me after death, if I
exist again at all—But stop—as I must be conscious of my past
feelings to be myself, and as this conscious being will be myself, how
if that consciousness should be transferred to some other being?
How am I to know that I am not imposed upon by a false claim of
identity? But that is ridiculous, because you will have no other self
than that which arises from this very consciousness. Why, then, this
self may be multiplied in as many different beings as the Deity may
think proper to endue with the same consciousness; which, if it can
be renewed at will in any one instance, may clearly be so in a
hundred others. Am I to regard all these as equally myself? Am I
equally interested in the fate of all? Or if I must fix upon some one of
them in particular as my representative and other self, how am I to
be determined in my choice? Here, then, I saw an end put to my
speculations about absolute self-interest and personal identity. I saw
plainly that the consciousness of my own feelings, which is made the
foundation of my continued interest in them, could not extend to
what had never been, and might never be; that my identity with
myself must be confined to the connection between my past and
present being; that with respect to my future feelings or interests,
they could have no communication with, or influence over, my
present feelings and interests, merely because they were future; that
I shall be hereafter affected by the recollection of my past feelings
and actions; and my remorse be equally heightened by reflecting on
my past folly and late-earned wisdom, whether I am really the same
being, or have only the same consciousness renewed in me; but that
to suppose that this remorse can re-act in the reverse order on my
present feelings, or give me an immediate interest in my future
feelings, before they exist, is an express contradiction in terms. It can
only affect me as an imaginary idea, or an idea of truth. But so may
the interests of others; and the question proposed was, whether I
have not some real, necessary, absolute interest in whatever relates
to my future being, in consequence of my immediate connection with
myself—independently of the general impression which all positive
ideas have on my mind. How, then, can this pretended unity of
consciousness which is only reflected from the past—which makes
me so little acquainted with the future that I cannot even tell for a
moment how long it will be continued, whether it will be entirely
interrupted by or renewed in me after death, and which might be
multiplied in I don’t know how many different beings, and prolonged
by complicated sufferings, without my being any the wiser for it,—
how, I say, can a principle of this sort identify my present with my
future interests, and make me as much a participator in what does
not at all affect me as if it were actually impressed on my senses? It is
plain, as this conscious being may be decompounded, entirely
destroyed, renewed again, or multiplied in a great number of beings,
and as, whichever of these takes place, it cannot produce the least
alteration in my present being—that what I am does not depend on
what I am to be, and that there is no communication between my
future interests, and the motives by which my present conduct must
be governed. This can no more be influenced by what may be my
future feelings with respect to it, than it will then be possible for me
to alter my past conduct by wishing that I had acted differently. I
cannot, therefore, have a principle of active self-interest arising out
of the immediate connection between my present and future self, for
no such connection exists, or is possible. I am what I am in spite of
the future. My feelings, actions, and interests, must be determined by
causes already existing and acting, and are absolutely independent of
the future. Where there is not an intercommunity of feelings, there
can be no identity of interests. My personal interest in any thing
must refer either to the interest excited by the actual impression of
the object, which cannot be felt before it exists, and can last no
longer than while the impression lasts; or it may refer to the
particular manner in which I am mechanically affected by the idea of
my own impressions in the absence of the object. I can, therefore,
have no proper personal interest in my future impressions, since
neither my ideas of future objects, nor my feelings with respect to
them, can be excited either directly or indirectly by the impressions
themselves, or by any ideas or feelings accompanying them, without
a complete transposition of the order in which causes and effects
follow one another in nature. The only reason for my preferring my
future interest to that of others, must arise from my anticipating it
with greater warmth of present imagination. It is this greater
liveliness and force with which I can enter into my future feelings,
that in a manner identifies them with my present being; and this
notion of identity being once formed, the mind makes use of it to
strengthen its habitual propensity, by giving to personal motives a
reality and absolute truth which they can never have. Hence it has
been inferred that my real, substantial interest in any thing, must be
derived in some indirect manner from the impression of the object
itself, as if that could have any sort of communication with my
present feelings, or excite any interest in my mind but by means of
the imagination, which is naturally affected in a certain manner by
the prospect of future good or evil.’
J. D. ‘This is the strangest tale that e’er I heard,
C. D. ‘It is the strangest fellow, brother John!’
THE FREE ADMISSION
The New Monthly Magazine.]

