sunflowers 2014
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BOTANICAL RESEARCH AND PRACTICES
SUNFLOWERS
GROWTH AND DEVELOPMENT,
ENVIRONMENTAL INFLUENCES
AND PESTS/DISEASES
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BOTANICAL RESEARCH AND PRACTICES
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BOTANICAL RESEARCH AND PRACTICES
SUNFLOWERS
GROWTH AND DEVELOPMENT,
ENVIRONMENTAL INFLUENCES
AND PESTS/DISEASES
JUAN IGNACIO ARRIBAS
EDITOR
New York
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Copyright 2014 by Nova Science Publishers, Inc.
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Library of Congress Cataloging-in-Publication Data
Sunflowers : growth and development, environmental influences and pests/diseases / editor: Juan
Ignacio Arribas (Electrical Engineering Department, Univ. Valladolid, Spain).
pages cm
Includes index.
1. Sunflowers. I. Arribas, Juan Ignacio.
QK495.C74S87 2014
583'.99--dc23
2014003599
Published by Nova Science Publishers, Inc. New York
ISBN: (eBook)
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CONTENTS
Preface vii
Chapter 1 An Introduction to the Sunflower Crop 1 Fabin Fernndez-Luqueo, Fernando Lpez-Valdez,
Mariana Miranda-Armbula, Minerva Rosas-Morales, Nicolaza Pariona and Roberto Espinoza-Zapata
Chapter 2 Floral Biology of Sunflowers: A Histological
and Physiological Analysis 19 Basudha Sharma, Rashmi Shakya and Satish C. Bhatla
Chapter 3 Development of Female Reproductive Structures and Apomixis
in Sunflowers 43 Olga N. Voronova
Chapter 4 Genetics and Genomics Applied to Sunflower Breeding 61 Carla Filippi, Jeremas Zubrzycki, Vernica La,
Ruth A. Heinz, Norma B. Paniego and H. Esteban Hopp
Chapter 5 Sunflower Genetic Resources: Interspecific Hybridization
and Cytogenetics in Prebreeding 95 Jovanka Atlagi and Sreten Terzi
Chapter 6 Functional Genomics and Transgenesis Applied to
Sunflower Breeding 131 Sebastian Moschen, Laura M. Radonic, Guillermo F. Ehrenbolger, Paula Fernndez, Vernica La, Norma B. Paniego, Marisa Lpez Bilbao,
Ruth A. Heinz and H. Esteban Hopp
Chapter 7 Disease Management in Sunflowers 165 Regina M. V. B. C. Leite
Chapter 8 Recent Advances for Developing Resistance against
Plasmopara halstedii in Sunflowers 187 Osman Radwan
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Contents vi
Chapter 9 Effects of Crop Management on the Incidence and Severity
of Fungal Diseases in Sunflowers 201 P. Debaeke, E. Mestries, M. Desanlis and C. Seassau
Chapter 10 Insect Pests of Sunflowers in Africa 227 Hannalene du Plessis
Chapter 11 Soil Amendments and Their Effects on Sunflower Growth 239 Fernando Lpez-Valdez, Fabin Fernndez-Luqueo, Perla Xchitl Hernndez-Rodrguez, Minerva Rosas-Morales
and Silvia Luna-Surez
Chapter 12 Nutrition and Fertilization of Sunflowers in Brazilian Cerrado 257 C. de Castro, F. A. Oliveira, A. Oliveira Junior
and N. P. Ramos
Chapter 13 Environmental Issues in the Sunflower Crop of Midwestern Brazil:
Diversification and Complementarities in the Biodiesel Chain 281 N. P. Ramos, A. M. M. Pires, C. C. A. Buschinelli,
H. B. Vieira, C. de Castro and G. S. Rodrigues
Chapter 14 Micro and Macro-Morphological Variation of Cosmos bipinnatus
and Cosmos bipinnatus var. Albiflorus in Sympatric Zones in
Central Mexico 297 M. Paniagua-Ibaez, A. Zepeda-Rodrguez, P. Mussali-Galante,
R. Ramrez-Rodrguez and E. Tovar-Snchez
Index 309
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To Juan Ignacio, Jr., Elena, Jr. and Elena
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PREFACE
We are all well aware that the importance of the sunflower (Helianthus Annuus) as a crop
has increased significantly in recent years, not only in the food industry but also as a natural
energy resource in oil production. I am, thus, very pleased to be able to present this
comprehensive monograph on a wide range of important issues regarding sunflowers, with an
emphasis on environmental influences, pests and diseases in order to maximise production
whilst minimising costs.
Contributors where selected based on their proven experience in the field of sunflowers.
Contributors submitted an extended abstract that was assessed for relevance. They were then
invited to contribute draft chapters. Each chapter underwent a stringent and thorough peer
review process by other experts in the field, with final approval by the editor who, thus, was
able to balance the topics from all contributors.
The book contains important original results. Each chapter deals with a different topic,
and draws, where appropriate, from studies and results previously published by the authors.
Authors were encouraged to complement their writing with original and high quality graphs,
charts, tables, figures, pictures and photographs.
Its my honour and pleasure to acknowledge the rigorous work carried out by all authors
in this book, and at the same time I am very grateful to them for trusting me in leading this
project in the role of the editor of their work. My thanks also go to the anonymous reviewers
who contributed their time so generously to this book, and without whom it would not exist.
I am also very grateful to Nova Science Publishers for inviting me to lead this book, and
thank them for the help and coverage provided during the whole time that this project lasted.
I really do hope that you find this book of interest and wish you enjoy its reading as much
as I have done through the whole editing process and as much I am sure all authors have done
while writing it.
The book is structured as follows: Chapter 1 introduces sunflowers. Chapters 2 and 3
detail the biology of a sunflower. Chapters 4, 5 and 6 explore the important topics of
sunflower production: genetics, genomics and the breeding of sunflowers. Chapters 7, 8, 9
and 10 address with other important aspects of sunflower production, pests and diseases.
Chapters 11 and 12 deal with sunflower nutrition and growth. Finally, Chapter 13 presents
environmental sunflower crop issues and Chapter 14 studies sunflower morphological
Cosmos variations.
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Juan Ignacio Arribas x
In Chapter 1, entitled An Introduction to the Sunflower Crop, Dr. Luqueno and colleagues
from the Natural Resources and Energy Group, Cinvestav-Saltillo, Mexico, present an
enjoyable historical perspective on sunflowers. Sunflower (Helianthus annuus L.) belongs to
the family Asteraceae. The sunflower plant originated in eastern North America. It is thought
to have been domesticated around 3000 B.C. by Native Americans. In the late 1800s the
sunflower was introduced in the Russian Federation where it became a food crop and Russian
farmers made significant improvements in the way that the sunflower was cultivated. Since
3000 B.C. a wide range of uses of sunflower have been reported throughout the world.
Sunflower is well known by its phytoremediation potential and by its seed oil content.
