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3499 P- FM 11/17/2000 8:45 AM Page i

PLANT BREEDING REVIEWS


Volume 20
3499 P- FM 11/17/2000 8:45 AM Page ii

Plant Breeding Reviews is sponsored by:


American Society for Horticultural Science
Crop Science Society of America
Society of American Foresters
National Council of Commercial Plant Breeders

Editorial Board, Volume 20


G. R. Askew
F. A. Bliss
M. Gilbert
3499 P- FM 11/17/2000 8:45 AM Page iii

PLANT BREEDING REVIEWS


Volume 20

edited by
Jules Janick
Purdue University

John Wiley & Sons, Inc.


NEW YORK / CHICHESTER / WEINHEIM / BRISBANE / SINGAPORE / TORONTO
3499 P- FM 11/17/2000 8:45 AM Page iv

This book is printed on acid-free paper.

Copyright © 2001 by John Wiley & Sons, Inc.

All rights reserved.

Published simultaneously in Canada.

No part of this publication may be reproduced, stored in a retrieval system or


transmitted in any form or by any means, electronic, mechanical, photocopying,
recording, scanning or otherwise, except as permitted under Sections 107 or 108 of the
1976 United States Copyright Act, without either the prior written permission of the
Publisher, or authorization through payment of the appropriate per-copy fee to the
Copyright Clearance Center, 222 Rosewood Drive, Danvers, MA 01923, (978) 750-8400,
fax (978) 750-4744. Requests to the Publisher for permission should be addressed to the
Permissions Department, John Wiley & Sons, Inc., 605 Third Avenue, New York, NY
10158-0012, (212) 850-6011, fax (212) 850-6008, E-Mail: PERMREQ @ WILEY.COM.

This publication is designed to provide accurate and authoritative information in regard


to the subject matter covered. It is sold with the understanding that the publisher is not
engaged in rendering professional services. If professional advice or other expert
assistance is required, the services of a competent professional person should be sought.

Library of Congress Catalog Card Number: 83-641963

ISBN 0-471-38788-6

ISSN 0730-2207

Printed in the United States of America

10 9 8 7 6 5 4 3 2 1
3499 P- FM 11/17/2000 8:45 AM Page v

Contents

List of Contributors vii

1. Dedication: Norman Willison Simmonds;


Plant Breeder, Teacher, Administrator 1
William Spoor and Frederick England

2. The Origin of Maize: Evidence for Tripsacum Ancestry 15


Mary W. Eubanks
I. Introduction 16
II. Theoretical Framework 18
III. Taxonomic Classification 22
IV. The Nature of the Evidence 31
V. Synthesis and Future Directions 55
Literature Cited 57

3. History of Public Onion Breeding Programs


in the United States 67
Irwin L. Goldman, Geoffrey Schroeck, and Michael J. Harvey
I. Introduction 68
II. History of Onion in the Americas 71
III. State Experiment Station Programs 95
IV. Future Prospects for Public Onion Breeding
in the United States 99
Literature Cited 101

v
3499 P- FM 11/17/2000 8:45 AM Page vi

vi CONTENTS

4. Current and Future Issues in Lettuce Breeding 105


Edward J. Ryder
I. Introduction 105
II. History of Lettuce Breeding 106
III. Breeding Efforts 107
IV. Lettuce Breeding for the Future 117
Literature Cited 130

5. Cactus Pear Domestication and Breeding 135


Candelario Mondragon Jacobo
I. Introduction 136
II. Origin and Early Development 138
III. Genetic Resources 141
IV. Breeding Objectives 145
V. Breeding Techniques 152
VI. Breeding Systems 158
VII. Summary and Future Prospects 161
Literature Cited 162

6. Somatic Hybridization and Applications


in Plant Breeding 167
Alexander A. T. Johnson and Richard E. Veilleux
I. Introduction 168
II. Recent Efforts in Somatic Hybridization 173
III. Conclusion 209
Literature Cited 212
Subject Index 227
Cumulative Subject Index 229
Cumulative Contributor Index 245
3499 P- FM 11/17/2000 8:45 AM Page vii

