Published 1978

SUNFLOWER SCIENCE AND TECHNOLOGY

 AGRONOMY
A Series 0/ Monographs
The American Society of Agronomy (ASA) and Academic Press published the first six books in this
series. Subsequent books were published by ASA alone, but in 1978 the associated societies, ASA, Crop Sci-
ence Society of America (CSSA), and Soil Science Society of America (SSSA), published Agronomy 19. The
books numbered 1 to 6 on the list below are available from Academic Press, Inc., III Fifth Avenue, New
York, NY 10003; those numbered 7 to 19 are available from ASA, 677 S. Segoe Road, Madison, WI 53711.

General Editor Monographs 1 to 6, A. G. NORMAN

1. C. EDMUND MARSHALL: The Colloid Chemistry of the Silicate Minerals, 1949
2. BYRON T . SHAW, Editor: Soil Physical Conditions and Plant Growth, 1952
3. K. D. JACOB: Fertilizer Technology and Resources in the United States, 1953
4. W. H. PIERRE and A. G. NORMAN, Editor: Soil and Fertilizer Phosphate in Crop Nutrition, 1953
5. GEORGE F. SPRAGUE, Editor: Com and Com Improvement, 1955
6. J. LEVITT: The Hardiness of Plants, 1956
---------------------
7. JAMES N. LUTHIN, Editor: Drainage of Agricultural Lands, 1957
General Editor, D. E. Gregg
8. FRANKLIN A. COFFMAN, Editor: Oats and Oat Improvement
Managing Editor, H. L. Hamilton
9. C. A. BLACK, Editor-in-Chie!. and D. D. EVANS, J. L. WHITE, L. E. ENSMINGER, and F. E.
CLARK, Associate Editors: Methods of Soil Analysis, 1965.
Part I-Physical and Mineralogical Properties, Including Statistics of Measurement and Sampling
Part 2-Chemical and Microbiological Properties
Managing Editor, R. C. Dinauer
10. W. V. BARTHOLOMEW and F. E. CLARK, Editor: Soil Nitrogen, 1965
Managing Editor, H. L. Hamilton
11. R. M. HAGAN, H. R. HAISE, and T. W. EDMINSTER, Editors: Irrigation of Agricultural Lands,
1967
Managing Editor, R. C. Dinauer
12. R. W. PEARSON and FRED ADAMS, Editors: Soil Acidity and Liming, 1967
Managing Editor, R. C. Dinauer
13. K. S. QUISENBERRY and L. P. REITZ, Editors: Wheat and Wheat Improvement, 1967
Managing Editor, H. L. Hamilton
14. A. A. HANSON and F. V. JUSKA, Editors: Turfgrass Science, 1969
Managing Editor, H. L. Hamilton
IS. CLARENCE H. HANSON, Editor: Alfalfa Science and Technology, 1972
Managing Editor, H. L. Hamilton
16. B. E. CALDWELL, Editor: Soybeans: Improvement, Production, and Use, 1973
Managing Editor, H. L. Hamilton
17. JAN VAN SCHILFGAARDE,Editor: Drainage for Agriculture,l974
Managing Editor, R. C. Dinauer
18. GEORGE F. SPRAGUE, Editor: Com and Com Improvement, 1977
Managing Editor, D. A. Fuccillo
19. JACK F. CARTER, Editor: Sunflower Science and Technology, 1978
Managing Editor, D. A. Fuccillo
SUNFLOWER SCIENCE
AN D TECH NOLOGY

EDITOR:

JACK F. CARTER

Editorial Committee
Gerhardt N. Fick Eric D. Putt
Donald L. Smith

Managing Editor
D. A. Fuccillo

Number 19 in the series

AGRONOMY

American Society of Agronomy,

Crop Science Society of America,
Soil Science Society of America, Inc., Publishers
Madison, Wisconsin, USA
1978
Copyright © 1978 by the American Society of Agronomy, Inc., Crop Science Society of
America, Inc., and Soil Science Society of America, Inc.