Because the sunflower has several potential markets, it is a good choice for growers on both
small and large scales. However, it has to be remembered that scientific, technical or
agricultural projects linked with sunflower have to include side effects elsewhere in order to
shape a sustainable future.
In Chapter 2, entitled Floral Biology of Sunflower - A Histological and Physiological
Analysis, Dr. Bhatla and colleagues from the Laboratory of Plant Physiology and
Biochemistry, Department of Botany, University of Delhi, India, introduce a meticulous
approach to the development of sunflower inflorescence as considered under three phases
listed next: inflorescence initiation, floret development and anther formation. Anthesis of disc
florets is a phytochrome-mediated response and is also modulated by phytohormones, such as
auxins and gibberellic acid. Dr. Bhatla and colleagues focus on the role of various
biomolecules, like glycoproteins, calcium, nitric oxide, reactive oxygen species, and
associated scavenging enzymes in relation to stigma maturation. Specific expression of
lignoceric acid (24:0) in the pollen coat and localization of lipase in pollen and stigma are
likely to have possible roles during pollen-stigma interaction. The phenomenon of self-
incompatibility and pseudo self-compatibility in sunflower has been discussed. The initial
processes accompanying pollen-stigma interaction and their regulation, especially the
adhesion of pollen on the stigma surface, hydration, formation of an "attachment foot" and
pollen tube germination in sunflower with respect to self-and cross-pollinated situations, has
also been dealt with in detail.
In Chapter 3, entitled Development of Female Reproductive Structures and Apomixis in
Sunflowers, Dr. Voronona from the Department of Embryology and reproductive biology,
Komarov Botanical Institute of RAS, Saint-Petersburg, Russia, presents an scrupulous visual
analysis of archesporial cells which are formed and gave rise to megaspore mother cells. The
meiotic divisions produced a linear tetrad of haploid megaspores and from one chalazal
megaspore a Polygonum-type embryo sac is formed. Under natural conditions the apomixis
phenomenon was hardly observed in genus Helianthus L. In addition, author shows how on
plants of CMS-lines a number of anomalies in development of female reproductive system
were detected, including such phenomena as total absence of embryo sac, apospory and
integumentary embryony. Lack of the main embryo sac and formation of additional
aposporous embryo sacs could be observed in the same ovule. Finally, investigation of the
early stages of ovule development showed that aposporous embryo sacs originated from the
same ovule subepidermal cells as a normal embryo sac.
In Chapter 4, entitled Genetics and Genomics Applied to Sunflower Breeding, Dr. Hopp
and colleagues from the Instituto de Biotecnologia, Centro de Investigaciones Veterinarias y
Agronomicas, Instituto Nacional de Tecnologia Agropecuaria, Hurlingham, Argentina,
present a thoughtful study about new breeding strategies based on molecular markers, like
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Preface xi
quantitative trait loci mapping, association mapping and genomic selection that are currently
being developed for commercial crop improvement. The need to increase efficiency and
precision, and save time, resources and efforts, has motivated the application of Marker
Assisted Selection (MAS) in sunflower breeding programs. Furthermore, nowadays, the focus
is on tolerance improvement to biotic and abiotic stresses and oil quality and yield increasing,
in order to reduce the gap between potential and actual sunflower production.
In Chapter 5, entitled Sunflower Genetic Resources Interspecific Hybridization and
Cytogenetics in Prebreeding, Drs. Atlagic and Terzic present a rigorous description of the
genus Helianthus by reviewing genetic resources, cytogenetic research and application in
breeding. Besides the review of available literature, research results of the Institute of Field
and Vegetable Crops are presented in detail since the establishment of its collection in 1980.
Experience collected during this period indicates the difficulties in collection maintenance,
interspecific crosses and isolation of desired genes. Nevertheless, genus Helianthus proved to
be a good source of material for the constant improvement of cultivated sunflower.
In Chapter 6, entitled Functional Genomics and Transgenesis Applied to Sunflower
Breeding, Dr. Hopp and colleagues from the Instituto de Biotecnologia, Centro de
Investigaciones Veterinarias y Agronmicas, Instituto Nacional de Tecnologia Agropecuaria,
Hurlingham, Argentina, introduce an interesting chapter where they analyze different
strategies which have been developed in the last decade from functional genomics and post
genomics disciplines to contribute to the elucidation of gene regulation and identification of
key metabolic pathways involved in the response to biotic and abiotic stresses in sunflower.
The state of the art of strategies for gene function, studies in silico and in planta, by stable
gene transfer or agroinfiltration in sunflower as well as in the model system for Asteraceae
species, lettuce, are discussed within the frame of their application in sunflower breeding.
In Chapter 7, entitled Disease Management in Sunflowers, Dr. Leite from Embrapa
Soybean, Brazil, presents an interesting approach regarding the most important sunflower
diseases and strategies for disease management. Sunflower can be affected by the presence of
diseases, which may, depending on climatic conditions that favor the occurrence of pathogens
and the infective process, lead to a significant reduction on yield and quality of product.
Disease management should be based on an integrated program, in order to give support for
the sustainability and competitiveness of the sunflower crop.
In Chapter 8, entitled Recent Advances for Developing Resistance against Plasmopara
Halstedii in Sunflowers, Dr. Radwan from the Department of Natural Resources and
Environmental Sciences, University of Illinois at Urbana-Champaign, Urbana, IL, USA,
presents an interesting visual approach to Downy Mildew disease, as one of the most
important diseases of sunflower, which leads to an economic yield loss. In last two decades,
different approaches of genetics and genomics have significantly contributed to better
understand sunflower-Plasmopara halstedii. This progress directed to development of
sunflower lines carrying resistance to different races of this pathogen.
In Chapter 9, entitled Effects of Crop Management on the Incidence and Severity of
Fungal Diseases in Sunflowers, Dr. Debaeke and colleagues from the Institut National de la
Recherche Agronomique (INRA), Toulouse, France, dissected the effects of crop
management on the incidence and severity of major fungal diseases in sunflower, including
downy mildew, phoma, phomopsis and sclerotinia. They deeply reviewed and discussed the
influence of sowing date, plant population, N fertilization, and irrigation on sunflower
diseases from numerous experiments conducted in France during the last twenty years. They
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Juan Ignacio Arribas xii
proposed indicators of canopy development and nutritional status that could be useful when
developing crop management systems with reduced chemical applications.
