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Jameel M. Al-Khayri · Shri Mohan Jain
Dennis V. Johnson Editors
Advances in Plant
Breeding Strategies:
Industrial and Food
Crops
Volume 6
Advances in Plant Breeding Strategies: Industrial
and Food Crops
Jameel M. Al-Khayri • Shri Mohan Jain
Dennis V. Johnson
Editors
Advances in Plant Breeding

Strategies: Industrial
and Food Crops
Volume 6
Editors
Jameel M. Al-Khayri Shri Mohan Jain
Department of Agricultural Biotechnology University of Helsinki
King Faisal University Helsinki, Finland
Al-Hassa, Saudi Arabia
Dennis V. Johnson
Cincinnati, OH, USA
ISBN 978-3-030-23264-1 ISBN 978-3-030-23265-8 (eBook)

https://doi.org/10.1007/978-3-030-23265-8
© Springer Nature Switzerland AG 2019

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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 6, subtitled Industrial and Food Crops, focuses on advances in
breeding strategies using both traditional and modern approaches for the improve-
ment of individual crops. Included in Part I are 11 industrial plant species utilized as
sources of raw materials for the production of industrial products including pulp and
wood crops (acacia), fiber (cotton, jute, and ramie), rubber (guayule and rubber
tree), and oil (jojoba and flax). In addition to agave which has industrial uses as a
source of raw materials for hard fiber, biofuels, and pharmaceutical products as well
as uses in food industries supplying raw materials for alcoholic beverages such as
tequila, the main table sugar source (sugarcane) is included. Part II of this volume
covers seven food plants selected for their utilization in food industries for the pro-
duction of chocolate (cacao), cooking oil (oil palm, safflower, sesame, and sun-
flower), and natural flavors (saffron and vanilla).
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 138 figures and 78 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 60 scientists
from 14 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 Industrial Crops

1 Genetics and Breeding of Tropical Acacias for Forest Products:
Acacia mangium, A. auriculiformis and A. crassicarpa �� 3
Arif Nirsatmanto and Sri Sunarti
2 Cotton (Gossypium hirsutum L.) Breeding Strategies�� 29
Saeed Rauf, Muhammad Shehzad, Jameel M. Al-Khayri,
Hafiz Muhammad Imran, and Ijaz Rasool Noorka
3 CRISPR/Cas9: A New Genome Editing Tool to Accelerate
Cotton (Gossypium spp.) Breeding �� 61
Muhammad N. Sattar, Zafar Iqbal, Sarbesh Das Dangol,
and Allah Bakhsh
4 Jute (Corchorus spp.) Breeding�� 85
Liwu Zhang, Aminu Kurawa Ibrahim, Sylvain Niyitanga,
Liemei Zhang, and Jianmin Qi
5 Ramie (Boehmeria nivea L. Gaud) Genetic Improvement �� 115
Pratik Satya, Sabyasachi Mitra, and Deb Prasad Ray
6 Genetic Improvement of Guayule (Parthenium argentatum
A. Gray): An Alternative Rubber Crop �� 151
Hussein Abdel-Haleem, Zinan Luo, and Dennis Ray
7 Biotechnological Advances in Rubber Tree (Hevea brasiliensis
Muell. Arg.) Breeding�� 179
Sankaran Sobha, Karumamkandathil Rekha, and Thomas K. Uthup
8 Genetics and Breeding of Jojoba [Simmondsia chinensis
(Link) Schneider] �� 237
Mohamed M. A. Khairi
vii
viii Contents
9 Flax (Linum usitatissimum L.) Genomics and Breeding�� 277

Frank M. You, Sylvie Cloutier, Khalid Y. Rashid,
and Scott D. Duguid
10 Breeding Strategies to Improve Production
of Agave (Agave spp.) �� 319
Kelly M. Monja-Mio, Miguel A. Herrera-Alamillo,
Lorenzo F. Sánchez-Teyer, and Manuel L. Robert
11 Sugarcane (Saccharum spp.): Breeding and Genomics�� 363
Shriram J. Mirajkar, Rachayya M. Devarumath, Ashok A. Nikam,
Kapil V. Sushir, Harinath Babu, and Penna Suprasanna
Part II Food Crops

12 Cacao (Theobroma cacao L.) Breeding�� 409
Frances Bekele and Wilbert Phillips-Mora
13 Oil Palm (Elaeis spp.) Breeding in Malaysia �� 489
Fadila Ahmad Malike, Mohd Din Amiruddin, Zulkifli Yaakub,
Marhalil Marjuni, Norziha Abdullah, Nor Azwani Abu Bakar,
Suzana Mustaffa, Mohd Mustakim Mohamad, Mohd Yazid Hassan,
Meilina Ong Abdullah, Ahmad Parveez Ghulam Kadir,
and Ahmad Kushairy Din
14 Safflower (Carthamus tinctorius L.) Breeding �� 537
Pooran Golkar and Somayeh Karimi
15 Advances in Sesame (Sesamum indicum L.) Breeding�� 577
Swapan K. Tripathy, Jayashree Kar, and Dibyabharati Sahu
16 Breeding Strategies for Sunflower (Helianthus annuus L.)
Genetic Improvement�� 637
Saeed Rauf
17 Saffron (Crocus sativus L.) Breeding:
Opportunities and Challenges�� 675
Majid Shokrpour
18 Vanilla (Vanilla spp.) Breeding�� 707
Alan H. Chambers
Index�� 735
Editors and Contributors
About the Editors
Prof. 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 regeneration
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 refereed
international journals and 25 review chapters and
edited 7 journal special issues. In addition, he edited
five reference books on date palm biotechnology and
utilization 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 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.
Prof. 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),
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.
Editors and Contributors xi
Prof. 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 international development agencies provid-
ing 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
Hussein Abdel-Haleem USDA-ARS, ALARC, Maricopa, AZ, USA

Norziha Abdullah Malaysian Palm Oil Board, Advanced Biotechnology and
Breeding Centre (ABBC), Persiaran Institusi, Kajang, Selangor, Malaysia
Meilina Ong Abdullah Malaysian Palm Oil Board, Advanced Biotechnology and
Jameel M. Al-Khayri Department of Agricultural Biotechnology, King Faisal
University, Al-Hassa, Saudi Arabia
Mohd Din Amiruddin Malaysian Palm Oil Board, Advanced Biotechnology and
Harinath Babu Molecular Biology and Genetic Engineering, Vasantdada Sugar
Institute, Pune, Maharashtra, India
Nor Azwani Abu Bakar Malaysian Palm Oil Board, Advanced Biotechnology
and Breeding Centre (ABBC), Persiaran Institusi, Kajang, Selangor, Malaysia
xii Editors and Contributors
Allah Bakhsh Department of Agricultural Genetic Engineering, Faculty of

Agricultural Sciences and Technologies, Nigde Omer Halisdemir University, Nigde,
Turkey
Frances Bekele Cocoa Research Centre (CRC), The University of the West Indies,
St. Augustine, Trinidad and Tobago
Alan H. Chambers Tropical Research and Education Center, Horticultural
Sciences Department, University of Florida, Homestead, FL, USA
Sylvie Cloutier Ottawa Research and Development Centre, Agriculture and Agri-
Food Canada, Ottawa, ON, Canada
Sarbesh Das Dangol Department of Agricultural Genetic Engineering, Faculty of
Agricultural Sciences and Technologies, Nigde Omer Halisdemir University, Nigde,
Turkey
Rachayya M. Devarumath Molecular Biology and Genetic Engineering,
Vasantdada Sugar Institute, Pune, Maharashtra, India
Ahmad Kushairy Din Malaysian Palm Oil Board, Advanced Biotechnology and
Scott D. Duguid Morden Research and Development Centre, Agriculture and
Agri-Food Canada, Morden, MB, Canada
Ahmad Parveez Ghulam Kadir Malaysian Palm Oil Board, Advanced
Biotechnology and Breeding Centre (ABBC), Persiaran Institusi, Kajang, Selangor,
Malaysia
Pooran Golkar Research Institute for Biotechnology and Bioengineering, Isfahan
University of Technology, Isfahan, Iran
Mohd Yazid Hassan Malaysian Palm Oil Board, Advanced Biotechnology and
Miguel A. Herrera-Alamillo Centro de Investigación Científica de Yucatán A.C.,
Mérida, Mexico
Aminu Kurawa Ibrahim Bast Fiber Biology Center, College of Crop Science,
Fujian Agriculture and Forestry University, Fuzhou, China
Hafiz Muhammad Imran Central Cotton Research Institute, Multan, Pakistan
Zafar Iqbal Central Laboratories, King Faisal University, Al-Ahsa, Kingdom of
Saudi Arabia
Jayashree Kar Department of Plant Breeding & Genetics, College of Agriculture,
Orissa University of Agriculture and Technology (OUAT), Bhubaneswar, Odisha,
India
Editors and Contributors xiii
Mohamed M. A. Khairi Faculty of Agriculture, Dongola University, Dongola,