ALL RIGHTS RESERVED UNDER THE U.S. COPYRIGHT LAW OF 1978 (P.L. 94-553).
Any and all uses beyond the "fair use" provision of the law require written permission from
the publishers and/or the authors; not applicable to contributions prepared by officers or
employees of the U.S. Government as part of their official duties.

Library of Congress Cataloging in Publication Data

Sunflower Science and Technology.

(Agronomy, a series of monographs; no. 19)

Includes bibliographies and index.
1. Sunflower. I. Carter, Jack F. 1919-
II. Series.
SB299.S9S86 338.1 '7' 59335578-16192
ISBNO-89118-054-0

The American Society of Agronomy, Ins;.

677 S. Segoe Road, Madison, Wisconsin, USA 53711

Printed in the United States of America

Reprinted with minor revisions, 1980

Frontispiece. Top, a field of hybrid sunflower in North Dakota; bottom, a sunflower head
visited by a bumblebee pollinator.
v
A Sunflower Seeds

A. Black
B. Black with grey stripe
C. Black with white stripe
D. Dark grey with white stripe
E. Grey with white stripe
F. White with grey stripe
G. White

B c

vi
 FOREWORD

When I was a small boy in Tennessee, my grandmother planted a few

sunflower seeds in the corner of the garden. She said they were planted for
the three B's: beauty, bees, and birds. How those plants did grow. They
held dominion over all other plants in the garden because of their size and
beauty.
My next remembered encounter with sunflower came in an excellent
restaurant in Shanghai, China, at the end of World War II. There I dis-
covered that sunflower seeds were not just for birds, but a delicious toasted
and salted hors d' oeuvre with the English translation of "time killer." One
needed a tough fingernail and patience to open them.
During graduate school I learned from Kansans that the sunflower was
a state flower, originated in America, and occasionally was planted in large
fields for commercial purposes. I was surprised in the 1960's suddenly to
start hearing and reading about the high oil sunflower cultivars being devel-
oped and grown in the USSR and their introduction to Canada and the
USA.
The sunflower might be described as the Golden Girl of American
Agriculture. For the plant was native born, acclaimed first in foreign lands,
and then returned home to a tumultuous welcome by both growers and re-
searchers. This amazing saga of the sunflower has not yet ended.
The American Society of Agronomy, the Crop Science Society of
America, and the Soil Science Society of America are pleased to bring to
you this monograph on an increasingly important world food crop. We ex-
press our deepest appreciation to Editor Jack Carter and the many authors
and reviewers for contributing their expertise and time.

Madison, Wisconsin JOHNNY W. PENDLETON

February 1978 President
American Society oj Agronomy

vii
 GENERAL FOREWORD

Publication of this Sunflower Monograph is evidence of the growing

importance of this crop to agronomy and crop science in the USA. This
book has been evolving for several years, from the inception of the idea for
a monograph, its suggestion to the Society's Monograph Committee, and
now to its materialization into book form by the industrious members of the
editorial committee and their able chairman, Dr. Jack Carter.
The editors and authors of this book contributed vast efforts to pro-
duce it. Together with their coauthors, and with the assistance of reviewers,
a splendid text was written to describe the current status of sunflower re-
search, and the historical significance, taxonomy, production, pests, and
various other aspects of this important crop.
Sunflower Science and Technology is the 19th monograph in the series
prepared by the American Society of Agronomy since 1949. The first six
volumes were published by Academic Press, Inc., of New York, but since
1957 the society has become the publisher. A complete list of the titles in the
series may be found among these first pages. The monographs represent a
significant and continuing activity of the American Society of Agronomy,
its officers, and its approximately 10,000 members located in 100 countries
around the world.
The American Society of Agronomy is associated with the Crop Sci-
ence Society of America and the Soil Science Society of America. The so-
cieties share many objectives and activities in promoting these branches of
agriculture and scientific disciplines. Members of the societies contribute
generously of their time and talents in producing various publications, in-
cluding monographs, and in pursuing other activities in the interest of
human welfare.
Sunflower Science and Technology is the first monograph cosponsored
by the three societies under a new agreement by these organizations. The
book should be of great interest and benefit to researchers, teachers, stu-
dents, producers, and other users. The societies consider it as one of their
major contributions to mankind because of the worldwide adaptation of
sunflower as a field crop and its growing importance in the USA. Through
the presentation of up-to-date scientific and practical mater:ial on this sub-
ject, the societies aim to make sunflower an even more useful and widely
grown crop for the benefit of all people.
In behalf of the society members and myself in particular, I sincerely
thank Dr. Carter for his successful performance as editor, the editorial
committee members, the many authors, Domenic Fuccillo, managing
editor, and all others who have contributed directly or indirectly to the ac-
complishments of this worthy project.