In Chapter 10, entitled Insect Pests of Sunflowers in Africa, Dr. du Plessis from the Unit
for Environmental Sciences and Management, North-West University, Potchefstroom, South
Africa, present a complete study regarding a number of insect species which have adapted to
cultivated sunflower as a source of food and have consequently become economically
important pests. However, although many insect species is associated with sunflower in
Africa, only few are considered to be of potential economic importance. Insects most
commonly reported as injurious to this crop, occur sporadically, but usually in high numbers.
The families Noctuidae, Tenebrionidae, Curculionidae, Pentatomidae and Orsillidae are the
most important. Various types of damage are caused to sunflower seedlings, but the damage
symptoms are specific to a particular pest species. Dr. du PLessis argues that these seedling
pests mainly constitute dusty surface beetles (Gonocephalum simplex), greater false wire
worms (Somaticus spp.), cutworms (Agrotis spp.) and ground weevils (Protostrophus spp.).
Total defoliation can be incurred by the Plusia looper, Trichoplusia orichalcea. Hemipterans
and the African bollworm, Helicoverpa armigera are the most important insect pests of
sunflower during the heading stages of crop development. Intensive feeding by hemipterans
during this development stage results in deformed heads, which delay flower opening. The
occurrence of the false chinch bug, Nysius natalensis on sunflower during the heading stage
onwards in South Africa, is similar to that of N. stali in Nigeria. Since high summer
temperatures prevail throughout the sunflower production area of South Africa and the most
favourable temperature range for N. natalensis development is between 26C and 38 C, the
potential for rapid population build-up by this pest during the sunflower production season is
good. It is likely that N. natalensis can become important in sunflower production in other
African countries with similar weather conditions too. The insect is polyphagous and a variety
of wild host plants, mainly weed species as well as crops such as grain sorghum play an
important role in sustaining its populations. Sunflower is not the preferred host but N.
natalensis lays its eggs on sunflower when its preferred host plants are removed or dead. This
behaviour explains the insects injuriousness to late-planted sunflower because weed species
have already senesced before the sunflower. This period often coincides with seed fill of late-
planted sunflower, providing an alternative for the insect for moisture, as well as seeds that
are necessary for reproduction of the pest. Weeding in and around sunflower during seed fill
of the crop, therefore results in destruction of the preferred host plants of N. natalensis, and
they consequently move to sunflower where they feed and cause damage. When considering
application of insecticides for control of this pest, it should take into consideration that N.
natalensis is highly mobile and continuous re-infestations could occur. Timing of insecticide
application is therefore important. African bollworm (H. armigera) is frequently present
during the reproductive stage of cultivated sunflower in Africa. The attractiveness of
sunflower to this pest is demonstrated by the traps use as a trap crop in and around organic
cotton fields in Tanzania. Larvae occur from the budding stage onwards. Levels of infestation
vary between localities and seasons, sporadically reaching epidemic proportions. Sunflower
has, however, the ability to compensate for head damage and along with the fact that
preferential feeding sites of H. armigera are not the achenes, a significant number of larvae
could be tolerated without any significant effect on yield. Actual damage is, therefore, the
only criterion that could be used in the determination of economic injury levels for control of
African bollworm on sunflower crop.
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Preface xiii
In Chapter 11, entitled Soil Amendments and Their Effects on Sunflower Growth, Dr.
Lopez and colleagues from the Centro de Investigacion en Biotecnologia Aplicada, Instituto
Politecnico Nacional, Tlaxcala, Mexico, present a novel approach to several forms of amend
or fertilize sunflowers as alternative to improves this important cultivar. In particular authors
are interested in the study of organic materials that could be applied to soil in order to
improve their properties and plant growth, keeping in mind that we must recycle or reuse this
kind of materials. Finally, the organic amendments could be a beneficial disposal approach
that must be considered.
In Chapter 12, entitled Nutrition and Fertilization of Sunflowers in Brazilian Cerrado,
Dr. Castro and colleagues present an interesting chapter centered in a particular region in
Brazil, which authors argue that is recognized as a major global food producer and despite the
fact that its agriculture occupies only less than 5 % of the national territory, the estimated
grain production for 2013/2014 growing season is 195 million tons. The Brazilian Cerrado is
the main agriculture expansion region in the country, driven by soybean cultivation in an area
of 13 million ha. In this tropical agricultural region, sunflower has great potential for
expansion and consolidation as an important component of sustainable crop rotation
production systems. This chapter addresses the major limiting soil fertility factors which
hinder the crop development and discusses fertilization management practices related to the
main limiting nutrients, like the macronutrients nitrogen, phosphorus and potassium and
micronutrients such as boron and molybdenum. Adequate management of soil acidity and
fertilization has been proved as a powerful tool to improve the natural conditions of acidic
and chemically poor arable tropical soils. In addition to the soil fertility assessment, leaf
analysis is essential for the proper interpretation of plant nutritional status, thereby enabling
better refinement of the crop nutritional management.
In Chapter 13, entitled Environmental Issues in the Sunflower Crop of Midwestern Brazil
Diversification And Complementarities in the Biodiesel Chain, Dr. Ramos and colleagues
from Embrapa Environment, Brazil, present an interesting study regarding the increase in
global demand for renewable energy, the production of oilseeds, including sunflower, as
feedstock for biodiesel. The increase in global demand for renewable energy has encouraged,
both directly and indirectly, the production of oilseeds, including sunflower, as feedstock for
biodiesel. In this scenario, authors argue that Brazil stands out for its excellent agronomic and
climatic conditions for growing these crops throughout its territory. Sunflower is considered
an interesting option and your production in the mid-western region of Brazil has done
valuable contributions as a second, and especially the production practices adopted by the
reference farmers of the country, rendering complementarities and diversification to both the
food and the bioenergy sectors.
In Chapter 14, entitled Micro and Macro-Morphological Variation of Cosmos Bipinnatus
and Cosmos Bipinnatus Var. Albiflorus in Sympatric Zones in Central Mexico, Dr. Paniagua
and colleagues from the Centro de Investigacion en Biodiversidad y Conservacion, Univ.