Contributors

Frederick England, Formerly of Scottish Plant Breeding Station, Pentland-


field, Midlothian, Scotland
Mary W. Eubanks, Department of Botany, Duke University, Durham, North
Carolina 27708-0338, [email protected]
Irwin L. Goldman, Department of Horticulture, 1575 Linden Drive, University
of Wisconsin–Madison, Madison, Wisconsin 53706,
[email protected]
Michael J. Harvey, USDA—Agricultural Research Service and Department of
Horticulture, 1575 Linden Drive, University of Wisconsin–Madison, Madi-
son, Wisconsin 53706
Candelario Mondragon Jacobo, Instituto Nacional de Investigaciones Fore-
stales y Agropecuarias, Guanajuato, Mexico
Alexander A. T. Johnson, Department of Horticulture, Virginia Polytechnic
Institute and State University, Blacksburg, VA 24061-0327
Edward J. Ryder, U.S. Agricultural Research Station, U.S. Department of
Agriculture, Agricultural Research Service, 1636 E. Alisal Street, Salinas,
California 93905, [email protected]
Geoffrey Schroek, Department of Horticulture, 1575 Linden Drive, University
of Wisconsin–Madison, Madison, Wisconsin 53706
William Spoor, Department of Biotechnology, Plant Science Division, Scot-
tish Agricultural College, West Mains Road, Edinburgh, EH9 3JG, Scotland
[email protected]
Richard E. Veilleux, Department of Horticulture, Virginia Polytechnic Insti-
tute and State University, Blacksburg, VA 24061-0327
Masahiko Yamada, Persimmon and Grape Research Center, National Institute
of Fruit Tree Science, Akitsu, Hiroshima 729-2494, Japan
Keizo Yonemori, Laboratory of Pomology, Graduate School of Agriculture,
Kyoto University, Sakyo-ku, Kyoto 606-8502, Japan, [email protected]
Zhao-Bang Zeng, Department of Statistics, North Carolina State University,
Raleigh, NC 27695-1932

vii
3499 P- FM 11/17/2000 8:45 AM Page viii

Norman W. Simmonds
3499 P-01 9/26/2000 1:07 PM Page 1

Dedication:
Norman Willison Simmonds
Plant Breeder, Teacher, Administrator
William Spoor
Department of Biotechnology, Plant Science Division, Scottish
Agricultural College, West Mains Road, Edinburgh, EH9 3JG, Scotland

Frederick England
Formerly of Scottish Plant Breeding Station, Pentlandfield,
Midlothian, Scotland

Norman W. Simmonds’s long and distinguished professional career has


centered around three major crops, namely, bananas, potatoes, and sugar
cane. Along the way, he has made additional substantial contributions
in other crops, to plant breeding in general, to general taxonomy, eco-
nomic botany, and tropical farming systems. In all these fields he has
contributed creatively and provocatively. His output has been enor-
mous, diverse, and profound.

CAREER

Norman Willison Simmonds was born in Bedford (England) on Decem-


ber 15, 1922. His father was a civil servant and his mother of Scottish
(Perthshire) farming stock, the Willisons, from whom he takes his mid-
dle name. Norman attended Whitgift School in Croydon from 1934 to
1940 and from there won an Open Exhibition (scholarship) to Downing
College, Cambridge. At school he was stimulated, as were so many bud-
ding botanists, by Cecil T. Prime, that notable teacher. While at Cam-
bridge, he was much influenced by Professor D. G. Catcheside, who

Plant Breeding Reviews, Volume 20, Edited by Jules Janick


ISBN 0-471-38788-6 © 2001 John Wiley & Sons, Inc.