Madison, Wisconsin MATTHIAS STELLY

April 1978 Editor-in-Chiej
ASA Publications

viii
 PREFACE

The colorful, single-headed sunflower of the Frontispiece is a valuable

and useful cousin of the native sunflower of North America. The native
sunflower was distributed widely across the Central Plains from north to
south and occurred rarely in much of the rest of the continent. The wide
genetic diversity of sunflower is still available in the indigenous popula-
tions, although population density and dispersion is decreased with in-
creased pressure of cultivation and urbanization, grazing by animals, etc.
The native sunflower was used by natives of North America, and archaeo-
logical evidence indicates large headed types with large seeds existed many
centuries ago. Sunflower was used for food in mixtures of cooked vegeta-
bles and in food "concentrates" by some native Americans of the Plains.
Sunflower was taken to Europe and Asia in the 16th century and
spread widely from west to east as an ornamental plant and as kitchen
garden food. It was selected first for large head and large seed types as a
food crop. Later, plant breeding techniques were used in the USSR and ad-
jacent countries to produce types with high oil percentage. Sunflower has
become the major annual oilseed crop of the USSR and other countries, in-
cluding Argentina.
The nonoilseed, confectionery, or food type sunflower was brought
back to North America by immigrants and cultivated in the Northern Plains
of North America. Sunflower has been grown in the Northern Great Plains
of the USA and in the Prairie Provinces of Canada for the last century. Pro-
duction areas fluctuated, but they were economically nonsignificant until
recent years. The introduction of the high oil types from the USSR, e.g.,
'Peredovik,' into Canada and then on to the USA, and favorable prices for
the oilseed sunflower for export mainly has led to sunflower production of
about 910,000 ha in the USA and 67,000 ha in Canada in 1977. Interest for
domestic use of sunflower oil as an edible salad and cooking oil, and in
manufactured products, because of its high unsaturation, is increasing. The
production first of improved, disease resistant, open-pollinated types, then
true hybrids from cytoplasmic male sterility/fertility restorer techniques in
the last 5 to 10 years has made sunflower economically competitive with
other crops. Recent surpluses of other crops in the Northern Plains of
North America have increased the competitiveness of sunflower in 1977 and
apparently for 1978.
The Monograph Committee of the American Society of Agronomy,
the Crop Science Society of America, and the Soil Science Society of Ameri-
ca and the Executive Committee of the societies considered the preparation
of a Sunflower Monograph for several years prior to authorizing its writing.
Sunflower has become much more important as a crop during the period of
planning and writing of the monograph, reaching approximately 910,000 ha
in 1977 in the USA, most of which is oilseed sunflower. The production of
sunflower is expected to increase further in 1978.
ix
 Since sunflower was an extremely minor crop in North America until a
few years ago, only a few plant and soil scientists had conducted research on
the crop. Botanists such as Charles B. Heiser, Jr., and taxonomists had con-
ducted the major studies about sunflower and provided the only detailed
publications, except for the pioneering research of Eric D. Putt and associ-
ates in Canada. Only a few scientists were available to write about sun-
flower in this monograph. Authors from public and private agencies, who
were distributed geographically as much as possible, were chosen to repre-
sent the various disciplines.
This first Sunflower Monograph in English is designed to cover the his-
torical aspects of the crop and the species, the economic and food function
of the crop, the usefulness of the products from sunflower, problems of
production and efforts made to improve sunflower, and its expected posi-
tion in commerce in North America and the world in the future. The mono-
graph is written to appeal to a wide audience in North America and to those
involved in sunflower production, merchandizing, processing, and sun-
flower improvement throughout the world.
The great contribution of writing and personal time by authors, the
editorial committee, and outside reviewers is acknowledged. Also acknowl-
edged is the special secretarial and other assistance of Mrs. Elaine Dobrinz,
Agronomy Department, North Dakota State University, Fargo. The Edi-
torial Committee and the societies also recognize the willingness of the
various public and private agencies to permit the authors and editorial com-
mittee to devote many hours of time to literature search, writing, and edi-
torial activities to produce this Sunflower Monograph. Lastly, the counsel
and advice of Domenic Fuccillo, managing editor, and of the staff at
Society Headquarters at Madison, Wisconsin, is acknowledged, as well as
the financial assistance of ASA, CSSA, and SSSA for various costs of the
Editor.