Autonoma Estado de Morelos, Morelos, Mexico, present a concise but at the same time
precise analysis of the morphological variations in various sunflower species in central
Mexico area. Mexico is considered one of the centers of diversification of the Asteraceae
family, which contains the greatest richness of flowering plants. Particularly, the Trans-
Mexican Volcanic Belt (TMVB) is a heterogeneous mountain belt, located in the central part
of the country in an eastwest direction, which has been considered a diversification site for
many genera due to the vast number of species that it contains, as well as its high degree of
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Juan Ignacio Arribas xiv
endemism. Cosmos bipinnatus is present in various types of vegetation along the TMVB
which is favored by disturbances. Taxonomic studies have documented that this sunflower
species presents white to lilac ligules. However, horticulturists have considered the white
variety as C. bipinnatus var. albiflorus. Still, there is no scientific evidence to support their
observations. Therefore, the goal of this study was to compare the micro and macro
morphology characters between C. bipinnatus individuals with white and lilac ligules to
determine a possible morphological differentiation between both phenotypes in sympatric
zones in the central Mexico region. Principal Component Analysis and Non-Metric
Multidimensional Scaling showed a clear morphological differentiation between both groups;
this pattern was consistent even when the ligules color was not considered for the statistical
analyses. Dr. Paniagua and colleagues results sign a possible speciation between these
phenotypes and support a taxonomic shift for the Mexican sunflower with white ligules to C.
bipinnatus var. albiflorus.
Juan Ignacio Arribas, PhD
Associate Professor of Electrical Engineering
University Valladoild, Spain
Valladolid, November 2013
Tel: +34 983423000
Fax: +34 983423667
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In: Sunflowers ISBN: 978-1-63117-347-9
Editor: Juan Ignacio Arribas 2014 Nova Science Publishers, Inc.
Chapter 1
AN INTRODUCTION TO THE SUNFLOWER CROP
Fabin Fernndez-Luqueo1,
, Fernando Lpez-Valdez2,
Mariana Miranda-Armbula2, Minerva Rosas-Morales
2,
Nicolaza Pariona1 and Roberto Espinoza-Zapata
3
1Natural Resources and Energy Group, Cinvestav-Saltillo, Coahuila, Mxico
2CIBA - Instituto Politcnico Nacional, Tepetitla de Lardizbal, Tlaxcala, Mxico
3Crop Breeding Department, UAAAN, Saltillo, Coahuila, Mxico
ABSTRACT
Sunflower (Helianthus annuus L.) belongs to the family Asteraceae. The Helianthus
genus contains 65 different species of which 14 are annual plants. The sunflower plant
originated in eastern North America. It is thought to have been domesticated around 3000
B.C. by Native Americans. In the late 1800s the sunflower was introduced in the Russian
Federation where it became a food crop and Russian farmers made significant
improvements in the way that the sunflower was cultivated. Since 3000 B.C. a wide
range of uses of sunflower have been reported throughout the world such as ornamental
plant, medicinal, alimentary, feedstock, fodder, dyes for textile industry, body painting,
decorations, and so on. Sunflower species are allelopathic in nature and this crop appears
to have a bright future, especially if the scientists can translate the cutting-edge research
into technologies that will reduce the reliance on synthetic herbicides, pesticides, and
crop protection chemicals. On the one hand sunflower is well known by its
phytoremediation potential, thus it can be speculated that the good tolerance of sunflower
towards pollutants coupled with an increased accumulation/degradation capacity might
contribute to an efficient removal of pollutants from soil and water; on the other hand
sunflower possesses the potential to develop bioenergy systems that allow for synergies
between food and energy production. Because the sunflower has several potential
markets, it is a good choice for growers on both small and large scales. However, it has to
be remembered that scientific, technical or agricultural projects linked with sunflower
have to include side effects elsewhere in order to shape a sustainable future.
* Corresponding author: F. Fernndez-Luqueo, Natural Resources and Energy Group, Cinvestav-Saltillo, Coahuila.
C. P. 25900, Mxico Tel.: +52 844 4389625; Fax: +52 844 4389610. E-mail address: cinves.cp.cha.luqueno@
gmail.com.
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F. Fernndez-Luqueo, F. Lpez-Valdez, M. Miranda-Armbula et al. 2
Keywords: Allelopathy, biodiesel, phytoremediation, renewable energy, sustainable
development, symbiosis
1. INTRODUCTION
Sunflower (Helianthus annuus L.) belongs to the family Asteraceae. Helianthus genus
contains 65 different species (Andrew et al., 2013). The name Helianthus, being derived from
helios (the sun) and anthos (a flower), has the same meaning as the English name Sunflower,
which has been given these flowers from a supposition that they follow the sun by day,
always turning towards its direct rays. The sunflower that most people refer to is H. annuus,
an annual sunflower. In general, it is an annual plant which possesses a large inflorescence
(flowering head), and its name is derived from the flower's shape and image, which is often
used to depict the sun. The plant has a rough, hairy stem, broad, coarsely toothed, rough
leaves and circular heads of flowers (Khaleghizadeh, 2011). The heads consist of many
individual flowers which mature into seeds on a receptacle base (Seghatoleslami et al., 2012).
Sunflower is the worlds fourth largest oil-seed crop and its seeds are used as food and its
dried stalk as fuel. It is already been used as ornamental plant and was used in ancient
ceremonies (Harter et al., 2004; Muller et al., 2011). Additionally, medical uses for
pulmonary afflictions have been reported. In addition, parts of this plant are used in making
dyes for the textile industry, body painting, and other decorations. Sunflower oil is used in
salad dressings, for cooking and in the manufacturing of margarine and shortening
(Kunduraci et al., 2010). Sunflower is used in industry for making paints and cosmetics. A
coffee type could be made with the roasted seeds. In some countries the seed cake that is left
after the oil extraction is used as livestock feed. In the Soviet Union the hulls are used for
manufacturing ethyl alcohol, in lining for plywood and growing yeast. The dried stems have
also been used for fuel. The stems contain phosphorous and potassium which can be
composted and returned to soil as fertilizer. Sunflower meal is a potential source of protein
for human consumption due to its high nutritional value and lack of anti-nutritional factors
(Fozia et al., 2008).
Sunflower was a common crop among American Indian tribes throughout North
America. Evidence suggests that the plant was cultivated by natives in present-day Arizona
and New Mexico about 3000 B.C. Some archaeologists suggest that sunflower may have been
domesticated before corn (NSA, 2013). Although the scientific consensus had long been that
sunflower was domesticated once in eastern North America, the discovery of pre-Columbian
sunflower remains at archaeological sites in Mexico led to the proposal of a second
domestication center in southern Mexico. However, evidences from multiple evolutionary
important loci and from neutral markets support a single domestication event for extant
cultivated sunflower in eastern North America (Blackman et al., 2011).