Sudan
Zinan Luo USDA-ARS, ALARC, Maricopa, AZ, USA
Fadila Ahmad Malike Malaysian Palm Oil Board, Advanced Biotechnology and
Marhalil Marjuni Malaysian Palm Oil Board, Advanced Biotechnology and
Shriram J. Mirajkar Department of Biotechnology, Dr. D. Y. Patil Arts,
Commerce and Science College, Pimpri, Pune, India
Sabyasachi Mitra Indian Council of Agricultural Research-Central Research
Institute for Jute and Allied Fibres, Barrackpore, Kolkata, India
Mohd Mustakim Mohamad Malaysian Palm Oil Board, Advanced Biotechnology
and Breeding Centre (ABBC), Persiaran Institusi, Kajang, Selangor, Malaysia
Kelly M. Monja-Mio Centro de Investigación Científica de Yucatán A.C., Mérida,
Mexico
Suzana Mustaffa Malaysian Palm Oil Board, Advanced Biotechnology and
Ashok A. Nikam Plant Tissue Culture Laboratory, Vasantdada Sugar Institute,
Pune, Maharashtra, India
Arif Nirsatmanto Center for Forest Biotechnology and Tree Improvement,
Yogyakarta, Indonesia
Sylvain Niyitanga Bast Fiber Biology Center, College of Crop Science, Fujian
Agriculture and Forestry University, Fuzhou, China
Ijaz Rasool Noorka Department of Plant Breeding and Genetics, College of
Agriculture, University of Sargodha, Sargodha, Punjab, Pakistan
Somayeh Karimi Pajoohan Spadan Research Company, Isfahan Science and
Technology Town, Isfahan, Iran
Wilbert Phillips-Mora Retired, Centro Agronómico Tropical de Investigación y
Enseñanza (Tropical Agricultural Research and Higher Education Center) (CATIE),
Turrialba, Costa Rica
Jianmin Qi Bast Fiber Biology Center, College of Crop Science, Fujian Agriculture
and Forestry University, Fuzhou, China
Khalid Y. Rashid Morden Research and Development Centre, Agriculture and
Agri-Food Canada, Morden, MB, Canada
Saeed Rauf Department of Plant Breeding and Genetics, College of Agriculture,
University of Sargodha, Sargodha, Punjab, Pakistan
xiv Editors and Contributors
Deb Prasad Ray Indian Council of Agricultural Research-National Institute of

Natural Fibre Engineering and Technology, Kolkata, India
Dennis Ray School of Plant Sciences, University of Arizona, Tucson, AZ, USA
Karumamkandathil Rekha Advance Centre for Molecular Biology and
Biotechnology, Rubber Research Institute of India, Rubber Board, Kottayam,
Kerala, India
Manuel L. Robert Centro de Investigación Científica de Yucatán A.C., Mérida,
Mexico
Dibyabharati Sahu Department of Plant Breeding & Genetics, College of
Agriculture, Orissa University of Agriculture and Technology (OUAT),
Bhubaneswar, Odisha, India
Lorenzo F. Sánchez-Teyer Centro de Investigación Científica de Yucatán A.C.,
Mérida, Mexico
Muhammad N. Sattar College of Agriculture and Food Science, King Faisal
University, Al-Ahsa, Kingdom of Saudi Arabia
Pratik Satya Indian Council of Agricultural Research-Central Research Institute
for Jute and Allied Fibres, Barrackpore, Kolkata, India
Muhammad Shehzad Department of Plant Breeding and Genetics, College of
Agriculture, University of Sargodha, Sargodha, Punjab, Pakistan
Majid Shokrpour Department of Horticulture, College of Agriculture and Natural
Resources, University of Tehran, Karaj, Iran
Sankaran Sobha Rubber Research Institute of India (Retd), Kerala, India
Sri Sunarti Center for Forest Biotechnology and Tree Improvement, Yogyakarta,
Indonesia
Penna Suprasanna Nuclear Agriculture and Biotechnology Division, Bhabha
Atomic Research Centre, Mumbai, India
Kapil V. Sushir Sugarcane Breeding Section, Vasantdada Sugar Institute, Pune,
Maharashtra, India
Thomas. K. Uthup Advance Centre for Molecular Biology and Biotechnology,
Rubber Research Institute of India, Rubber Board, Kottayam, Kerala, India
Swapan K. Tripathy Department of Agricultural Biotechnology, College of
Agriculture, Orissa University of Agriculture and Technology (OUAT),
Bhubaneswar, Odisha, India
Zulkifli Yaakub Malaysian Palm Oil Board, Advanced Biotechnology and
Editors and Contributors xv
Frank M. You Ottawa Research and Development Centre, Agriculture and Agri-
Food Canada, Ottawa, ON, Canada
Liemei Zhang Bast Fiber Biology Center, College of Crop Science, Fujian Agriculture
Liwu Zhang Bast Fiber Biology Center, College of Crop Science, Fujian Agriculture
Part I
Industrial Crops
Chapter 1
Genetics and Breeding of Tropical Acacias
for Forest Products: Acacia mangium,
A. auriculiformis and A. crassicarpa
Arif Nirsatmanto and Sri Sunarti
Abstract Industrial forest plantations are playing an important role in the forestry
sector for both economic and environmental reasons. Some fast- growing species of
tropical acacias, such as Acacia mangium, A. auriculiformis, A. crassicarpa and
Acacia hybrids are among the most dominant trees in forest industrial plantations in
Southeast Asia countries: Indonesia, Malaysia, Vietnam, Thailand and Papua New
Guinea. These trees are mainly planted to supply raw materials for pulp and paper,
sawn lumber, the plywood industry and wood energy. Improvement through opti-
mizing of silvicultural practices, exploring genetics and breeding of acacias has
been gradually taking place. Although progress is still slow, as compared to agricul-
tural crops, due to the age at rotation, larger tree size and extensive site management,
some achievements in genetics and breeding of acacias have provided a significant
benefit and impact in the operational scale of forest industries. Genetic diversity of
acacias is low and most breeding is practiced conventionally under a recurrent selec-
tion strategy practiced successively from the first generation to advanced generation
breeding cycles. Recently, breeding of acacias is also being developed through
hybridization, application of biotechnology and molecular breeding.
Keywords Conventional breeding · Tree improvement · Molecular breeding ·

hybridization · Biotechnology · Forest industry
1.1 Introduction
The genus Acacia consists of trees and shrubs and is in the family Leguminosae,
subfamily Mimosoideae. The genus includes more than 1000 species which are
naturally distributed in Africa, the Americas, Asia and Australia (Anonymous 1982).
Mangium (Acacia mangium) auriculiformis (A. auriculiformis) and crassicarpa
A. Nirsatmanto (*) · S. Sunarti

Center for Forest Biotechnology and Tree Improvement, Yogyakarta, Indonesia
e-mail: arifnirsatmanto@biotifor.or.id; narti_nirsatmanto@yahoo.com
© Springer Nature Switzerland AG 2019 3