Fargo, North Dakota JACK F. CARTER

January 1978 Editor

JACK F. CARTER: B.S., University of Nebraska; M.S., Washington State University;

Ph.D., University of Wisconsin; Professor and Chairman, Agronomy Department, North
Dakota State University (NDSU), Fargo. editor of publications on sunflower and adminis-
trator in sunflower research programs by USDA-NDSU in North Dakota.
x
 CONTRIBUTORS

JEROME F. BESSER, Fish and Wildlife Service, U.S. Department of In-

terior, Building 16, Denver Federal Center, Denver, CO 80225
JACK F. CARTER, Agronomy Department, North Dakota State Universi-
ty, Fargo, NO 58102
DA VID W. COBIA, Agricultural Economics, North Dakota State Uni-
versity, Fargo, NO 58102
D. GORDON DORRELL, Agriculture Canada, Research Station, Box
3301, Morden, Manitoba ROG 110
HARRY O. DOTY, JR., Economics, Statistics, and Cooperatives Service,
U.S. Department of Agriculture, Washington, D.C. 20250
GERHARDT N. FICK, SIGCO Sunflower Products, Box 150, Brecken-
ridge, MN 56520
CHARLES B. HEISER, JR., Botany Department, Indiana University,
Bloomington, IN 47401
HARVEY J. HIRNING, Agricultural Engineering Department, North
Dakota State University, Fargo, NO 58102
JOHN A. HOES, Agriculture Canada, Research Station, Box 3301,
Morden, Manitoba ROG 110
VERNON L. HOFMAN, Agricultural Engineering Department, North
Dakota State University, Fargo, NO 58102
PAULDEN F. KNOWLES, Department of Agronomy, University of Cali-
fornia, Davis, CA 95616
JAMES R. LOFGREN, Dahlgren, Inc., Crookston, MN 56716
DARNELL R. LUNDSTROM, Agricultural Engineering Department,
North Dakota State University, Fargo, NO 58102
ERIC D. PUTT, Agriculture Canada, Research Station, Box 3301,
Morden, Manitoba ROG 110
ROBERT G. ROBINSON, Department of Agronomy and Plant Genetics,
University of Minnesota, St. Paul, MN 55108
RONALD T. SCHULER, Agricultural Engineering Department, Universi-
ty of Minnesota, St. Paul, MN 55108
J. T. SCHULZ, Department of Entomology, North Dakota State Universi-
ty, Fargo, NO 58102
DONALD L. SMITH, CallWest Seeds, P. o. Box 1428, Woodland, CA
95696
ERNEST D. P. WHELAN, Agriculture Canada, Research Station, Leth-
bridge, Alberta T11 4B1
DAVID E. ZIMMER, Science and Education Administration, U.S. Depart-
ment of Agriculture, Coastal Plains, Research Station, Tifton, GA 31794

xi
 Conversion Factors for English and Metric Units
To convert To convert
column 1 column 2
into column 2, Column 1 Column 2 into column I,
multiply by multiply by