The objective of this chapter is to present and discuss a summary about the huge amount
of information in which the sunflower is the main subject. The chapter aims to assist people
involved in all aspects of sunflower management, including conservation, agriculture, mining,
energy, food production, health and other industries, to obtain a broad knowledge of
sunflower and of its ecosystem services.
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An Introduction to the Sunflower Crop 3
2. BOTANICAL AND MORPHOLOGICAL DESCRIPTION
Sunflowers are botanically classified as Helianthus annuus L. (Table 1). They are large
plant and are grown throughout the world because of their relatively short growing season.
Sunflower is an annual herb, with a rough, hairy stem, 3 to 12 feet high, broad, coarsely
toothed, rough leaves, 3 to 12 inches long and circular heads of flowers, 3 to 6 inches wide in
wild specimens and often a foot or more in cultivation. The flower-heads are composed of
many small tubular flowers arranged compactly on a flattish disk: those in the outer row have
long strap-shaped corollas, forming the rays of the composite flower. Each sunflower head, or
inflorescence, is actually composed of two types of flowers. What appears to be yellow petals
around the edge of the head are actually individual ray flowers. The face of the head is
comprised of hundreds of disk flowers, which each form into a seed (achene).
The basic chromosome number for the Helianthus genus is 17. Diploid, tetraploid and
hexaploid species are known. There are only 14 annual species of Helianthus. Plant breeders
have made interspecific crosses within the genus and have transferred such useful characters
as higher oil percentage, cytoplasmic male sterility for use in production of hybrids, and
disease and insect resistance to commercial sunflower.
Table 1. Scientific classification of H. annuus L.; this genus counts 65 different species
Taxa
Kingdom Plantae
Subkingdom Viridaeplantae
Infrakingdom Streptophyta
Division Tracheophyta
Subdivision Spermatophytina
Infradivision Angiospermae
Class Magnoliopsida
Superorder Asteranae
Order Asterales
Family Asteraceae
Subfamily Helianthoideae
Tribe Heliantheae
Genus Helianthus
Specie annuus
The taxonomic classification has been in place since 1753.
3. PRODUCTION
In recent years, the sunflower cultivated area has been steadily increasing due to the
breeding of dwarf high yielding hybrids that also facilitate mechanization and the emphasis
given to polyunsaturated acids for human consumption. Global production grew steadily in
last 25 years (PSD-USDA, 2011), and FAO expect a total world output close to 60 million
tons towards 2050. The four largest producers (Russia, Ukraine, European Union and
Argentina) account for 70% of global volume, with an exponential growth of production in
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the last ten years in the Black Sea region, with increased acreage an higher yields achieved by
the replacing old varieties by hybrid seeds.
According to data from FAOSTAT (FAOSTAT, 2011) Russia Federation ranked first
producing ca. 9.7 millions of tons of sunflower seeds or 26% of the world total. Ukraine and
Argentina ranked second and third place with 8.6 and 3.6 tons of sunflower seeds,
respectively. France, Romania, China, Bulgaria, Hungary, Turkey, and Spain produced
between 1.0 and 1.9 millions of tons of sunflower seeds (Table 2). The United States
produced ca. 1.0 millions of tons of sunflower seeds, or 5% of the worlds total production.
That is enough to make the United States rank eleventh in that category. South Africa ranked
twelfth producing ca. 0.9 millions of tons of sunflower seeds.
Table 2. The highest twelve sunflower seed producing countries
in the world during 2011
Place Countries Production (tons)
1 Russia Federation 9,696,450
2 Ukraine 8,670,500
3 Argentina 3,671,750
4 France 1,882,450
5 Romania 1,789,330
6 China 1,700,000
7 Bulgaria 1,439,700
8 Hungary 1,374,780
9 Turkey 1,335,000
10 Spain 1,084,300
11 United States of America 924,550
12 South Africa 860,000
Russia followed by Ukraine are harvesting almost half of the world sunflower seed production. The
total sunflower seed production is reaching ca. 35 millions of tons
Data source: data obtained from FAOSTAT (2011).
According to FAO (FAO, 2010), there are some key production parameters which have
to be known by farmers throughout the world:
Sunflowers are grown in warm to moderate semi-arid climatic regions of the world from Argentina to Canada and from central Africa to the Commonwealth of
Independent States (Esmaeli et al., 2012; Onemli, 2012).
Frost will damage sunflowers at all stages of growth. The plant grows well within a temperature range of 20-25C; temperatures above 25C reduce yields and oil
content of the seeds (Thomaz et al., 2012).
Plants are drought-resistant, but yield and oil content are reduced if they are exposed to drought stress during the main growing and flowering periods. Sunflowers will
produce moderate yields with as little as 300 mm of rain per year, while 500-750 mm
are required for better yields (Gholamhoseini et al., 2013; Ghaffari et al., 2012).
Sunflowers adapt to a wide variety of soil, but perform best on good soils suitable for maize or wheat production (Radanielson et al., 2012).
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An Introduction to the Sunflower Crop 5
Sunflower plant density of 5-8 plants per m2 is required to form the optimum leaf area for plant photosynthesis. Kernel weight (40-80 g per 1000 kernels) and the
average number of kernels in a sunflower head (1200-1500) are the other most
important yield component (Seassau et al., 2012; Emami-Bistghani et al., 2012).
Sunflower growth depends more on nitrogen than any other nutrient. Due to its deep rooting system, sunflower is able to use nitrogen from soil layers that are inaccessible
to wheat, corn or other field crops. The plant requires a maximum of 150 kg of
nitrogen per hectare to produce a three tons ha-1
yield. Over fertilization may lead to
sunflower lodging. Phosphorous, potassium, boron, magnesium and molybdenum are
also needed to achieve the best yields (Jabeen and Ahmad, 2012; Babaeian et al.,
2011).
The average fatty acid composition of oil from temperate sunflower crops is 55-75% linoleic acid and 15-25% oleic acid. Protein content is 15-20% (Aznar-Moreno et al.,
2013; Ali and Ullah, 2012).
Planting in the Western Balkan countries, Eastern Europe and countries of the Former Soviet Union takes place during March and April (Zheljazkov et al., 2012;
Saleem et al., 2008).
Sunflower has one of the shortest growing seasons of the major economically important crops of the world. Early maturing varieties are ready for harvesting 90 to
120 days after planting, and late maturing varieties 120 to 160 days after planting.