J. M. Al-Khayri et al. (eds.), Advances in Plant Breeding Strategies:
Industrial and Food Crops, https://doi.org/10.1007/978-3-030-23265-8_1
4 A. Nirsatmanto and S. Sunarti
(A. crassicarpa) are among the most dominant tropical acacias cultivated for the
forest industries in Southeast Asia countries. These three acacias have a natural
distribution in Australia, Papua New Guinea and Papua Province, Indonesia (Gunn
and Midgley 1991; Thomson 1994), usually occurring in overlapping populations.
Natural interspecific hybridization is potentially taking place among the acacias,
especially between A. mangium × A. auriculiformis (Griffin et al. 2015a, b; Kha
2001; Sunarti et al. 2013).
Mangium is a major species for industrial forest plantation growth in the
Southeast Asian countries of Indonesia, Malaysia, Vietnam, Thailand and Papua
New Guinea (Harwood and Nambiar 2013; Turnbull et al. 1997). It was introduced
as an exotic fast-growing tree for forest plantations beginning in the 1960s in Sabah,
followed by Malaysia, Indonesia, Vietnam and Thailand (Pinyopusarerk et al. 1993;
Turnbull et al. 1997). The main purpose of wood production is to supply raw mate-
rial for pulp and paper, plywood, particleboard and sawn lumber. This species is
also widely planted for other forestry purposes, such as reforestation and soil reha-
bilitation, and categorized as a multipurpose species. Recently it was reported that
the total area of mangium plantations is around 1,350,000 ha, mostly in Indonesia,
Malaysia and Vietnam, with the main purpose to supply raw material for the pulp
and paper industry (Harwood and Nambiar 2014). The wood can also be combined
into pulp mixtures including Eucalyptus globulus to obtain enhanced pulp proper-
ties (Wu et al. 2018).
Auriculiformis was introduced earlier than mangium, in the 1930s, particularly
for land rehabilitation (Turnbull et al. 1997). Later it was cultivated for industrial
purposes, mostly to supply raw material for sawn lumber. Due to higher levels of
lignin, wood density and extractive contents in the wood of this species, there is a
reduction in pulp yield (Jahan et al. 2008), making it less desirable for the pulp and
paper industry. However, it has potential for hybridization with A. mangium to pro-
duce the hybrid, A. mangium × A. auriculiformis, which is suitable for pulp and
paper needs (Kha 2001; Sunarti et al. 2013). Most auriculiformis plantations are
managed by small growers using fewer silvicultural practices as compared to man-
gium (Hai et al. 2008; Harwood and Nambiar 2014; Huong et al. 2015). Like man-
gium, auriculiformis is also categorized as a multipurpose species and planted for
reforestation and soil rehabilitation.
Although auriculiformis has had a long history since its introduction, the precise
area of plantations is unclear because this species is mostly planted by small local
growers with uncertain silvicultural practices and harvesting rotation. In Indonesia,
auriculiformis is planted along with other acacia species or mixed with other tree
species and crops; the time of harvest depends on the farmers’ needs without con-
sideration of the fixed age of the trees. Natural regeneration is practiced following
the selective successive harvesting rotation. Unlike in Indonesia, auriculiformis
plantations in Vietnam are better organized and managed in terms of silvicultural
practices and harvesting (Hai et al. 2008; Harwood and Nambiar 2014; Huong
et al. 2015).
1 Genetics and Breeding of Tropical Acacias for Forest Products 5
Crassicarpa was first introduced as an alternative species in the pulp and paper
industry in the 1990s (Turnbull et al. 1997). In drylands, the growth of crassicarpa
is suboptimal as compared to mangium. However, following the enlargement of
plantations into wetlands, crassicarpa was found to be one of the acacias adaptable
to peat soils. It was reported that the area of plantations of this species is around
700,000 ha, mostly in Indonesia (Harwood and Nambiar 2013; Turnbull et al. 1997).
Most acacia plantation areas were established beyond the natural distribution of
the tree species, and thus there were introduced as exotics. Seeds for plantations
were first collected from natural populations, mainly in eastern Indonesia, Papua
New Guinea and northern Australia (Gunn and Midgley 1991). At the time of initial
plantation establishment, stand productivity was low due to improper silvicultural
practices and unselected planting stock. Following the particular importance of
high-productivity acacia plantations, efforts were made to adopt sound silvicultural
practices; breeding was then carried out and has resulted in efficient and effective
plantation management (Griffin et al. 2015a, b; Harwood et al. 2015). Mangium is
the most advanced in terms of silvicultural practices among the three Acacia spe-
cies, followed by crassicarpa and auriculiformis. Intensive and comprehensive
research studies on nursery growth and silvicultural practices, such as cultivation,
plantation management, fertilizing, weeding, pest and disease control, harvesting,
wood properties and manufacturing processes in acacias have been conducted for
20–40 years with varied progress among the three Acacia species.
Breeding is one of the important factors in improving acacia plantations. After
initial introduction and recognition that the acacias have potential for industry, early
improvement through breeding was also pursued. Considering the need to supply
large quantities of planting stock, genetic materials were sought and collected from
a wide range of the natural distribution areas for establishing initial genetic trials,
such as seed stands, provenance stands and first-generation seed orchards, in many
sites representative of plantation regions. Although some advanced generation
breeding programs were recently achieved, application of biotechnology and
molecular breeding remains challenging. The role of certain research institutions
and research projects in developing genetic and breeding is recognized for the four
tropical acacias; such as the Center for Forest Biotechnology and Tree Improvement
(CFBTI) in Indonesia, the Forest Research Institute Malaysia (FRIM), the Institute
of Forest Tree Improvement and Biotechnology (FTIBRI) in Vietnam and the
ACIAR Project on pest and disease of tropical acacias in Indonesia and Vietnam
(ACIAR-FST/2014/068) (see Appendix I).
In this chapter, information on the genetics, achievements and challenges of
breeding Acacia mangium, A. auriculiformis and A. crassicarpa) and the hybrid
(A. mangium × A. auriculiformis) is presented. The chapter provides a history of the
introduced species, cultivation status and breeding, biodiversity, future challenges
in the involvement of biotechnology and molecular breeding.
1.2 Silviculture and Conventional Breeding
1.2.1 Current Cultivation Practices
The four acacias mentioned above are highly adaptable to a wide range of tropical
regions, from sites with adverse conditions of dry and degraded soil, soil with a low
pH and low rainfall, up to sites with good fertile soil and high rainfall (Mackey
1996; Pinyopusarerk et al. 1993; Turnbull et al. 1997). Although the main objective
is to develop forest industries, most acacia plantations were initially established in
degraded soils as part of reforestation and soil rehabilitation programs. Concerning
the importance of high productivity of plantations, considerable efforts were under-
taken to improve cultivation practices by assessing the suitability of site conditions.
Due to the variability of site conditions, cultivation techniques in practice differ
from site to site according to company preferences.
The main objectives typical of industrial acacia plantation in drylands are that
they are planted mostly on degraded lands such as grasslands (Imperata cylindrica)
and to some degree in logged-over areas (secondary forest). In the first rotation,
fully mechanized site preparation is practiced in grassland areas by ploughing, fol-
lowed by harrowing (Otsamo et al. 1996; Siregar et al. 1999). In the case of scru-
bland and logged-over areas, site preparation is through slashing, felling and
chopping. Debris is then piled at the sides of the planting block. For subsequent
rotations, mechanical site preparation is not done. After timber harvest, weeds and
naturally-regenerated acacia seedling are slashed, followed by herbicide application
(Hardiyanto and Nambiar 2014). Unlike in drylands, plantation areas for acacias in
wetlands mostly take place in peat land where the only adaptable species is crassi-
carpa. Peat land drainage is implemented by controlling water levels at around
50–80 cm below the peat surface through a canal system (Tropenbos, APRIL and
Ministry of Forestry 2010).
Planting stock for acacia plantations is mostly provided by raising seedlings
from seed. In mangium, large-scale multiplication using stem cuttings from young
seedlings is also practiced by some companies (Wong and Yuliarto 2014).
Genetically-improved seed for seedling preparation is mostly collected from first-
and second-generation breeding cycles (Harwood et al. 2015; Kurinobu and
Rimbawanto 2002; Nirsatmanto et al. 2014, 2015; Werren 1991). In the A. mangium
× A. auriculiformis hybrid, planting stock is prepared only through the large-scale
multiplication of shoot and stem cuttings from selected clones. Ageing effects com-
monly occur in the multiplication of shoot and cutting for acacias in which the root-
ing ability of a cutting decreases as hedge plants or stool plants get older (Ahmad
1991; Monteuuis et al. 1995). Therefore, maintenance of juvenility is essential to
the success of cuttings in acacias.
Stand density is one of the important factors in the silvicultural practices of aca-
cia plantations. It is determined at the beginning of planting by the spacing arrange-
ment. Considering crown development, growth and stand productivity, the initial
plant spacing for acacias is 3 × 3 m, or a stand density of 1100 trees ha−1, mainly for
the pulp and paper industry (Hardiyanto 2004). Further silvicultural treatments such
as thinning and singling in dryland plantation areas are necessary to produce fine,
straight logs if the wood is destined for sawn lumber (Beadle et al. 2007), plywood
and particleboard.
Some acacias, such as mangium, are nitrogen-fixers (Galiana et al. 1990). This
character will affect the fertilizer application in plantations, especially the levels of
N. The first application of fertilizer when planting acacias is at planting time, com-
monly using TSP (triple superphosphate) or SP-36 (Hardiyanto 2004). Fertilizer is
put into the planting hole and then the surface is covered by soil. This is done before
planting the seedling to avoid direct contact between the seedling roots and the fer-
tilizer. One month after planting, additional fertilizer is applied using NPK which is
allocated in the dip around 10 cm from the planted seedling. In some companies, the
additional fertilizer is also applied 2–3 times within the first 2 years. In the subse-
quent rotation, the levels of N application will be much reduced due to fixed nitro-
gen added to the soil from trees in previous rotations. Weeding should be practiced
periodically to improve early seedling growth by reducing intense weed competi-
tion. Weeding can be manual or by the application of herbicides, and optimally done
before fertilizer application.
In the past, the growth of the three Acacia species was commonly crooked with
multi-stemmed leaders (occurrence of more than one main stem in one tree).
However some recent breeding processes have succeeded in improving the stem
form to be straighter and with a single stem (Harwood et al. 2015). For pulp and
paper, multi-stemmed trees are not a matter of serious concern and thinning is not
necessary.
The harvesting rotation for acacia depends on the purposes of the end-product.
With fast growing tree species for biomass production, such as pulp paper and wood
energy, the length of rotation is shorter, around 2–6 years. In the case of wood-based
products, such as furniture, particle board and other sawn lumber products, the rota-
tion is longer at more than 10 years. In some regions, especially when planted by the
local people, a combination of the purposes of sawn lumber and wood energy is
practiced, in which the main stem is used to furnish logs for sawn lumber while the
trimmings are used for wood energy. Wood processing technology is important for
producing good lumber from acacias, especially with respect to wood drying. This
is because the water content of the wood is high and the lumber will easily split and
crack if improperly dried, especially for younger harvested timber (Tenorio et al.
2012; Yamamoto et al. 2003).
1.2.2 Current Problems and Challenges
There are at least two main problems in plantation development of the four acacias.
The first is the low stand productivity due to substandard silvicultural practices,
particularly under adverse condition either in drylands or wetlands. The second is
the increase of plantation tree damage due to pests and diseases.
Under current silvicultural practices, the productivity of acacia plantations has