Length
0.621 kilometer, km mile,mi 1.609
1.094 meter, m yard, yd 0.914
0.394 centimeter, cm inch, in. 2.540
Area
0.386 kilometer', km' mile2 ,mF 2.590
257.1 kilometer', km' acre, acre 0.00405
2.471 hectare, ha [Om km') acre, acre 0.405
Volume
0.00973 meter], m3 acre-inch 102.8
3.532 hectoliter, hi cubic foot, ft' 0.2832
2.838 hectoliter, hi bushel, bu 0.352
1.057 liter quart (liquid), qt 0.946
Mass
1.102 ton (metric) ton (English) 0.9072
220.5 quintal, q pound,lb 0.00454
2.205 kilogram, kg pound,lb 0.454
Yield or Rate
0.446 ton (metric)/hectare ton (English)/acre 2.242
0.892 kglha lb/acre 1.121
0.892 quintal/hectare hundredweight/acre 1.121
Pressure
14.22 kg/em' lb/inch', psi 0.0703
14.50 bar Ib/in.', psi 0.06895
0.9869 bar atmosphere, atm· 1.013
0.9678 kg/cm' atmosphere, atm· 1.033
14.70 atmosphere, atm· Ib/in.', psi 0.06805
• An "atmosphere" may be specified in metric or English units.
Temperature
1.80C+32 Celsius, C Fahrenheit, F 0.555(F - 32)

Note From the Editorial Committee:

Some Standard Sunflower Terminology for the Future

SUNFLOWER: the crop, not sunflowers, that is, discard the "s". Compare to wheat. corn,
soybean, cotton, etc.
OILSEED: sunflower grown primarily for extraction of edible oil from the seeds, plus other
supplementary uses.
NONOILSEED: sunflower grown primarily for human food, i.e., so-called confectionery
uses, or birdfood, other petfood, etc.
ACHENE: the ripened sunflower ovary.
"SEED": equivalent of achene in colloquial use.
KERNEL: the seed with hull or ovary wall removed (commonly called nutmeat, a term to be
discouraged).
TO HULL OR DEHULL: means removal of the hull or ovary wall from the seed or
"kernel"
CUL TIVAR: equals variety in this monograph.

xii
 CONTENTS
Page
FOREWORD ................................................. vii
GENERAL FOREWORD ...................................... viii
PREFACE................................................... ix
CONTRIBUTORS............................................ xi

Chapter 1 History and Present World Status .................... .

ERIC D. PUTT

Sunflower Among North American Indians ................... 1

Early Evidence. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Uses of Sunflower by the Indians .......................... 2
European Adoption ....................................... 3
First Introductions ...................................... 3
Spread through Europe.. . . . .. . . .. . . . . . . . . ... . .. . . . . . . . . . . 4
Evolution as a Crop in Russia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Evolution as a Crop in Other European Countries ... . . . . . . . . . 6
Present Importance in Europe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Sunflower in North America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Reintroduction from Europe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Early Uses. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Evolution of Sunflower as an Oilseed Crop. . . . . . . . . . . . . . . . . . 9
Current Importance of Nonoilseed Types ................... 13
Sunflower in Other Continents. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14
Sunflower in Asia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14
Sunflower in Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16
Sunflower in Africa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17
Sunflower in South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19
Sunflower in the World Oilseed Economy . . . . . . . . . . . . . . . . . . . .. 20
World Production of Annual Oil seeds .. . . . . ... . ... . . ... . . .. 20
Continental Production of Sunflower. . . . . . . . . . . . . . . . . . . . . .. 20
Sunflower Production Ranked by Country . . . . . . . . . . . . . . . . .. 25
Sunflower in World Trade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 25

Chapter 2 Taxonomy of Helianthus and Origin of Domesticated

Sunflower .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 31
CHARLES B. HEISER, JR.