Delayed harvesting causes unwelcome changes in oil quality, with an increase in free
fatty acid content. The seeds are ready to harvest when the heads turn black or brown
and the seed moisture content reaches 10-12%. Grain combines are fairly easily
adapted for the harvesting of sunflower by the addition of a head snatcher (Borbely et
al., 2008).
Depending on climatic and cultivation conditions, yields can vary from as much as 600 to 3000 kg ha
-1; irrigation is a key factor for obtaining high yields (Chigeza et
al., 2013; Khan et al., 2013; Akhtar et al., 2012).
Table 3 shows the oil yields in gallons per acre of oil producing crops, the yields will
vary in different agroclimatic zones. Sunflower produces 98 Gal oil acre-1
. That is enough to
make the sunflower rank twenty-third in that category. Additionally, higher-yielding oil crops
like safflower, mustards and sunflower have significant rotational benefits. For example, deep
safflower and sunflower roots help break up hardpan and improve soil tilth.
4. GROWTH AND DEVELOPMENT
Sunflower is a broadleaf plant that emerges from the soil with two large cotyledons
(Rawat et al., 2010). The emergence will take four to five days when planted an inch deep in
warm soil, but will take a few days longer in cooler soils or when planted deeper. Soil
crusting can make it difficult for the large seedlings to push out of the soil. Sunflowers grow
rapidly, producing large and rough leaves. Current sunflower varieties reach an average
height of six feet, varying between five and seven feet depending on planting date and soil
conditions (Saensee et al., 2012). After reaching their full height and blooming, heads on
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commercial cultivars turn downwards, designed to make it harder for birds to eat the seed.
Commercial sunflowers have flowers that are self-compatible for pollination, meaning they
do not require a pollinating insect, although some studies have shown bee pollinators
providing a slight yield boost (de Carvalho and de Toledo, 2008). Some farmers prefer
sowing their rows from north to south so that the capitula can lean into the row space, rather
than bumping against an adjacent plant, causing some seed to fall (Olowe and Adeyemo,
2009).
Table 3. Oil producing crops
Number Crop Scientific name Yield (Gal oil acre-1
)
1 Oil palm Elaeis guineensis Jacq. 610
2 Macauba palm Acrocomia aculeata Jacq. 461
3 Pequi Caryocar brasiliense Camb. 383
4 Buriti palm Mauritia flexuosa L. 335
5 Oiticia Licania rigida Benth 307
6 Coconut Cocos nucifera L. 276
7 Avocado Persea americana Mill. 270
8 Brazil nut Bertholletia excelsa Humb & Bonpl. 245
9 Macadamia nut Macadamia ternifolia F.V. Muell. 230
10 Jatrofa Jatropha curcas L. 194
11 Babassu palm Orbignya martiana Mart. 188
12 Jojoba Simmondsia chinensis Link 186
13 Pecan Carya illinoensis Wangenh. 183
14 Bacuri Platonia insignis Mart. 146
15 Castor bean Ricinus communis L. 145
16 Ghoper plant Euphorbia lathyris L. 137
17 Pissava Attalea funifera Mart. 136
18 Olive tree Olea europea L. 124
19 Rapessed Brassica napus L. 122
20 Opium poppy Papaver somniferum L. 119
21 Peanut Arachis hypogea L. 109
22 Cocoa Theobroma cacao L. 105
23 Sunflower Helianthus annuus L. 98
24 Tung oil tree Aleurites fordii Hemsl. 96
Yields of common energy crops are associated with biodiesel production. This is not related to ethanol
production, which relies on starch, sugar, and cellulose content instead of oil yields.
Experiments have been carried out to improve the growth and development of sunflower
under natural or stress conditions (Gerardo et al., 2013; Nasim et al., 2011; Da Silva et al.,
2012). Naz and Bano (2013) reported that the adverse effects of salt stress on sunflower
growth could be alleviated by foliar application of salicylic acid alone or in combination with
Azospirillum and Pseudomonas inoculations (Table 4). Gholamhoseini et al. (2013) shown
that the application of Glomus musseae and Glomus hoi could be critical in the cultivation of
sunflowers under arid and semi-arid conditions, where water is the most important factor in
determining plant growth and yield. Additionally, Akbari et al. (2011) reported that
inoculating the sunflower seeds with plant-growth promoting rhizobacteria increased the
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An Introduction to the Sunflower Crop 7
qualitative and quantitative properties of sunflower significantly, as compared to the control
treatment.
Table 4. Recent uses of the sunflower during the last years; main or alternative uses
make evident the diversity of sunflower
Area Description References
Food
Blends of high linoleic sunflower oil with
selected cold pressed soils.
(Ramadan, 2013)
Production of florets of sunflower. (Liang et al., 2013)
Tocopherols and phytosterols for the human
food market.
(Fernndez-Cuesta et al.,
2012)
Sunflower flour as a rich source of high
quality proteins.
(Levic et al., 2012)
Protein hydrolysis using proteases. (Tavano, 2013)
Animal Feed
Sunflower products fed to finishing pigs. (Gonzlez-Vega and Stein,
2012)
Ingestive behavior and physiological
responses of goats fed with sunflower cake.
(Agy et al., 2013)
Nutritional value of sunflower meal on broiler
chickens.
(Moghaddam et al., 2012)
Potential nutritive value as source of feed for
ruminants in Kenya.
(Osuga et al., 2012)
Energy
Methane production. (Fernndez-Cegr et al., 2013;
Todorovic et al., 2013)
Biodiesel production. (Iriarte and Villalobos, 2013;
Iglesias et al., 2012)
Bioenergy: biotechnology progress and
emerging possibilities.
(Gonzlez-Rosas et al., 2013)
Anaerobic digestion of sunflower oil cake. (De la Rubia et al., 2013)
Oil production. (Spinelli et al., 2012)
Sustainability
Of sunflower cultivation within the EU
Renewable Energy Directive.
(Spugnoli et al., 2012)
Sustainable sunflower processing. (Weisz et al., 2013)
Economic sustainability of sunflower
production.
(Keskin and Dellal, 2011)
Symbiosis and Plant-Growth Promoting Rhizobacteria
Effect of arbuscular mycorrhizal inoculation
on sunflower.
(Naz and Bano, 2013; Audet
and Charest, 2013;
Gholamhoseini et al., 2013)
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Table 4. (Continued)
Area Description References
Symbiosis and Plant-Growth Promoting Rhizobacteria
Bacterial inoculation speeds zinc release from
ground tire rubber.
(Khoshgoftarmanesh et al.,
2012)
A strain of Bacillus subtilis stimulates
sunflower growth.