been increased in some regions. In addition, breeding practices have also succeeded
in improving plantation productivity (Hardiyanto 2014; Harwood and Nambiar
2014; Nirsatmanto et al. 2015). However, there is still a large gap between supply
and the total demand. In this case, expansion of plantations is necessary. Very large
variations of planting-site conditions means the deployment of high-productivity
acacia plantations is limited, although the key species have been reported to be
highly adaptive to varied soil types. Growth will be more pronounced in a site with
favorable soil conditions and climate, such as well-drained soil properties and
higher annual rainfall (>1800 mm/year) (Mackey 1996). For peat areas, given the
current issues regarding carbon emissions and peat fires, water levels in peatland
drainage should be reassessed for sustainable peatland management. For instance,
Indonesian Government Regulation number 71/2014 on the Protection and
Management of Peat Ecosystems, declares that the water levels can be as high as
40 cm below the peat surface. However, that water level will suppress the growth of
crassicarpa, the main acacia commonly planted in peat areas. This situation has
become one of the most serious problems for crassicarpa plantations. It is necessary
to have the appropriate silvicultural technique to encourage high-productivity aca-
cia plantations in adverse environments. Plant exploration to identify adaptive geno-
types is also an important challenge in acacia breeding.
In the past, there were no serious and significant impacts of pest and disease on
acacias. However, recently acacia plantations are facing very serious threats both
from pests and diseases resulting in very significant damage to plantations, mostly
in mangium (Brawner et al. 2015; Harwood and Nambiar 2014; Harwood et al.
2015; Tarigan et al. 2011). Squirrels and monkeys are pests that seriously threaten
mangium plantations. A large proportion of the plantations in Sumatra Island,
Indonesia are attacked by these pests (Hardiyanto 2014). Diseases are another very
serious threat, mainly in mangium. Root rot associated with Ganoderma sp. and
wilt disease with Ceratocystis sp., cause high mortality in mangium plantations. For
example, Suryantini and Wulandari (2018) studied the virulence level of three
Ganoderma isolates to infect mangium seedlings, which revealed that G. lucidum
isolated from mangium is of highest virulence, compared to G. boninense isolated
from the oil palm and G. applanatum isolated from the rubber tree. Pathogen infec-
tions are commonly associated with tree wounds due to silvicultural practices, such
as branch pruning and singling of multi-stemmed trees, as well as wounds from
pests and wind damage. Exploring for resistant genotypes to such diseases is another
important challenge in acacia breeding.
Acacia plantations for pulp and paper industries are usually monocultural with
limited genetic diversity that is probably more susceptible to the consequences of
climate change (Pawson et al. 2013). Climate change may impact directly on forest
diversity through changes in temperature, rainfall, forest fires, and pest and disease
attacks. Indirectly, climate change will strongly affect plantation forests requiring
changes in forest management practices to mitigate the influences of climate change
on productivity. Furthermore, Pawson et al. (2013) suggested that the management
of forest plantations in future should take into account the plant rotation, weed
c ontrol and silvicultural practices and also established multi-species plantations for
climate change mitigation and adaptation. Mixed plantations of clones, which are
structured on a mosaic with variation in age classes, will be more productive,
healthier and increase the biodiversity (Lindgren 1993), while increasing resilience
to climate change (Pawson et al. 2013). Booth et al. (2014) examined the challenge
on how climate change may affect locations in Southeast Asia suitable for growing
mangium, auriculiformis and crassicarpa by considering a short rotation, with the
relatively frequent opportunity to change planting stock and provenance resources.
Covering a large area, an acacia plantations plays an important role in biological
carbon mitigation through increased concentration of CO2 by sequestering consid-
erable amounts of carbon in biomass and soil (Kalita et al. 2016).
1.2.3 Improvement Strategy
Some improvement strategies are underway to overcome current problems of acacia

plantations. Modified silvicultural practices related to successive second and third
rotation plantations are being applied, particularly to improve conditions through
reducing soil compaction from mechanical harvesting, appropriate fertilizer appli-
cation and land preparation for the next planting. To increase the quantity of wood
production, plantations are being enlarged through the expansion of plantation areas
to include minimally-suited areas of soil fertility, soil drainage and rainfall.
The effort to manipulate genetics through breeding is another strategy to over-
come problems. Although most breeding is still taking place conventionally through
selection in successive advanced generations and hybridization, some involvement
of biotechnology such as polyploid breeding and genomic selection has also been
initiated (Baurens et al. 2004; Griffin et al. 2015a, b; Wang and Cui 2000). All
breeding strategies will be focused on improving genotypes which show higher pro-
ductivity both in growth and wood properties, resistance to pests and diseases and
tolerance to adverse conditions.
1.2.4 Traditional Breeding Methodologies and Limitations
Breeding strategies using a recurrent selection system is regularly practiced in aca-

cia and includes selection, mating and propagation, and repeated in successive gen-
erations of breeding (Namkoong et al. 1988). In each generation, the process is
implemented by establishing a seedling seed orchard for three simultaneous objec-
tives: (1) progeny testing, (2) seed production and (3) a genetic base for successive
generations. The breeding strategy employs open-pollinated progeny from plus
trees. This strategy is adopted due to the early flowering habit at 2–3 years of age,
and the difficulty in vegetative propagation, as well as the limitation of available
breeding techniques using controlled pollination (Wright 1976).
Due to the difficulty of full pedigree control, breeding populations of the four
acacias are managed under multiple populations. In this strategy, a large quantity of
genetic material for breeding is divided into several subpopulations of sublines,
instead of one single large population (Barnes 1984; McKeand and Beineke 1980;
van Buijitenen and Lowe 1979). Sublines are then created by establishing several
small breeding groups, and mating among the parent trees with some degree of
inbreeding is only allowed within the respective subline (McKeand and Beineke
1980). On the other hand, maintaining outbred offsprings will be directed in a seed
production orchards consisting of a combination of a few selected superior seed
parents from a respective subline. To avoid relatedness among sublines and to adjust
the timing of flowering, a sublining system of tested families by provenance is rec-
ommended (Nikles 1989).
Selection for genetic gain in breeding population of acacias can be based on
either phenotypic or genotypic values (Falconer 1981). Regarding the seedling seed
orchards with multiple-tree plots, the genetic gain will be realized within-plot as
well as by family selection (Wright 1976). In spite of the low potential of gain as
compared with that by family selection, roguing the orchard by within-plot selec-
tion is a first step to improve genetic quality of the acacias.
A subsequent selection procedure conducted after the within-plot selection in
seedling seed orchards of acacias is family selection. The primary purpose of this
type of selection is to improve the genetic quality of seed produced in the orchards
by culling trees in the poor families. Unlike the within-plot selections that were
done largely dependent on breeder intuition, the family selection can be regarded as
more controllable, because it is theoretically oriented and straightforward. However,
in practice, due to the low genetic diversity in acacias, something short of modifica-
tion in family selection is desirable to reduce some risk associated with selection
intensity, relationship between selection intensity and genetic variability, and differ-
ent outcrossing rates. After finishing the family selection, seedling seed orchards are
used as orchards for operational seed production.
Beyond the strategy toward advance generation breeding populations using the
sublining system in acacia breeding, selection of plus trees is conducted in each
subline. Selected trees are considered as candidate before designation as plus trees,
and they should be examined not only based on growth and form trait, but also on
other essential traits such as flowering habit, wood property as well as pedigree.
Candidate plus trees should be confirmed on their family origin to reduce the prog-
ress of inbreeding in the next generation, where a maximum of two plus trees per
family are allowed to be tested in the subsequent generation. To maintain wide
genetic diversity in the orchards and to increase genetic gain, new selected plus trees
from other populations within the same provenance region of the acacias can be
added into subsequent generation of the breeding population (Borralho and
Dutkowski 1996). From two successive generational breeding cycles, improved
seed of mangium could result in a genetic gain of stand volume of 30–50% over the
unimproved seed, and the second generation breeding showed 13% higher than the
first generation breeding (Nirsatmanto et al. 2015).
Early selection is often practiced in seedling seed orchard establishments to
increase the genetic gain per unit of time and thus the economic return on tree
improvement activities (Lambeth 1980; McKeand 1988). To evaluate the efficiency
of early selection in acacia seedling seed orchards, assessment of the genetic param-
eters is necessary. This is because genetic gain by this type of early selection is well
known to be primarily dependent on the magnitude of heritability and the genetic
correlation between selection age and rotation age (Cotteril and Dean 1988;
McKeand 1988). Nevertheless, genetic parameters may change markedly as trees
grow and develop (Balocchi et al. 1993; Namkoong and Conkle 1976). Likewise,
for many breeding programs which are concerned with volume improvement, tree
height was found to be good early predictor and selection trait for volume yield at
rotation age of the acacias due to its typically higher heritability and stronger
juvenile-mature correlation (Lambeth et al. 1983). Nirsatmanto et al. (2012)
reported that early selection of mangium based on tree height brought more gain per
year than later direct selection on 8-year rotation with optimal age for early selec-
tion found at 2 years of age.
Besides shortening the breeding cycle, large additional gain in productivity in
acacias also can be achieved through selection of the best provenance for a given
site and purpose. Therefore, acacia seedling seed orchards should be made up using
families of plus trees selected from good provenances which proved to be promising
based on previous provenance trials. This is because the acacias are introduced exot-
ically into plantation areas far from their natural distribution. Hence, available prov-
enance information of the acacias will become an important factor prior in seedling
seed orchard establishment. A study of projected increase in stand volume of intro-
duced provenances of mangium showed that those from Papua New Guinea and
Queensland increased the genetic gain by 17–26% over the local land race
(Nirsatmanto et al. 2014).
Although it has come a long way and made significant achievements in tradi-
tional breeding, work remains in acacia breeding due to their limitations. As
described in preceding paragraphs, traditional breeding in acacias faces difficulty in
controlled crosses among the parents and the ageing effect in propagation. In addi-
tion, breeding was explored mostly through the use of potential genetic variation
which was reported at a low level. This low variation is the most current limitation
in improving acacias through traditional breeding. This situation is aggravated by
the decrease of acacia populations in natural areas due to illegal logging or other
biological and physical threats. In this case, an infusion of genetic material from
new collections into the advanced generation breeding population could not be suc-
cessively practiced to maintain a large genetic variation. As a result, the potential for
increasing productivity through the breeding will stop.
1.2.5 Role of Biotechnology
In tree breeding programs, biotechnology advances include new methods in vegeta-