[July, 1830.
A free Admission is the lotos of the mind: the leaf in which your
name is inscribed as having the privileges of the entrée for the season
is of an oblivious quality—an antidote for half the ills of life. I speak
here not of a purchased but of a gift-ticket, an emanation of the
generosity of the Managers, a token of conscious desert. With the
first you can hardly bring yourself to go to the theatre; with the last,
you cannot keep away. If you have paid five guineas for a free
admission for the season, this free admission turns to a mere slavery.
You seem to have done a foolish thing, and to have committed an
extravagance under the plea of economy. You are struck with
remorse. You are impressed with a conviction that pleasure is not to
be bought. You have paid for your privilege in the lump, and you
receive the benefit in driblets. The five pounds you are out of pocket
does not meet with an adequate compensation the first night, or on
any single occasion—you must come again, and use double diligence
to strike a balance to make up your large arrears; instead of an
obvious saving, it hangs as a dead-weight on your satisfaction all the
year; and the improvident price you have paid for them kills every
ephemeral enjoyment, and poisons the flattering illusions of the
scene. You have incurred a debt, and must go every night to redeem
it; and as you do not like being tied to the oar, or making a toil of a
pleasure, you stay away altogether; give up the promised luxury as a
bad speculation; sit sullenly at home, or bend your loitering feet in
any other direction; and putting up with the first loss, resolve never
to be guilty of the like folly again. But it is not thus with the possessor
of a Free Admission, truly so called. His is a pure pleasure, a clear
gain. He feels none of these irksome qualms and misgivings. He
marches to the theatre like a favoured lover; if he is compelled to
absent himself, he feels all the impatience and compunction of a
prisoner. The portal of the Temple of the Muses stands wide open to
him, closing the vista of the day—when he turns his back upon it at
night with steps gradual and slow, mingled with the common crowd,
but conscious of a virtue which they have not, he says, ‘I shall come
again to-morrow!’ In passing through the streets, he casts a sidelong,
careless glance at the playbills: he reads the papers chiefly with a
view to see what is the play for the following day, or the ensuing
week. If it is something new, he is glad; if it is old, he is resigned—but
he goes in either case. His steps bend mechanically that way—
pleasure becomes a habit, and habit a duty—he fulfils his destiny—he
walks deliberately along Long-acre (you may tell a man going to the
play, and whether he pays or has a free admission)—quickens his
pace as he turns the corner of Bow-street, and arrives breathless and
in haste at the welcome spot, where on presenting himself, he
receives a passport that is a release from care, thought, toil, for the
evening, and wafts him into the regions of the blest! What is it to him
how the world turns round if the play goes on; whether empires rise
or fall, so that Covent Garden stands its ground? Shall he plunge into
the void of politics, that volcano burnt-out with the cold, sterile,
sightless lava, hardening all around? or con over the registers of
births, deaths, and marriages, when he may be present at Juliet’s
wedding, and gaze on Juliet’s tomb? or shall he wonder at the throng
of coaches in Regent-street, when he can feast his eyes with the
coach (the fairy-vision of his childhood) in which Cinderella rides to
the ball? Here (by the help of that Open Sessame! a Free Admission),
ensconced in his favourite niche, looking from the ‘loop-holes of
retreat’ in the second circle, he views the pageant of the world played
before him; melts down years to moments; sees human life, like a
gaudy shadow, glance across the stage; and here tastes of all earth’s
bliss, the sweet without the bitter, the honey without the sting, and
plucks ambrosial fruits and amaranthine flowers (placed by the
enchantress Fancy within his reach,) without having to pay a tax for
it at the time, or repenting of it afterwards. ‘He is all ear and eye, and
drinks in sounds or sights that might create a soul under the ribs of
death.’ ‘The fly,’ says Gay, ‘that sips treacle, is lost in the sweets’: so
he that has a free admission forgets every thing else. Why not? It is
the chief and enviable transfer of his being from the real to the
unreal world, and the changing half of his life into a dream. ‘Oh!
leave me to my repose,’ in my beloved corner at Covent Garden
Theatre! This (and not ‘the arm-chair at an inn,’ though that too, at