(Lpez-Valdez et al., 2011)
Remediation
Biodegradation of PAHs. (Tejeda-Agredano et al.,
2013)
Plant response to lead. (Doncheva et al., 2013)
Metal accumulation on sunflower. (Mahmood et al., 2013; Hao et
al., 2012)
Fertilization, pesticides and environment
Foliar fertilization with molybdenum. (Skarpa et al., 2013)
Fertilization affects the agronomic traits of
high oleic sunflower hybrid.
(Mohammadi et al., 2013)
Gas exchange in sunflower plants. (Da Silva et al., 2013)
Effect of different nitrogen level on yield
components.
(Rafiei et al., 2012)
Biological control
Encrusting offers protection against
phytotoxic chemicals.
(Szemruch and Ferrari, 2013)
Biological control of Macrophomina
phaseolina on sunflower.
(Ullah, 2010)
Allelopathic effects
On growth of rice and subsequent wheat crop. (Bashir et al., 2012)
On seed germination and seedling growth of
Trianthema portulacastrum.
(Rawat et al., 2012)
Health
In vivo evaluation of an oral health toothpaste
with sunflower oil.
(Schafer et al., 2007)
Health benefits of the sunflower kernel. (Holliday and Phillips, 2001)
5. SUNFLOWER ALLELOPATHY
Sunflower species are allelopathic in nature; as well cultivated sunflower has great
allelopathic potential and inhibits weed-seedling growth of velvet leaf, thorn apple, morning
glory, wild mustard and other weeds (Macas et al., 1998a). Two members of the genus
Helianthus contain a great quantity of allelopathic compounds. H. annuus is well known for
its allelopathic compounds, including sesquiterpene lactones, heliespirones A, annoionones,
helibis-abonols and heliannols (Macas et al., 1998b). Heliannols A, D and E have special
relevance due to high phytotoxic activity (Macas et al., 1999).
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An Introduction to the Sunflower Crop 9
Figure 1. Some molecular structures of allelopathic compounds presents in sunflower cultivars. A)
Annuithrin (sesquiterpene lactone) or Niveusin C, a growth inhibitor. B) Furanoheliangoline, a
biologically active molecule. C) Germacranolide, a toxic sesquiterpene lactone (a potent feeding
deterrents).
Helianthus tuberosus contains helian-gine and H. annuus contains a sesquiterpene
lactone; a heliangolide [Annuithrin or Niveusin C (Figure 1A)] (a growth inhibitor);
furanoheliangolide [(Figure 1B) a biologically active]; three additional sesquiterpene
lactones: the known compound niveusin B, a germacranolide (Figure 1C) (the tifruticin-type);
a 3-ethoxy-niveusin B; an ethoxyheliangolide (Spring et al., 1982) and coumarins (only
accumulate in healthy sunflower plants as a response to the variation in environmental
conditions that affect field-grown plants). In sunflower, it was reported that the concentrations
of scopolin exceeded those in both infected and uninfected plants (Gutirrez-Mellado et al.,
1996).
Scopoletin have been described as phytoalexins and allelopathic compounds, being
accumulated in response to fungal and parasitic plant infection, insect attack, mechanical
injury and treatment with abiotic elicitors such as sucrose and CuCl2, and plant hormones;
besides scopoletin has also been shown to have a physiological activity, including the
promotion of stomatal closure in sunflower and inhibition of bud growth in pea at very low
concentrations (Gutirrez-Mellado et al., 1996).
Annuithrin was tested using a bioassay with Avena straight growth test. The addition of a
concentration range from 50 to 180 M resulted in a linear reduction of growth between 10
and 90%. In fact, annuithrin was shown to have antibacterial qualities. However, fungi and
yeast were either less inhibited or not inhibited (minimal inhibitory concentration, MIC 45 g
mL-1
on Bacillus brevis; MIC 90 g mL-1
on Proteus vulgaris; MIC 90 g mL-1
on
Eremothecium ashbyi; Macas et al., 1996). In addition, in vivo DNA and RNA synthesis in
cells of the ascitic form of Ehrlich carcinoma was drastically reduced by annuithrin (at an
annuithrin concentration of 20 g mL-1
about 50% inhibition of DNA synthesis and about
75% inhibition of RNA synthesis) (Spring et al., 1981).
It is well known that there are examples of allelopathic cover crops being used for weed
management in other crops, as well as other cultural methods to employ allelopathy (Duke,
2010). However, there are still no cultivars of crops being sold with allelopathic properties as
a selling point (Cheema and Khaliq, 2000; Tesio and Ferrero, 2010). Enhancement or
impartation of allelopathy in crops through the use of transgenes could eventually be used to
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produce such a cultivar. The study of allelopathic crops appears to have a bright future,
especially if the scientists can translate the cutting-edge research into technologies that will
reduce the reliance on synthetic herbicides, pesticides, and crop protection chemicals. Tesio
and Ferrero (2010) reported that the use of allelopathic traits from crops or cultivars with
important weed inhibition qualities, together with common weed control strategies, can play
an important role in the establishment of sustainable agriculture. It has to be noted that
allelopathy may also be another component of desired improved weed management. It will
not solve all weed problems in any field, but may help considerably to reduce the population
of weeds in the fields (Labrada, 2008).
6. PHYTOREMEDIATION WITH SUNFLOWER
Phytoremediation consists of mitigating pollutant concentrations in contaminated soils,
water, or air, with plants able to contain, degrade, or eliminate contaminants and its
derivatives (Malaviya and Singh, 2012). H. annuus is a plant with not only food and energy
values, but also with phytoremediation potential (Seth et al., 2011; Mukhtar et al., 2010). It is
one of the most widely studied plants for heavy metal phytoremediation (Kara et al., 2013).
However, it is well known that sunflower is able to contain, degrade or eliminate metals
(Chen et al., 2012; Ker and Charest, 2010; Lee and Yang, 2010), polycyclic aromatic
hydrocarbons (Tejeda-Agredano et al., 2013; Gan et al., 2009) and polychlorinated biphenyls
(Fiebig et al., 1997) from soil or water. Investigations with H. annuus have revealed that
several heavy metals, including lead, cadmium, copper, zinc and cobalt, accumulate at high
concentrations in shoots as well as in roots. Heavy metal uptake is minor in seeds than in
roots and shoots. However, few attempts have been made to use plant-growth promoting
rhizobacteria to facilitate phytoextraction and cadmium uptake in H. annuus planted in
cadmium-contaminated soil (Prapagdee et al., 2013). Sunflower is a documented metal
accumulator and its growth on contaminated soil for simultaneous remediation and further
energy production has been studied (Marques et al., 2013; Madejon et al., 2003). The good
tolerance of sunflower toward pollutants coupled with an increased accumulation/degradation
capacity might contribute to an efficient removal of pollutants from soil and water. Clearly it
is not an easy job, thus scientists of multidisciplinary areas have to work hard. Additionally,
there is a lack of knowledge concerning the pollutants accumulation and antioxidant
responses during the growth and development of sunflowers.