tive propagation through in vitro, molecular biology through DNA markers and
genomic DNA, polyploidy breeding and genetic transformation (Barrens et al.
2004; Burdon 1994; Griffin et al. 2015a, b). The role of biotechnology is expected
to overcome some limitations and constraints in the breeding of forest tree species,
such as shorter time for selection, maintain the juvenility of stock plants for vegeta-
tive propagation, and give more accurate means for modifying and combining of
target genes (Burdon 1994).
Depending on the end-product uses, economic rotation of acacias can be har-
vested at around 6–10 years of age. Final selection in the breeding process is com-
monly practiced at the midpoint of rotation at around 4–5 years of age. Considering
the genetic information from molecular studies such as quantitative trait loci (QTL),
the selection procedure could be potentially started at early stages in the nursery
(Burdon 1994). However, the time lag to proceed into the subsequent generation
cycles based on the reproductive ability in the acacias is around 2–3 years
(Nirsatmanto et al. 2012). Therefore, complete control of flowering using biotech-
nology, such as through induction of precocious flowering, is necessary to shorten
the time lag. In another case, technology using species-specific SNP markers will
also be useful to confirm acacia hybrid trees and for screening mass propagated
ramets of selected acacia hybrid clones (Asif et al. 2017).
Other biological limitations of acacia breeding include the difficulty in making
controlled cross-pollination due to the typical flowers, which are hermaphroditic
with many tiny flowers in an inflorescence (Ibrahim 1993). An emasculation pro-
cess is needed to make controlled crosses (Griffin et al. 2010). But then, due to the
tiny flowers with polyad pollen, emasculation is difficult, although that does not
mean it cannot be done. Therefore, other alternative methods for gene recombina-
tion are necessary, and hence biotechnology has great potential to be applied in
acacia breeding, such as through genetic transformation, polyploid breeding, and
genome and molecular breeding. In addition, the developed biotechnology could be
used to optimize the low genetic diversity in the breeding of acacias.
Asexual propagation is commonly practiced to rapidly deploy genetically
improved stock obtained from breeding programs. However the ageing effect influ-
ences low rates of success of clonal propagation in acacias (Ahmad 1990; Monteuuis
et al. 1995). This is because material from mature stock plants usually results in a
low level of rooting capability. In this case, maintaining the juvenility of stock
plants for the mass scale clonal propagation is important. Developed technology
through in vitro culture and somatic embryogenesis (SE) can help greatly not only
to maintain the stock plants in juvenile stages, but also to accelerate the increased
number of multiplied planting stock from vegetative propagation (Andrade et al.
2011; Hazubska-Przybyl and Bojarczu 2016).
The technique of tissue culture propagation of the four acacias has been devel-
oped well, and successfully produced plantlets for mass plantations (Ahmad 1991;
Galiana et al. 2003; Monteuuis 2004; Yang et al. 2006). Propagation by tissue
culture of mangium was successfully practiced using nodal explants which were
excised from 1-month old aseptically germinated seedlings and cultured on full
strength MS basal medium (Murashige and Skoog 1962) supplemented with
6-benzylamino purine (BAP) or kinetin (Ahmad 1991). The types of auxin hor-
mones such as indole butyric acid (IBA), alpha-naphthalene acetic acid (NAA)
and Serradix 3 were used for root formation. Multiplication from juvenile material
of mangium showed better rooting ability than the mature genotypes
(Monteuuis 2004).
In vitro multiplication and rooting formation on acacia hybrids requires different
medium as practiced on mangium, additional suitable regulators were necessary in
the culture medium. The average of multiplication and rooting ability of the Acacia
hybrid was higher than mangium but varied greatly among the clones (Galiana et al.
2003). The survival rate during acclimatization in the nursery under a misting sys-
tem was also higher.
The first successful crassicarpa propagation from an excised 2-month-old phyl-
lode in vitro seedling was achieved using MS basal media supplemented with
1-phenyl-3 (thiadiazol-5-yl), thidiazuron and α-naphthalene acetic acid (NAA)
(Yang et al. 2006). The rooting medium from elongated adventitious shoots was MS
media supplemented with 3-indolebutyric acid (IBA). The shoots produced 96.5%
of rooting plantlets which were successfully acclimatized in a nursery bed on soil
media with an 80% survival rate.
1.3 Germplasm Biodiversity and Conservation
1.3.1 Germplasm Diversity
The natural distribution of the three acacias is in northern Australia, southeastern

Papua Province, Indonesia and southwestern Papua New Guinea (Gunn and Midgley
1991; Thomson 1994). The populations are separated into two larger groups on
New Guinea Island and Australia, with some overlapping sites among the three
acacias. The distinct differences in geographic distribution among the two islands is
indicative of the differences in germplasm diversity for the three acacias. Genetic
diversity among the three species is typically similar, and generally categorized as
low, with mangium having the least genetic diversity among the three (Moran et al.
1988, 1989; Widyatmoko et al. 2010).
Among the two distinct natural distributions of mangium, Queensland, Australia
and New Guinea Island (Papua Province and PNG), the Papua population is reported
as having higher genetic diversity as compared to Australia (Butcher et al. 1998). In
Australia, a decline in diversity occurs from populations in the north (Cape York) to
populations farther south in Townsville and Daintree.
The natural distribution of auriculiformis is geographically wider than mangium.

It covers not only populations in New Guinea Island and the northern part of
Queensland, but also Northern Territory, Australia (Turnbull et al. 1997). It is
reported that the populations from Queensland are closely related to the populations
of New Guinea Island, rather than to the Northern Territory (Wickneswari and
Norwati 1993). In addition, it was reported that the genetic diversity of auriculifor-
mis is higher as compared to mangium and crassicarpa (Widyatmoko et al. 2010).
The natural distribution of crassicarpa covers a similar ranges to mangium, which is
New Guinea Island and northern Queensland. It was also reported that the genetic
diversity of crassicarpa is low and comparable mangium.
1.3.2 Species Characterization and Phylogeny
In general, the three acacias are somewhat similar in morphology. However, the dif-
ferences among the species can be recognized visually, mostly through the leaf,
stem and bark pattern, and reproductive morphologies. According to Laksmi and
Gopakumar (2009), morphological keys for A. mangium include rhytidome bark
rectangular in shape; length of internode, short (~1.5 cm; leaves most spaciously
arranged); phyllode, broad (~5.4 cm) and looking more like an entire leaf, rough
textured due to the presence of four nerves on the dorsal side of the phyllode, which
can be prominently perceived by touch; flower, creamy- white color; pod, highly
coiled; funicle, orange color. In A. auriculiformis the rhytidome bark is rectangular
in shape; phyllode, short (~12.6 cm), sickle-shaped; pod, highly coiled; funicle,
curved and enclosing more than half of the seed, reddish-orange color. For A. crassi-
carpa: the rhytidome bark is square in shape; length of internode long (~3.3 cm)
which means that leaves are arranged more compactly on the branchlets; phyllode,
thick with a grayish tint and sickle-shaped; flower, greenish-yellow color; pod, flat
and uncoiled; prominent protrusion in the middle of the seed; funicle, creamy-white
color. However, dissimilar site and climate may show slight differences in morphol-
ogy for the three species, especially for the bark color and rythidome shapes. Some
morphology keys of the four tropical acacias grown in Central Java, Indonesia are
presented in Figs. 1.1 and 1.2. Seed morphology of the three tropical acacias is
presented in Fig. 1.3.
Rather than in morphological and biochemical markers, the revolution in bio-
technology is in the molecular basis for DNA sequencing, which has become an
important tool for accelerating an accurate understanding of phylogenetic and
genetic diversity in a species. Random amplified polymorphic DNA (RAPD),
amplified fragment length polymorphism AFLP and restriction fragment length
polymorphism (RFLP) are among the most common technologies employed in phy-
logenetic studies. In crops and horticultural species, phylogenetic studies using
molecular markers have become a common tool and have resulted in the completion
of several thousand phylogenetic analyses.
Fig. 1.1 Bark color and rhytidome shape from mature trees grown in Central Java, Indonesia for
(a) Acacia mangium, (b) A. auriculiformis (c) Acacia hybrid A. mangium × A. auriculiformis, (d)
A. crassicarpa, (e) the phyllode shapes from the respective species: (1) A. crassicarpa, (2) A. man-
gium, (3) Acacia hybrid, (4) A. auriculiformis. (Photographs by A. Nirsatmanto)
Fig. 1.2 Flower morphology of (a) Acacia mangium, (b) A. auriculiformis, (c) Acacia hybrid A.
mangium × A. auriculiformis, (d) A. crassicarpa. (Photographs by A. Nirsatmanto)
Fig. 1.3 Seed and funicle morphology of Acacia mangium, A. auriculiformis and A. crassicarpa
compared with other Acacia species. (Photograph by A. Nirsatmanto)
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naimisissa vai ei… Ja minkä minä nyt sille voin, että rakastan sinua
— etten saa sinua mielestäni? Eikä se sitäpaitsi niin kauhea asia ole,
kaikilla paremmilla ihmisillä on joku toinen, jota he rakastavat —
vaikka ovatkin naimisissa. Älä siis…
(Lähenee Annia.)
ANNI katkeran ivallisesti, siirtyen poispäin.
Mestarillahan on toisia, montakin — jos se kerran on välttämätöntä

paremmille ihmisille! Miksi te siis minua kiusaatte?
MESTARI teennäisen moittivasti.
Voi Anni! Sinäkö syytät minua kiusaamisesta, vaikken ole tahtonut

sinulle muuta kuin hyvää? Sinä et välitä vähääkään minun
tunteistani, lähetit lahjanikin sillälailla takaisin. — (Ottaa taskustaan
kellon ja lähenee taas Annia.) — Ota tämä edes… otathan Anni…
ANNI tuskastuneena.
Välitättekö te minun tunteistani, kun yhtämittaa puhutte sitä

samaa, vaikka monasti olen sanonut etten voi teistä pitää? Miksi te
nytkin tyrkytätte väkisin lahjaanne kun minä en sitä tahdo?
MESTARI
Enhän voinut aavistaa, että sinä tulet siitä entistä katkerammaksi.
Odotin minä edes ystävällistä sanaa, katsetta — ajattelin, että sinä
sentään lopulta alat ymmärtää, tulet minun ystäväkseni… Minä
järjestäisin kaikki niin hyvin, saisit kaksinkertaisen palkan, ja…
ANNI punastuen harmista ja häpeästä.
Te tahtoisitte tuollaisella kellolla ostaa minut, mutta se ei tapahdu!