Generic Description. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 33
Generic Relations ......................................... 33
Key to Sections of Helianthus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 34
Key to Annual Species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 37
Key to Types of H. annuus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 44
xiii
 Archaeological Sunflower . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 45
Origin and Development of the Domesticated Sunflower. . . . . . . .. 49
Summary..... ......... . . .. ... ... . .... ..... . ...... . . ..... 52

Chapter 3 Morphology and Anatomy . . . . . . . . . . . . . . . . . . . . . . . . . .. 55

P.F.KNOWLES

Stern.. . . . . . . . . . ...... . ............. . . ..... . . . . . . . ..... .. 55

Dimensions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 55
Internode Elongation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 56
Anatomy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 58
Branching. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 60
Root...... . . . .. . . .. ...... . . . . . . . . .... . . . . ........ . . . . . .. 61
Dimensions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 61
Primary Growth ........................................ 62
Lateral Roots. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 62
Secondary Growth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 62
Leaves................................................... 63
Morphology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 63
Anatomy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 65
Involucral Bracts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 66
Flower Bracts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 66
Heliotropism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 66
Inflorescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 67
Floral Initiation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 67
Inflorescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 71
Ray Flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 72
Disk Flowers ........................................... 73
Anthesis ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 74
Anther and Pollen Development. . . . . . . . . . . . . . . . . . . . . . . . . .. 76
Male Sterility. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 76
Embryogenesis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 77
Achenes ................................................. 80
Pericarp ........ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 81
Seed................................................... 85

Chapter 4 Production and Culture. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 89

R. G. ROBINSON

Adaptation and Physiological Characteristics of the

Sunflower Plant. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 89
Light and Photosynthesis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 89
Respiration. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 90
Net Assimilation Rate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 91

xiv
 Temperature.... .......... .... ..... ....... ...... ........ 91
Photoperiod. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 92
Latitude Effects. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 92
Water Requirement. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 93
Drought Tolerance.... .... ..... ........... ....... ....... 93
Soil................................................... 94
Phenology ............................................. 95
Planting Practices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 95
Dates and Depth of Planting . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 96
Plant Populations and Planting Rates ...................... 98
Arrangement of the Population ............................ 101
Crop Sequence and Rotation ................................ 104
Volunteer Sunflower and Its Control . . . . . . . . . . . . . . . . . . . . . .. 105
Effect of Preceding Crop on Sunflower ..................... 106
Effect of Sunflower on the Following Crop. . . . . . . . . . . . . . . . .. 108
Soil Management. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 109
Tillage and Seedbed Preparation. . . . . . . . . . . . . . . . . . . . . . . . . .. 109
Elemental Composition and Requirements of Sunflower. . . . . .. 112
Absorption of Nutrients. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 114
Organic Matter Maintenance. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 115
Fertilizer Trials ......................................... 115
Fertilizer Placement ..................................... 116
Fertilizer Recommendations .............................. 118
Irrigation .......... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 118
Weed ControL ............................................ 124
Mechanical Control. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 125
Herbicides ............................................. 128
Pollination ............................................... 132

Chapter 5 Harvesting, Handling, and Storage of Seed ............. 145

R. T. SCHULER, H. J. HIRNING, V. L. HOFMAN, AND
D. R. LUNDSTROM

Harvesting Sunflower Seed ................................. 145

Harvesting Attachments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 146
Combine Adjustment .................................... 149
Field Loss . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 151
Combine Capacity ....................................... 151
Power Requirements ..................................... 152
Desiccants .............................................. 152
Drying and Handling Sunflower Seed ......................... 153
Drying Systems ......................................... 158
Moisture Determination. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 161
Seed Handling Systems ................................... 162
Storage of Sunflower Seed .................................. 164
Sunflower for Silage ....................................... 166

xv
Chapter 6 Insect Pests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 169
J. T. SCHULZ

Head-Infesting Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 170

Sunflower Moth, Homoeosoma electellum ..... . . . . . . . . . . . .. 170
European Sunflower Moth, Homoeosoma nebulella .......... 180
Banded Sunflower Moth, Phalonia hospes .................. 181
Sunflower "Budworm", Suleima helianthana ................ 183
Seed Weevils, Smicronyxjulvus and S. sordidus . ............. 189
A Sunflower Curculio, Haplorhynchites aeneus .............. 191
Sunflower Seed Midges. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 194
A Tephritid Complex .................................... 197
Foliage and Stem Feeding Species ............................ 201
Sunflower "Budworm"-Suleima helianthana . .............. 201
Sunflower Beetle-Zygogramma exc/amationis .............. 201
Painted Lady, C~nthis cardui ............................. 203
Sunflower Maggot, Strauzia longipennis . . . . . . . . . . . . . . . . . . .. 204
Stem Weevil, Cylindrocopterus adspersus . . . . . . . . . . . . . . . . . .. 207
Cerambycid Complex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 209
Root Infesting Insects ...................................... 210
Carrot Beetle, Bothynus gibbosus .................. -....... 210
A Root-Infesting Weevil, Baris strenua ..................... 214
Other Insect Pests ......................................... 216
Potential Pest Species ...................................... 217
(Color photos follow page 210)