7. SUNFLOWER AS A RENEWABLE ENERGY SOURCE
Thousands of years ago, people in many regions throughout the world began to process
vegetable oils, utilizing whatever food stuffs they had on hand to obtain oils for a variety of
cooking purposes. The Chinese and Japanese produced soy bean oil as early as 2000 B.C.,
while southern Europeans had begun to produce olive oil by 3000 B.C. In Mexico and North
America, sunflower seeds were roasted and beaten into a paste before being boiled in water;
the oil that rose to the surface was skimmed off (FAO, 2010). During the last decade, an
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An Introduction to the Sunflower Crop 11
increased attention would be observed being paid on the use of sunflower as renewable
energy source.
Oilseed sunflower is quickly gaining popularity as a feedstock crop for biodiesel because
it shares several positive agronomic features with other common oil crops such as canola and
soybean; yields well in a variety of conditions, and can be grown easily and profitably at both
small farm and large field scales. It is well known that a number of crops can be used for both
food and bioenergy production such as sunflower (Kibazohi et al., 2012). Under some
circumstances, the potential exist to develop bioenergy systems that allow for synergies
between food and energy production. Integrated food and energy systems could produce food
crops while simultaneously addressing energy needs (Bogdanski et al., 2010).
There is a trend world-wide to grow crops in short rotation or in monoculture (such as
sunflower), particularly in conventional agriculture (Bennett et al., 2012). This practice is
becoming more prevalent due to a range of factors including economic market trends,
technological advances, government incentives, and retailer and consumer demands. Land-
use intensity will have to increase further in future in order to meet the demands of growing
crops for both bioenergy and food production, and long rotations may not be considered
viable or practical. Notwithstanding, evidence indicates that crops grown in short rotations or
monoculture often suffer from yield decline compared to those grown in longer rotations or
for the first time (Zambrano-Navea et al., 2012). Numerous factors have been hypothesized as
contributing to yield decline, including biotic factors such as plant pathogens, deleterious
rhizosphere microorganisms, mycorrhizas acting as pathogens, and allelopathy or autotoxicity
of the crop, as well as abiotic factors such as land management practices and nutrient
availability (Sun et al., 2011). This section identifies gaps in our understanding about the
energy production of biomass and the interaction of the ecosystems. Additionally, it has to be
remembered that each bioenergy development projects have to include side effects elsewhere
in order to shape a sustainable future.
CONCLUSION
Sunflower was domesticated in eastern North America and since 3000 B.C. this crop was
bred by natives. Thenceforth a wide range of uses of sunflower have been reported
throughout the world. Sunflowers are a permanent source of food, oilseed and biofuels
because they are well adapted to a variety of conditions and often require fewer agricultural
inputs than other more common crops, while under some circumstances, the potential exist to
develop bioenergy systems that allow for synergies between food and energy production.
Because the sunflower has several potential markets, it is a good choice for growers in both
small and large scales. However, scientific, technical or agricultural projects linked with
sunflower have to include environmental side effects such as pollution, greenhouse gases
emissions, salinization, or energy consumption elsewhere in order to shape a sustainable
future.
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In: Sunflowers ISBN: 978-1-63117-347-9
Editor: Juan Ignacio Arribas 2014 Nova Science Publishers, Inc.
Chapter 2
FLORAL BIOLOGY OF SUNFLOWERS:
A HISTOLOGICAL AND PHYSIOLOGICAL ANALYSIS
Basudha Sharma, Rashmi Shakya and Satish C. Bhatla* Laboratory of Plant Physiology and Biochemistry, Department of Botany,
University of Delhi, Delhi, India
ABSTRACT
The development of sunflower inflorescence can be considered under three phases,
namely inflorescence initiation, floret development and anther formation. Floret
primordia appear at the rim of the receptacle where ray or disc florets are generated. Disc
florets are arranged in Fibonacci series whereby a spiral pattern emerges as new florets
arise in rows of bumps consisting of a bract and a floret. Floral morphogenesis in
sunflower occurs according to the ABC model, whereby genes of the MADS box are
activated. Anthesis of disc florets is a phytochrome-mediated response and is also
modulated by plant hormones, such as auxins. The disc florets are hermaphrodite and
protandrous in nature, whereas the ray florets are sterile, incomplete and have an
attractive, fused and flag-like corolla. Stigma in sunflower is semi-dry in nature,
producing lipid rich exudates in the crevices of the adjacent papillae. Stigma undergoes
physiological maturity with the passage of development from bud, staminate and, finally
to the pistillate stage. The production of extracellular lipid rich secretions is initiated at
the staminate stage of stigma development and increases at the receptive stage through
the availability of elaioplasts and endoplasmic reticulum network in the basal regions of
the papillae. Transfer cells, earlier identified only in the wet type of stigma, are also
present in the transmitting tissue of sunflower stigma. Neutral esters and triacylglycerols
(TAGs) are the major lipidic constituents in pollen grains and stigma, respectively.
Lignoceric acid (24:0) and cis-11-eicosenoic acid (20:1) are specifically expressed only
in the pollen coat. Similar long-chain fatty acids have earlier been demonstrated to play a
significant role during the initial signalling mechanism leading to hydration of pollen
grains on the stigma surface. Lipase activity is expressed both in the pollen grains and
stigma papillae. Stigma exhibits a better expression of acyl-ester hydrolase activity the
pollen grains. Specific expression of lignoceric acid (24:0) in the pollen coat and
* Corresponding author: Professor S.C.Bhatla; E mail: [email protected].
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Basudha Sharma, Rashmi Shakya and Satish C. Bhatla 20
localization of lipase in pollen and stigma are likely to have possible roles during pollen-
stigma interaction. During the course of stigma development in sunflower, a correlation is
evident in the accumulation of reactive oxygen species (ROS), nitric oxide (NO) and the
activities of ROS scavenging enzymes [superoxide dismutase (SOD) and peroxidase
(POD)]. Mn-SOD (mitochondria localized) and Cu/Zn-SOD (cytoplasmic) exhibit
differential expression during the staminate stage of stigma development. An increase in
total SOD activity at the staminate stage is followed by a