Minä en tee koskaan sellaista rikosta! Olenhan jo sanonut, että pidän
toisesta — että minulla on sulhanen, Kaarlo…
MESTARI silmät välähtäen; hillitsee itsensä tekeytyen

hämmästyneeksi.
Kas, kun en lainkaan muistanut! Minähän kuulin jo tuon asian! Ja

kuulin minä sen yhteydessä muutakin… — (Vaikenee, katsoen
Annia, ja kun tämä ei puhu mitään, jatkaa ikäänkuin säälien.) — Niin,
olen vaan kuullut sellaista, että sinun tuota… täytyisi mennä… —
(Hän venyttää tahallaan sanojaan, tuijottaen kaiken aikaa
läpitunkevasti Annia, ja lopulla vaikenee.)
ANNI tuskastuen.
Mihinkä niin?
MESTARI tuijottaa yhä Annia, lausuen painokkaasti.
Niin — että sintin täytyisi mennä hänen kanssaan naimisiin —

mahdollisimman pian… Onko se totta?
ANNI punastuu ja huudahtaa silmät säkenöiden.

Kuinka mestari voi luulla minusta sellaista! Luuletteko minut
sellaiseksi?
(Kääntyy poispäin.)
MESTARI myhähtää ensin tyytyväisenä, sitten lausuu ankarasti.
Mitä sinä teeskentelet? Kyllähän minä tiedän, että nuoret

tehtaalaiset keskenään ihan tuota… ihan…
ANNI keskeyttää tulisesti.
Mutta minä en!
MESTARI jatkaa aivankuin samaa lausetta ja hänen pienissä

silmissään välkähtää kavala, ivallinen ilme.
Ihan suutelevat… No, etkö sinä siis koskaan ole sitä tehnyt?
ANNI kääntyy hämmentyneenä toisaalle, eikä vastaa mitään.
MESTARI
Etkö sinä voi vastata? Vai myönnätkö sen vaitiolollasi?
ANNI sammaltaa suuttuneena, häveten.
Miten mestari kysyy sellaista..? Eihän se kuulu… tuota…
MESTARI keskeyttää ilkeästi.
Minulleko? Vai ei kuulu minulle? Hyvä! Minä taas luulen, että se

kuuluu hiukan minullekin. — Minä huomaan, että sinä olet juuri
hänen tähtensä kohdellut minua kuin kerjäläistä! Kiusannut minua
kylmyydelläsi ja samaan aikaan suudellut ja rakastellut häntä. Eikö
niin? Vai kiellätkö, ettet rakasta tuota nulikkaa?
ANNI seisoo häneen selin, äänettömänä ja hämmentyneenä.
MESTARI yhä kiihtyen.
Jaha! Sinä myönnät sen! Vai niin. Tuota lurjusta minä siis saan
kiittää kaikesta. Tuollaisen tähden olet sinä siis pitänyt pilkkanasi
minua, naureskellut sen kanssa takanapäin minua — minua! —
(Uhkaavasti.) — Mutta sen täytyy nyt loppua.
ANNI naivisti, pelokkaasti.
Ei herra mestari… en minä ole koskaan nauranut teitä. Eikä

Kaarlo ole lurjus… hän on niin hyvä, siivo ja rehellinen… Kaikki
pitävät…
MESTARI kiukkuisesti.
Hän on lurjus — kirottu lurjus! Ja minä en enää kärsi sitä peliä,

jota te yhdessä harjoitatte! Huomispäivänä minä ajan hänet pois
tehtaasta, koko läänistä! Se on nyt sanalla sanottu.
ANNI änkyttää tolkuttomasti.
Oi herra mestari, älkää ajako häntä pois työstä — — hän ei ole

mitään pahaa tehnyt… Aina tehnyt työnsä hyvin, ollut säännöllinen ja
altis ja…
MESTARI keskeyttää ärjäisten.
Vaiti, sanon minä! Minä en viitsi kuunnella sinun lörpötyksiäsi.

Huomenna hän lähtee — ja' samalla sinä, isäsi, veljesi, kaikki —
koko roikka!
ANNI vaikenee ikäänkuin jähmettyneenä; sitten sammaltaa

tuskallisesti.
Isä… isäkin…? Ei, ei… Mihinkä isä ja äiti sitten joutuvat tällaisena
työttömänä aikana — ja niin monta lasta… Älkää sentään eroittako
isää työstä… Hyvä mestari…
MESTARI keskeyttää järkähtämättömästi.
Se riippuu nyt kokonaan sinusta. Jos olet järkevä, niin ehkä annan
heidän jäädä. — (Pahanenteisesti.) — Mutta ellet… ellet sinä…
ANNI keskeyttää tuskallisesti.
Älkää, herra mestari! Isä on ollut niin kauan täällä — koko ikänsä
ja, ja… — (Miltei vaikeroiden.) — Niin, ettekö te sitten ymmärrä, että
minä en voi, minä en voi…
MESTARI
No, sitten ei siitä enää kannata puhua. — (Ankarasti.) — Minä en

anna itseäni pilkata!
ANNI
Voi hyvä mestari, mitä te aiotte… Armahtakaa nyt tämä kerta,

älkää…
(Purskahtaa itkuun.)
MESTARI antaa Annin itkeä hetkisen, tuijottaa vain kummallisen
tutkivasti. Sitten hänen katseensa muuttuu neuvottomaksi, hän
lähenee, taputtaa hyväillen Annin olkapäätä ja hänen äänessään
värähtää levottomuus ja hellyys.
Auni älä itke, en voi sitä kestää… Anni ota edes tämä ja ole hiukan
ystävällinen minulle. Ota nyt Anni, äläkä itke Anni. Katsos miten
kaunis se on — perätkin ovat kullasta… Se maksaa kolmattasataa
markkaa…
ANNI nyyhkyttäen.
Miten minä voisin sen ottaa, sanoa että olen saanut sen teiltä?
Jokainen ajattelisi heti, että…
MESTARI kiihkeästi kuiskien.
Eihän sinun tarvitse sitä heti näyttää ja sitten jälkeenpäin voit

keksiä jotain, sanoa että olet kauan säästänyt sitä varten. Ota Anni
— minä pyydän. — (Työntää kellon Annia taskuun, sanoo
mahdollisimman lempeästi.)
Kas niin, Anni. — Älä nyt itke. Minä en voi katsella sinua itkevänä
— se koskee niin sydämeeni. Jospa vain tietäisit, miten silloin kärsin
— ja aina… Ja sinä voisit lopettaa kärsimyksen! kun vaan tahtoisit —
voisit muuttaa kaikki hyväksi… meille molemmille…
ANNI yhä itkien.
Mutta sehän olisi valhe… minä tuntisin aina, että olen tehnyt
rikoksen. Ei, minä en voi ottaa sitä… Ottakaa se takaisin…
MESTARI tarttuu hänen käsiinsä puhuen yhä kiihkeämmin.

Rakas Anni! Onko sekin mielestäsi rikos? Jospa vain tietäisit,
miten syvästi sinua rakastan — niin et puhuisi noin… Minähän
tahdon tehdä kaikki, mitä tahansa sinun hyväksesi, kun vain olisit
toisenlainen… Anni, sinä saisit suuremman onnen — — —. sinun ei
enää koskaan työssä tarvitsisi työssä käydä… minä korottaisin sinut
konttoristiksi. Opettaisin sinulle kirjanpidon, veljesi pääsisi
monttöriksi ja isäsi alityönjohtajaksi eikä huominen palkanalennus
lainkaan koskisi heitä. Niin, sen järjestäminen on minulle helppoa —
se olen minä, joka tässä tehtaassa määrään! Ajattele siis mikä valta
minulla on! Etkö edes omaistesi tähden koeta olla järkevä — ajatella
minunkin mieltäni…?
ANNI on monta kertaa koettanut keskeyttää mestaria; huudahtaa

epätoivoisesti.
Herra mestari… Älkää puhuko enää… minä en voi! Kuinka minä

sitten enää voisin olla täällä, katsoa ketään silmiin?
MESTARI huomaa Annissa horjuvaisuutta ja jatkaa yhä

kiihkeämmin.
Oi Anni, minkätähden sinä kiusaat minua? Eikö sinulla ole

lainkaan sydäntä? Sinä et yhtään ajattele, miten paljon olen kärsinyt
sinun tähtesi… Älä nyt enää ole tuollainen! Tule nyt edes katsomaan
minun huonettani. Ja luota minuun…
(Koettaa vetää Annia huoneeseensa.)
ANNI ponnistelee vastaan, rukoillen hätäisesti ja tuskallisesti.
Oi mestari hyvä, älkää nyt… Antakaa minun olla… Lakatkaa…