other times, and under different circumstances, is not without its
charms,) is to me ‘the throne of felicity.’ If I have business that would
detain me from this, I put it off till the morrow; if I have friends that
call in just at the moment, let them go away under pain of bearing
my maledictions with them. What is there in their conversation to
atone to me for the loss of one quarter of an hour at the ‘witching
time of night?’ If it is on indifferent subjects, it is flat and insipid; if it
grows animated and interesting, it requires a painful effort, and
begets a feverish excitement. But let me once reach, and fairly
establish myself in this favourite seat, and I can bid a gay defiance to
mischance, and leave debts and duns, friends and foes, objections
and arguments, far behind me. I would, if I could, have it surrounded
with a balustrade of gold, for it has been to me a palace of delight.
There golden thoughts unbidden betide me, and golden visions come
to me. There the dance, the laugh, the song, the scenic deception
greet me; there are wafted Shakspear’s winged words, or Otway’s
plaintive lines; and there how often have I heard young Kemble’s
voice, trembling at its own beauty, and prolonging its liquid tones,
like the murmur of the billowy surge on sounding shores! There I no
longer torture a sentence or strain a paradox: the mind is full without
an effort, pleased without asking why. It inhales an atmosphere of
joy, and is steeped in all the luxury of woe. To show how much
sympathy has to do with the effect, let us suppose any one to have a
free admission to the rehearsals of a morning, what mortal would
make use of it? One might as well be at the bottom of a well, or at the
top of St. Paul’s for any pleasure we should derive from the finest
tragedy or comedy. No, a play is nothing without an audience, it is a
satisfaction too great and too general not to be shared with others.
But reverse this cold and comfortless picture—let the eager crowd
beset the theatre-doors ‘like bees in spring-time, when the sun with
Taurus rides’—let the boxes be filled with innocence and beauty like
beds of lilies on the first night of Isabella or Belvidera, see the flutter,
the uneasy delight of expectation, see the big tear roll down the cheek
of sensibility as the story proceeds—let us listen to the deep thunder
of the pit, or catch the gallery’s shout at some true master-stroke of
passion; and we feel that a thousand hearts are beating in our
bosoms, and hail the sparkling illusion reflected in a thousand eyes.
The stage has, therefore, been justly styled ‘a discipline of humanity’;
for there is no place where the social principle is called forth with
such strength and harmony, by a powerful interest in a common
object. A crowd is everywhere else oppressive; but the fuller the play-
house, the more intimately and cordially do we sympathise with
every individual in it. Empty benches have as bad an effect on the
spectator as on the players. This is one reason why so many mistakes
are made with respect to plays and players, ere they come before the
public. The taste is crude and uninformed till it is ripened by the
blaze of lighted lamps and the sunshine of happy faces: the cold,
critical faculty, the judgment of Managers and Committees asks the
glow of sympathy and the buzz of approbation to prompt and guide
it. We judge in a crowd with the sense and feelings of others; and
from the very strength of the impression, fancy we should have come
to the same unavoidable conclusion had we been left entirely to
ourselves. Let any one try the experiment by reading a manuscript
play, or seeing it acted—or by hearing a candidate for the stage
rehearse behind the scenes, or top his part after the orchestra have
performed their fatal prelude. Nor is the air of a play-house
favourable only to social feeling—it aids the indulgence of solitary
musing. The brimming cup of joy or sorrow is full; but it runs over to
other thoughts and subjects. We can there (nowhere better) ‘retire,
the world shut out, our thoughts call home.’ We hear the revelry and
the shout, but ‘the still, small voice’ of other years and cherished
recollections is not wanting. It is pleasant to hear Miss Ford repeat
Love’s Catechism, or Mrs. Humby[23] sing ‘I cannot marry Crout’: but
the ear is not therefore deaf to Mrs. Jordan’s laugh in Nell; Mrs.
Goodall’s Rosalind still haunts the glades of Arden, and the echo of
Amiens’ song, ‘Blow, blow, thou winter’s wind,’ lingers through a
lapse of thirty years. A pantomime (the Little Red Riding-Hood)
recalls the innocence of our childish thoughts: a dance (the Minuet