Chapter 7 Diseases........................................... 225

D. E. ZIMMER AND J. A. HOES

Downy Mildew ............................................ 226

Causal Organism. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 226
Life Cycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 226
Symptomology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 227
Epidemiology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 228
Distribution and Economic Importance. . . . . . . . . . . . . . . . . . . .. 229
Races and Host Resistance. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 230
Control. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 231
Rust of Sunflower. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 231
Causal Organism. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 231
Symptomology. . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 232
Distribution and Economic Importance. . . . . . . . . . . . . . . . . . . .. 232
Life Cycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 233
Disease Cycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 233
Host Range .............................................. 234
Physiological Races and Host Resistance. . . . . . . . . . . . . . . . . . .. 234
Control. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 235
Sclerotinia Wilt and Head Rot. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 236
Causal Organism. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 236
xvi
 Host Range ............................................. 237
Life Cycle .............................................. 237
Symptomology .......................................... 238
Disease Cycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 238
Mechanism of Pathogenesis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 240
Distribution and Economic Importance ..................... 240
Survival of Sclerotia ..................................... 241
Control. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 242
Verticillium Wilt .......................................... 244
Causal Organism. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 244
Symptomology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 244
Distribution and Economic Importance. . . . . . . . . . . . . . . . . . . .. 245
Host Range and Specialization ............................ 246
Disease Cycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 246
Control ................................................ 247
Charcoal Rot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 249
White Blister Rust . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 249
Alternaria Diseases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 250
Phialophora Yellows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 251
Phoma Black Stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 252
Septoria Leaf Spot. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 252
Botrytis Head Rot or Grey Mold. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 252
Rhizopus Head Rot ........................................ 253
Damping-Off, Root Rots, and Stem Rots ..................... 253
Powdery Mildew . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 254
Bacterial Diseases ......................................... 254
Mycoplasms and Viruses ................................... 255
(Color photos follow Page 258)

Chapter 8 Birds and Sunflower. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 263

JEROME F. BESSER

Bird Species that Damage Sunflower ......................... 264

Extent of Loss . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 267
Factors Affecting Loss ......... " ......................... 269
Protecting Sunflowers from Bird Damage. . . . . . . . . . . . . . . . . . ... 271
Cultural Practices ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 271
Mechanical Means of Crop Protection. . . . . . . . . . . . . . . . . . . . .. 272
Chemical Frightening Methods ............................ 275
Summary .. , . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 277

Chapter 9 Breeding and Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 279

GERHARDT N. FICK

Breeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 280
Objectives .............................................. 280
Breeding Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 280
xvii
 Methods for Improving Cultivars .......................... 280
Inbreeding and Heterosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 283
Methods for Producing Hybrids ........................... 290
Recurrent Selection for Improving Source Populations. . . . . . .. 295
Genetic Resources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 297
Breeding Techniques. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 302
Genetics ......... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 308
Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 308
Yield and Yield Components. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 308
Plant Growth Characteristics. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 309
Flower Characteristics ................................... 315
Seed and Kernel Characteristics. . . . . . . . . . . . . . . . . . . . . . . . . . .. 319
Disease Resistance. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 322
Insect Resistance ........................................ 325
Broomrape Resistance ................................... 328
Tolerance to Bird Depredation ............................ 328
Tolerance to Adverse Environments ........................ 328
Gene Linkages .......................................... 329
Progress in the United States and Canada and Challenges
for the Future. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 329

Chapter 10 Cytology and Interspecific Hybridization. . . . . . . . . . . . .. 339