Minä koetan, lupaan ajatella… Älkää…
MESTARI kähisevin äänin.
Anni, minä pyydän, rukoilen sinua! Ajattelu, mikä onni sinulle tulisi!
Sinä et tiedä, miten hyvin kävisi omaisillesi — etkö sinä tahdo edes
heidän parastaan…?
ANNI näyttää menehtyneeltä ja katsahtaa hätääntyneenä ovelle.
Kyllä, kyllä tahtoisin, mutta en voi, en voi… Päästäkää, minä

lupaan ajatella — minä tulen huomennakin, kun vain päästätte!
Päästäkää! — (Kun mestari ei laske, kirkasee hän äkkiä.) — Apua,
Kaarlo!… Kaarlo!
MESTARIN sieramet laajenevat, silmät pienenevät

naskalinteräviksi ja hänen äänessään kuvastuu sekaisin intohimo,
pelko, raivo ja voitonriemu — se muistuttaa miltei käärmeen
sähinääa.
Vaiti! Jos vielä kerran huudat — niin se lurjus viedään täältä

ijäksi…
ANNI änkyttää kauhusta vavisten.
Mitä, mitä…? Se ei ole mahdollista…
MESTARI
Se on mahdollista! Hän on täällä yhdistyksessä puhunut

sillätavoin, että jos vihjaisen kaupungin santarmeille, niin hänet
viedään tavallista kauemmaksi — josta hän ei palaja koskaan.
Kuinka sinä uskalsit huutaa häntä — pilkata minua? Sillä hän on
minun kädessäni!
ANNI rukoillen.
Mestari… antakaa anteeksi jumalan tähden… En tahtonut

huutaa… en itsekään ymmärrä… Älkää antako ilmi häntä! Älkää,
hyvä mestari, älkää! Minä kuolisin… Minä teen mitä tahansa, kun
vaan…
MESTARI pidätetyllä voiton riemulla.
En minä odottanut noin suurta loukkausta sinulta. Mutta minä

annan vielä senkin anteeksi, jos näytät, ettet tahtonut minua pilkata.
Jos huomenna työajan jälkeen jäät tänne, niin minä lupaan, että
omaisesi saavat jäädä työhön entisellä palkalla, ja kaikki tulee
niinkuin olen puhunut.
ANNI hätäisesti sammaltaen.
Kyllä minä sitten tulen huomenna… huomenna… — (Kuin

havahtuen.) —
Ei, ei — en minä sentään voi — se olisi liian kamalaa.
MESTARI uhkaavasti.
Mitä, pilkkaatko sinä vielä silläkin tavalla? Ensin lupaat ja sitten

kiellät — aivan kuin ärsytät! Ei — se on liikaa! Minä lähetän hänestä
kirjelmän vielä tänään!
ANNI parahtaa epätoivoisesti.
Oi älkää, jumalan tähden! Minä tulen… lupaan tulla…
MESTARI raskaasti.
Tuletko siis varmasti! Jos sinä vielä tässä pettäisit minut, niin
silloin en enää armahda ketään! Tuletko?
ANNI
Minä tulen…
(Vaipuu kalpeana polvilleen.)
MESTARI nostaa Annin ylös alkaen taluttaa huoneeseensa,

toistellen
Anni, Anni, sinä voit pahoin… Tule ottamaan vettä… tule minun
huoneeseeni.
ANNI epätoivoisesti.
Oi älkää mestari… jumalan tähden en voi tulla, nyt… Päästäkää!
MESTARI käheästi, uhaten.
Älä vain huuda! Muista mitä siitä seuraa…
ANNI kuin sekopäisenä.
Hu, en minä huuda… kun ette vain… mestari hyvä — minä lupaan
tulla huomenna varmasti. — (Mestari vie hänet puoliväkisin sisään
sulkien oven.)
Oven takaa kuuluu Annin pelokas nyyhkytys ja rukoileva

kuiskaava ääni, vuoroin taas mestarin matala, kiihkosta käheä
kuiske katkonaisena, epäselvänä.
— Älä pelkää — istu — voit pahoin…

— Ei — hyvä mestari — menee ohi…
— Maista nyt — virkistää…
— En voi — viiniä — päästäkää…
— Älä nyt — ymmärrä — omaisesi…
Sen jälkeen ei enää eroita sanoja; ne hukkuvat kaukaiseen

kohinaan joka vähitellen lähenee; sen synnyttävät satojen askelten
jyminä ja ihmisjoukon sekava puheensorina, josta ei vielä sanoja
eroita. Se lähenee, kiihtyy ja lopulta pysähtyy ulkopuolelle ja sen
seasta kuuluu kiihtyneitä
HUUTOJA
— Ne aikovat tietysti tappaa meidät nälkään!
— Tietysti! Eihän sillä palkalla elä…
— Mutta se on pirullista, ettei sillä heti kuolekaan!
— Älä ole siitä huolissasi — kyllä tämän alennuksen jälkeen kuolet

heti!
— Se on selvä se!
— Kun oikein nälkäpiiskuri suunnittelee, niin kai se tepsii!
— Oikein! Tietysti se mestarin työtä on!
— Tietysti! Sehän syksylläkin oli ehdottanut!
— Menkää sisään! Ei se tästä parane!

— Menköön Korpi ja Suonpää sisään! Kyllä me odotamme täällä!
— Halonen myöskin!
Kuuluu Halosen vastustava ääni.
— Mene nyt vain! Ei se sinua syö!
— Ei se ainakaan purematta nielase!
— Ei kukaan paljaista luista huoli!
— Mene sinäkin Emil — silloin se ei kuitenkaan kaikkia syö!
— Mene vaan! Olet vähän lihavampikin!
Joitakin katkeria naurahduksia.
Melu hiljenee ja sitten aukee ovi; Korpi, hänen poikansa Emil,

Halonen ja Kaarlo Suonpää astuvat sisään. Edellinen on
suurikokoinen, vielä voimakas mies, vaikka ankara työ ja huolet
ovatkin uurtaneet syviä vakoja hänen kasvoihinsa; Halonen on kovin
laiha, kalpeilla kasvoillaan masennus ja arkuus; Suonpää on
solakka, jäntevävartaloinen nuorukainen, jolla on älykkäät ja
voimakkaat kasvot.
Ulkoa kuuluu edelleen joukon hillittyä sorinaa.
KORPI kuin pettyneenä.
Täällähän ei ole ketään…
EMIL pilkallisesti hymähtäen.

Onkohan kettu paennut pesästään!
KAARLO
No, odotetaan täällä vaikka huomisaamuun asti! Sitä asiaa ei

jätetä.
KORPI hiljaa, lujasti.
Sitä ei voi jättää. Jos nyt vielä alennettaisiin palkat kolmannella

osalla, silloin tulee monelle suurperheiselle suorastaan
nälkäkuolema.
HALONEN
Niin se on, ei sillä saa…
(Hän vaikenee, sillä mestarin huoneen ovi aukenee ja kynnykselle

ilmestyy Anni, hätääntyneenä, vapisevana, vaatteet ja tukka hieman
epäjärjestyksessä. Kaikki aivankuin jähmettyvät hämmästyksestä.
Äänettömyys.)
KORPI hiljaa, käheästi.
Sinä täällä… Mitä sinä teit siellä?
ANNI nyyhkyttäen.
Minä olin… olin tomuja pyyhkimässä…
KORPI ärjäisten.
Valehtelet! Sano heti, mitä sinulla on täällä tekemistä!

ANNI kuin menehtyneenä.
Mestari… tuota noin, käski… — (Huomaa Kaarlon, pelästyy yhä

enemmän ja sopertaa vavisten.) — Kaarlo… täällä…?
KAARLO on seissyt kalpeana ja liikkumattomana — niinkuin raivo

ja hämmästys olisi lamauttanut hänet.
Niin, minä olen täällä… Mutta mitä sinä teet täällä? Mitä sinä olet
tehnyt…?
ANNI tolkuttomasti.
Jumala auttakoon! En minä mitään…
KAARLO raivoisasti.
Mitä sinä olet tehnyt? Vastaa minulle. Oletko sinä, sinä…? Onko,
onko se totta?
MESTARI tulee huoneestaan ja karjaisee röyhkeästi.
Mitä tämä merkitsee? Mitä te täällä rähisette? Ulos jokainen!
KAARLO
Konna!… Mitä te olette tehnyt hänelle?
(Aikoo hyökätä mestarin kimppuun, mutta Korpi saa kiinni hänen

olkapäästään.)
MESTARI lähenee pari askella ja ähkyy pelästyksestä ja raivosta

vavisten.
Vai niin… vai niin! Vai sinä hyökkäät minun kimppuuni! No, no…
Te näitte sen, te voitte todistaa sen! — (Kaikille.) — Ja ettekö
kuulleet, mitä sanoin? Ulos!
KAARLO hiljaa, raskaasti.
Te ette pääse tästä asiasta niin helpolla kuin luulette.
KORPI Kaarlolle hiljaa.
Tule järkiisi. Ole hiljaa! — (Mestarille.) — Emme me sillä tavalla

ulos lähde. Olemme tulleet puhumaan tärkeistä asioista.
MESTARI
Mistä asioista? — (Kuuntelee jonkun sekunnin ja huomaa joukon

hillityn sorinan ulkopuolella.) — Mitä asioita teillä on? Keitä, mitä
rähinää tuolla on?
KORPI
Siellä on koko tehtaan työväki ja me olemme tulleet kaikkien

puolesta ilmoittamaan, ettemme voi hyväksyä sitä palkanalennusta.
Me emme tule sentään toimeen muutamalla pennillä. Me vaadimme,
että palkat pysytetään ennallaan.
HALONEN avuttomasti.
Minullakin on kahdeksan lasta…
MESTARI koettaa tekeytyä ivalliseksi.