de la Cour) throws us back to the gorgeous days of Louis XIV. and tells
us that the age of chivalry is gone for ever. Who will be the Mrs.
Siddons of a distant age? What future Kean shall ‘strut and fret his
hour upon the stage,’ full of genius and free from errors? What
favourite actor or actress will be taking their farewell benefit a
hundred years hence? What plays and what players will then amuse
the town? Oh, many-coloured scenes of human life! where are ye
more truly represented than in the mirror of the stage? or where is
that eternal principle of vicissitude which rules over ye, the painted
pageant and the sudden gloom, more strikingly exemplified than
here? At the entrance to our great theatres, in large capitals over the
front of the stage, might be written Mutability! Does not the curtain
that falls each night on the pomps and vanities it was withdrawn
awhile to reveal (and the next moment all is dark) afford a fine moral
lesson? Here, in small room, is crowded the map of human life; the
lengthened, varied scroll is unfolded like rich tapestry with its quaint
and flaunting devices spread out; whatever can be saved from the
giddy whirl of ever-rolling time and of this round orb, which moves
on and never stops,[24] all that can strike the sense, can touch the
heart, can stir up laughter or call tears from their secret source, is
here treasured up and displayed ostentatiously—here is Fancy’s
motley wardrobe, the masks of all the characters that were ever
played—here is a glass set up clear and large enough to show us our
own features and those of all mankind—here, in this enchanted
mirror, are represented, not darkly, but in vivid hues and bold relief,
the struggle of Life and Death, the momentary pause between the
cradle and the grave, with charming hopes and fears, terror and pity
in a thousand modes, strange and ghastly apparitions, the events of
history, the fictions of poetry (warm from the heart); all these, and
more than can be numbered in my feeble page, fill that airy space
where the green curtain rises, and haunt it with evanescent shapes
and indescribable yearnings.
‘See o’er the stage the ghost of Hamlet stalks,
Othello rages, Desdemona mourns,
And poor Monimia pours her soul in love.’
Who can collect into one audible pulsation the thoughts and
feelings that in the course of his life all these together have
occasioned; or what heart, if it could recall them at once, and in their
undiminished power and plenitude, would not burst with the load?
Let not the style be deemed exaggerated, but tame and creeping, that
attempts to do justice to this high and pregnant theme, and let tears
blot out the unequal lines that the pen traces! Quaffing these
delights, inhaling this atmosphere, brooding over these visions, this
long trail of glory, is the possessor of a Free Admission to be blamed
if ‘he takes his ease’ at the play; and turning theatrical recluse, and
forgetful of himself and his friends, devotes himself to the study of
the drama, and to dreams of the past? By constant habit (having
nothing to do, little else to think of), he becomes a tippler of the dews
of Castaly—a dram-drinker on Mount Parnassus. He tastes the
present moment, while a rich sea of pleasure presses to his lip and
engulfs him round. The noise, the glare, the warmth, the company,
produce a sort of listless intoxication, and clothe the pathos and the
wit with a bodily sense. There is a weight, a closeness even, in the air,
that makes it difficult to breathe out of it. The custom of going to the
play night after night becomes a relief, a craving, a necessity—one
cannot do without it. To sit alone is intolerable, to be in company is
worse; we are attracted with pleasing force to the spot where ‘all that
mighty heart is beating still.’ It is not that perhaps there is any thing
new or fine to see—if there is, we attend to it—but at any time, it kills
time and saves the trouble of thinking. O, Covent Garden! ‘thy
freedom hath made me effeminate!’ It has hardly left me power to
write this description of it. I am become its slave, I have no other
sense or interest left. There I sit and lose the hours I live beneath the
sky, without the power to stir, without any determination to stay.
‘Teddy the Tiler’ is become familiar to me, and, as it were, a part of
my existence: ‘Robert the Devil’ has cast his spell over me. I have
seen both thirty times at least, (no offence to the Management!) and
could sit them out thirty times more. I am bed-rid in the lap of
luxury; am grown callous and inert with perpetual excitement.
——‘What avails from iron chains
Exempt, if rosy fetters bind as fast?’
I have my favourite box too, as Beau Brummell had his favourite