ERNEST D. P. WHELAN
Chromosome Morphology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 339
Euploidy and Aneuploidy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 340
Microsporogenesis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 341
Megasporogenesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 346
Male Sterility. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 348
Mutagenesis .............................................. 351
Interspecific Hybridization ................................. 352
Hybridization Techniques ................................ 352
Interspecific Hybrids ..................................... 353
Section Annui (Table 2) .................................. 353
Section Ciliares Subsections Pumili and Ciliaris (Table 3) . . . . .. 356
Section Divaricati Subsection Divaricati (Table 4) ............ 358
Section Divaricati Subsection Gigantei (Table 5) .............. 362
Section Divaricati Subsection Microcephali (Table 6) . . . . . . . . .. 364
Section Divaricati Subsection Angustifoli (Table 7) ........... 365
Section Divaricati Subsection A trorubentes (Table 8) .......... 365

Chapter 11 Planting Seed Production ........................... 371

DONALD L. SMITH
Hybrid Seed .............................................. 372
Production Procedures ................................... 372
Seed Management ....................................... 380
Open-Pollinated Seed ...................................... 384
xviii
Chapter 12 Production Costs and Marketing ..................... 387
DAVID W. COBIA

Economics of Farm Production .............................. 387

Production Patterns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 387
A verage Yields. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 390
Production Costs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 391
Relative Profitability. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 392
The Marketing System ..................................... 392
Marketing Channel. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 392
The Pricing Mechanism .................................. 396
Grades and Standards. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 398
Major Industry and Research Organizations ................... 402
State and Federal Research and Extension Agencies ........... 402
Government Statistical Reporting Services ................... 403
The Sunflower Association of America ..................... 403
The International Sunflower Association .................... 404
The North Dakota State Sunflower Council ................. 404
National Cottonseed Products Association, Inc ............... 404

Chapter 13 Processing and Utilization of Oilseed Sunflower ........ 407

D. GORDON DORRELL

Seed ..................................................... 407

Chemical Composition ................................... 408
Oil ...................................................... 408
Physical Characteristics .................................. 408
Chemical Composition ................................... 408
Extraction and Processing of Oils .......................... 414
Margarine Production ................................... 418.
Utilization ............................................. 420
Nutrition ............................................... 422
Meal .................................................... 423
Chemical Composition ................................... 423
Utilization as Animal Feed ................................ 429
Concentrates and Isolates ................................. 430
Other Products ........................................... 436
Chapter 14 Sunflower for Confectionery Food, Birdfood,
and Petfood. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 441
JAMES R. LOFGREN

Processing Procedures ..................................... 441

Quality of Nonoilseed Sunflower ............................ 443
In-Shell Sunflower Seed .................................. 443
Hulling Sunflower Seed .................................. 446
Laboratory Methods to Evaluate Hulling Quality ............. 448
xix
 Nutritive Value of Sunflower Kernels ....................... 452
Uses for Sunflower Kernels in Cooking ..................... 453
Quality of Birdfeed (Petfood) Sunflower .................... 453
Byproducts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 454
Kernel Fragments ....................................... 454
Sunflower Meal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 454
Sunflower Hulls ......................................... 454

Chapter 15 Future of Sunflower as an Economic Crop in

North America and the World .. ..................... 457
HARRY O. DOTY, JR.

Changes in the Oilseed Industry .............................. 458

Increased Food Production ............................... 458
Relative Value of Oil and Meal . . . . . . . . . . . . . . . . . . . . . . . . . . .. 459
Synthetics. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 460
Substituting Fats and Oils ................................. 460
New Processing Methods ................................. 461
Polyunsaturated Fatty Acids . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 462
Linoleic Acid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 462
Oilseed Sunflower Markets ................................. 463
Food Fat Markets ....................................... 463
Industrial Oil Markets .................................... 472
World Trends in Fats and Oils Markets ..................... 473
High Protein Meal Markets ............................... 477
World Competition for Sunflower .... . . . . . . . . . . . . . . . . . . . . . .. 481
Soybean ............................................... 481
Oil Palm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 483
Peanut. ................................................ 484
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 485

Index ........................................................ 489

xx