Vai on Halonenkin mukana tuollaisessa! — (Kääntyen Korpeen.)
— Ja oikein vaaditte! Vai niin. Mutta palkkojen määrääminen on
herra patruunan asia, eikä hän kuuntele teidän vaatimuksianne!
HALONEN mutisee mestarin puheen aikana pelästyneenä.
Kun on lapsia, tuota… niin ei silloin…
KORPI
Jonkun täytyy meitä kuunnella. Muuten emme voi elättää

lapsiamme. Ja jollei muu auta, niin meidän täytyy lopettaa työt…
MESTARI pahaenteisen hitaasti, pilkallisesti hymyillen.
Ahaa! Sinä kiihotat siis julkisesti lakkoon. Hyvä. Sinä siis lopetat
tehtaan työt… Mutta minä luulen, että töiden lopettaminen on
myöskin herra patruunan asia! Ilmoittakaa siis hänelle itselleen
uhkauksenne — jos hän ottaa teidät vastaan!
KORPI
Kuulkaa herra mestari. Kun te olette itse tämän alennuksen

ehdottanut patruunalle, niin teidän velvollisuutenne on myös ilmoittaa
hänelle sen seurauksista. Enkä minä ole ketään kehoittanut lakkoon
tai muuhun. Mutta pitääkö meidän siis jättää kaikki sikseen ja tulla
kiltisti työhön jollette suvaitse edes ilmoittaa patruunalle… Mitä te
arvelette, toverit?
KAIKKI
— Silloin teemme lakon — ja heti aamulla!

— Niin teemme! Emme voi muuta.
— Meidän täytyy — jollette muuten ilmoita.
MESTARI pöyhkeästi.
Vai lakon te nyt teette? Mutta vakavasti puhuen ei se kaipaa

mitään ilmoittamista. Herra patruunan järkähtämätön päätös on, että
palkat alennetaan niinkuin julistuksessa on säädetty — eikä se tule
muuttumaan teidän rähinänne vuoksi.
ANNI peittää käsillään kasvonsa, purskahtaen itkuun.
Voi tätä elämää…
KORPI
Mutta me emme enää aio vapaaehtoisesti nälkään kuolla! —

(Katsahtaen itkevään Anniin ja jatkaa yhä kasvavalla katkeruudella
ja tuskalla.) — Ja sinä, sinä… mitä sinä olet tehnyt tyttärelleni?
Vieläkö sinä hänetkin tahdot uhriksesi! Eivätkö entiset riitä…? Ei siitä
vielä ole viikkoakaan kun viimeinen haudattiin — siellä se makasi
meidän aitassa jäätyneenä…
MESTARI on muuttunut kamalan näköiseksi kuullessaan nuo

murhaavat syytökset; mutta äkkiä hän keskeyttää syyttäjänsä,
karjaisten raivoisasti.
Mitä? Mitä? Mitä tämä oikein on? — Mitä sinä tarkoitat? Te olette
minun konttorissani! Ulos! Ulos! Ulos!
EMIL hurjasti.
Vieläkö sinä karjut! Ellet nyt ole hiljemmin, niin — niin piakkoin
tulee toiset odottamattomat hautajaiset…
KAARLO hiljaa, hammasta purren.
Tietäkää nyt, että teidän täytyy lopettaa mustat tekonne!

Ymmärrättekö te?
KORPI hilliten itsensä.
Älkää viitsikö sanoa enää mitään hänelle — puheet eivät tässä

auta!
Eikä väkivaltaisuudet! Niistä olisi vain heille hyötyä. — (Lujasti.) —
Loppu tästä on kyllä tehtävä — mutta toisella tavalla!
(Menevät ulos, paitsi Kaarlo, ja ovi jää raolleen. Joukon sorina

lakkaa, syntyy miltei kuolemanhiljaisuus. Sitten kuuluu Korven
mielenliikutuksesta väräjävä ääni.)
KORPI
Työtoverit! Mestari ei lupaa edes ilmoittaa patruunalle

vaatimustamme; ivailee vain ja sanoo olevansa varma siitä, että se
hylätään. Mitä on meidän tehtävä? Onko meidän tyyninä kärsittävä,
että palkat vieläkin alennetaan? — että meidän lapsemme joutuvat
kitumaan nälässä? Mitä te ajattelette, toverit?
KIIHKEITÄ HUUTOJA
— Ei — me emme anna enää alentaa palkkoja!
— Syksyllähän niitä juuri alennettiin…

— Me olisimme raukkoja, jos vieläkin…
— Ei enää…
— Lakko on meidän tehtävä!
— Ja huomispäivänä!
— Lakkoon, lakkoon!
— Ja nyt vaaditaan mestari pois!
— Oikein! Mestari pois!
KORPI
Se on todella kamalaa, että hän edelleenkin saa rauhassa jatkaa

vanhaa peliään. Vuosikausia on hän kiusannut, hätyytellyt ja
raiskannut tyttäriämme, vaimojamme. Heidän turvallisuutensa ja
kunniansa ei enää merkitse mitään. Hän on saanut aikaan niin paljo
onnettomuutta, että sen pitäisi jo riittää! — (Epäröiden.) — Minä
pahoin pelkään, että hän vielä tänään on tehnyt konnantyön, joka…
Hänet keskeyttävät kiihkeät, uhkaavat
HUUDOT
— Mitä hän taas on tehnyt?
— Pois mestari tehtaalta!
— Piiskuri pois!
— Pois se sika!

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PDF Advances in Plant Breeding Strategies Industrial and Food Crops Volume 6 Jameel M Al Khayri Ebook Full Chapter

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Advances in Plant Breeding Strategies

Industrial and Food Crops Volume 6

Advances in Plant Breeding Strategies Nut and Beverage

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 978-3-030-23264-1 ISBN 978-3-030-23265-8 (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 Industrial Crops

9 Flax (Linum usitatissimum L.) Genomics and Breeding���������������������� 277

Part II Food Crops

About the Editors

Prof. Jameel M. Al-Khayri is a Professor of Plant

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

Prof. Shri Mohan Jain is a Consultant and Plant

Prof. Dennis V. Johnson is a Consultant and Former

Hussein Abdel-Haleem USDA-ARS, ALARC, Maricopa, AZ, USA

Allah Bakhsh Department of Agricultural Genetic Engineering, Faculty of

Mohamed M. A. Khairi Faculty of Agriculture, Dongola University, Dongola,

Deb Prasad Ray Indian Council of Agricultural Research-National Institute of

Arif Nirsatmanto and Sri Sunarti

Keywords Conventional breeding · Tree improvement · Molecular breeding ·

A. Nirsatmanto (*) · S. Sunarti

© Springer Nature Switzerland AG 2019 3

1.2 Silviculture and Conventional Breeding

1.2.1 Current Cultivation Practices

1.2.2 Current Problems and Challenges

Under current silvicultural practices, the productivity of acacia plantations has

1.2.3 Improvement Strategy

Some improvement strategies are underway to overcome current problems of acacia

1.2.4 Traditional Breeding Methodologies and Limitations

Breeding strategies using a recurrent selection system is regularly practiced in aca-

1.2.5 Role of Biotechnology

In tree breeding programs, biotechnology advances include new methods in vegeta-

1.3 Germplasm Biodiversity and Conservation

1.3.1 Germplasm Diversity

The natural distribution of the three acacias is in northern Australia, southeastern

The natural distribution of auriculiformis is geographically wider than mangium.

1.3.2 Species Characterization and Phylogeny

MESTARI niinkuin ei kuulisikaan.

Miten olet jaksanut viime päivinä? Oliko sinulla hauska joulu?

Ja lahjoja tuli kai oikein paljon?

Kuinka mestari niin luulee? En minä saanut ainoatakaan lahjaa.

Todellakin! Ja minä luulin niitä tulleen oikein summassa kun ei

Ymmärtäähän mestari, ettei minun sovi ottaa sellaisia lahjoja…

MESTARI kuin ihmetellen.

Mikä siinä olisi sopimatonta? Jos tahdon antaa lahjan ystävälleni

ANNI keskeyttää, harmista punastuen.

ANNI katkeran ivallisesti, siirtyen poispäin.

Mestarillahan on toisia, montakin — jos se kerran on välttämätöntä

MESTARI teennäisen moittivasti.

Voi Anni! Sinäkö syytät minua kiusaamisesta, vaikken ole tahtonut

Välitättekö te minun tunteistani, kun yhtämittaa puhutte sitä

ANNI punastuen harmista ja häpeästä.

Te tahtoisitte tuollaisella kellolla ostaa minut, mutta se ei tapahdu!

MESTARI silmät välähtäen; hillitsee itsensä tekeytyen

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

Part I Industrial Crops

9 Flax (Linum usitatissimum L.) Genomics and Breeding�� 277

Part II Food Crops

1.2 Silviculture and Conventional Breeding

1.2.1 Current Cultivation Practices

1.2.2 Current Problems and Challenges

1.2.3 Improvement Strategy

1.2.4 Traditional Breeding Methodologies and Limitations

1.2.5 Role of Biotechnology

1.3 Germplasm Biodiversity and Conservation

1.3.1 Germplasm Diversity

1.3.2 Species Characterization and Phylogeny