leg; one must decide on something, not to be always deciding.
Perhaps I may have my reasons too—perhaps into the box next to
mine a Grace enters; perhaps from thence an air divine breathes a
glance (of heaven’s own brightness), kindles contagious fire;—but let
us turn all such thoughts into the lobbies. These may be considered
as an Arabesque border round the inclosed tablet of human life. If
the Muses reign within, Venus sports heedless, but not unheeded
without. Here a bevy of fair damsels, richly clad, knit with the Graces
and the Hours in dance, lead on ‘the frozen winter and the pleasant
spring!’ Would I were allowed to attempt a list of some of them, and
Cowley’s Gallery would blush at mine! But this is a licence which
only poetry, and not even a Free Admission can give. I can now
understand the attachment to a player’s life, and how impossible it is
for those who are once engaged in it ever to wean themselves from it.
If the merely witnessing the bustle and the splendour of the scene as
an idle spectator creates such a fascination, and flings such a charm
over it, how much more must this be the case with those who have
given all their time and attention to it—who regard it as the sole
means of distinction—with whom even the monotony and
mortifications must please—and who, instead of being passive,
casual votaries, are the dispensers of the bounty of the gods, and the
high-priests at the altar?

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Advances in Plant Breeding Strategies

Nut and Beverage Crops Volume 4

Advances in Plant Breeding Strategies Industrial and

Advances in Plant Breeding Strategies Cereals Volume 5

Advances in Plant Breeding Strategies Legumes Volume 7

Plant Breeding Reviews: Volume 40 1st Edition Jules

Plant breeding reviews: Volume 43 1st Edition Irwin

Plant Breeding Reviews: Volume 41 1st Edition Irwin

Advances in Plant Ecophysiology Techniques Adela M.

Current Technologies in Plant Molecular Breeding A

Advances in Plant Breeding

ISBN 978-3-030-23111-8 ISBN 978-3-030-23112-5 (eBook)

© Springer Nature Switzerland AG 2019

Contemporary plant breeders no longer need to rely solely on traditional method-

Al-Hassa, Saudi Arabia Jameel M. Al-Khayri

Part I Nut Crops

9 Advances in Edible Pine Nut Trees (Pinus spp.)

Part II Beverage Crops

Jameel M. Al-Khayri is a Professor of Plant

Legislative Council) for the 2009–2012 term. Currently,

Shri Mohan Jain is a Consultant and Plant

Dennis V. Johnson is a Consultant and Former

Naima Ait Aabd National Institute for Agricultural Research (INRA)-CRRA,

Rachid Bouharroud National Institute for Agricultural Research (INRA)-CRRA,

Pedro Martínez-Gómez CEBAS-CSIC, Murcia, Spain

Sunil Samal College of Horticulture, Chiplima, Odisha University of Agriculture

Pedro J. Martínez-García, Angela S. Prudencio, Raquel Sánchez-Pérez,

Keywords Almond · Breeding · Epigenetic marks · Genomics · Molecular

P. J. Martínez-García (*) · A. S. Prudencio · R. Sánchez-Pérez · F. P. De Los Cobos

© Springer Nature Switzerland AG 2019 3

1.2 Origin of Cultivation

A wide dissemination of cultivated almond is notable today; several authors have

1.3 Almond Production

Table 1.1 Global almond Country Production (mt)

1.4 Genetic Diversity and Conservation

Although, the wild almond progenitor is unknown, a significant reduction in genetic

1.5 Classical Almond Breeding

1.6 Mutation Breeding

1.7 Drought Resistance

1.8 Nutraceutical Values

Table 1.2 Nutrient Nutrient Value

1.9 Development of DNA Markers

is this single consideration, that we never know what is to happen to

The New Monthly Magazine.]

I have my favourite box too, as Beau Brummell had his favourite

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Al-Hassa, Saudi Arabia Jameel M. Al-Khayri

Part I Nut Crops

9 Advances in Edible Pine Nut Trees (Pinus spp.)

Part II Beverage Crops

1.2 Origin of Cultivation

1.3 Almond Production

1.4 Genetic Diversity and Conservation

1.5 Classical Almond Breeding

1.6 Mutation Breeding

1.7 Drought Resistance

1.8 Nutraceutical Values

1.9 Development of DNA Markers