1
3499 P-01 9/26/2000 1:07 PM Page 2

2 W. SPOOR AND F. ENGLAND

encouraged his interest in genetics and cytogenetics. After a distin-


guished undergraduate career, he was awarded a First class degree in the
Natural Sciences Tripos part II (Botany, with a strong bent to genetics
and plant breeding). In 1943 he was granted a Colonial Agricultural
Scholarship and studied at Cambridge and the Imperial College of Trop-
ical Agriculture (ICTA) in Trinidad and in 1944 obtained his B.A. from
the University of Cambridge. This was followed by an M.A. in 1948 and
later, in 1966, a Sc.D. The introduction to tropical agriculture afforded
by the time spent in Trinidad was to be the start of a life-time interest
in tropical agriculture, the crops and the demands, following his
employment as a Botany lecturer at ICTA in 1945. His stay in the West
Indies was to last for 15 years, ending in 1959 as Senior Cytogeneticist,
Banana Research Scheme. During this period he established himself as
a vigorous researcher, initially with K. S. Dodds, developing a banana
breeding strategy through constructed diploids crossed to triploids,
thence to tetraploids. It was also during this time that he started to
develop ideas on genetic resources, conservation, and utilization fol-
lowing two major collecting trips to East Africa (1948) and the Pacific,
Malaysia, Thailand, and North India (1954/55). Material from these trips
is now proving of value as later generation parents, but at the time led
to enhanced evolutionary understanding of the group. This extensive
experience led to two key books, the standard monograph Bananas
(1959, 1966, with R. H. Stover in 1987), long regarded as the banana
researcher’s bible, and Evolution of the Bananas (1962) in addition to
40 papers published during his period in Trinidad.
In 1959, Norman Simmonds returned to the United Kingdom as Head
of the Potato Genetics Department at the John Innes Institute at Hertford,
rejoining K. S. Dodds, the then Director. He characteristically threw
himself into research in this new crop challenge, publishing an array of
papers ranging over tuber dormancy, seed germination, polyploidy, cal-
lus differentation, virus transmission, chimeral and other mutants, link-
age studies, and disease resistance. It was during this period that he
developed the concept of genetic base broadening, now found to be fun-
damental and effective for potatoes. This has proved to be a valuable
recurring theme in much of his later work as it became obvious that such
an approach had general applicability to a very wide range of crop
species. It is one of Norman’s regrets that, while his ideas on base broad-
ening have received wide theoretical acceptance, they have been rela-
tively little applied in practice, although enjoying considerable success
in sugar cane improvement and to a lesser extent in oil palm and rub-
ber development. While at the John Innes Institute, he started two very
3499 P-01 9/26/2000 1:07 PM Page 3

1. DEDICATION 3

important consultancies: on West Indies sugar breeding in Barbados


and on rubber in Malaysia. This broadening of his crop interests allowed
him to bring experiences in other clonal crops to the benefit of these two
economically important tropical plantation crops. These consultancies
among others allowed him to build up a wide circle of friends and
acquaintances. He realized that breeders of particular crops have much
to gain from crosslinking with the experience of others. This developed
over the years and could be said to have culminated in the book Evolu-
tion of Crop Plants (1976), edited by Norman Simmonds but consisting
of the authoritative contributions of almost 86 authors, experts in their
field, in addition to six chapters contributed by himself. This extremely
popular book has done much to provoke interest in crop plants and the
systematic approach required by the editor (introduction, cytotaxonomic
background, early history, recent history, and prospects) has proved to
be particularly valuable to educators wherever crop evolution and breed-
ing is taught. The original book has now been replaced by a second edi-
tion (Smartt and Simmonds, 1995) which, in addition to the necessary
updating, has retained the much valued presentation formula.
In 1965, Norman Simmonds moved northwards nearer to his family’s
origin, when he took on the post of Director of the Scottish Plant Breed-
ing Station, then at Pentlandfield on the outskirts of Edinburgh. This
demanding role, at a time of increasing pressure for new varieties suited
to Scottish conditions left little time for personal research and many of
the ideas initiated at the John Innes Institute, had to be left to others to
pick up. However, he did make time to re-establish his education links,
developing teaching initiatives with the Botany Department of the Uni-
versity of Edinburgh and this thread would be carried through to the
final phase of his professional career when he joined the staff of the
Edinburgh School of Agriculture in 1976, then under the enlightened
direction of Professor Noel Robertson. This return to academia allowed
Norman to bring his knowledge and experience to bear in a wide range
of areas. One of his first activities was to produce a new and much
needed textbook on plant breeding (Principles of Crop Improvement,
1979). This provided students and practitioners with a synthesis of cur-
rent breeding approaches along with a bibliography that would allow the
motivated into the current literature in any particular area. This has
proved to be a very worthy and highly regarded addition to the texts cur-
rently available on plant breeding and, having recently undergone some
modifications, has appeared as a 2nd Edition (Simmonds and Smartt,
1999). During this period and continuing well after his formal retirement
in 1982 at the age of 60, he developed consultancies and took ever wider
3499 P-01 9/26/2000 1:07 PM Page 4

4 W. SPOOR AND F. ENGLAND

interests in tropical crops at large. He travelled widely throughout the


tropics, to some 20 countries, partly as Chairman of the Quinquennial
Review of the International Board for Plant Genetic Resources and later
on behalf of FAO and the World Bank. For the last he prepared a review
(1983/84) which was assimilated into Bank policy though not without
sociological controversy. In addition to the above he continued to make
valuable contributions to consultancies carried out for Sugar Cane Breed-
ing, in the West Indies; Rubber Research Institute of Malaysia; Booker-
Daehnfeldt, Denmark; and Copersucar, Brazil, to name but a few.
Although well into his retirement now, Professor Simmonds remains
no less active academically and continues to review and publish on a
wide range of topics, not only within his broad subject area, but also in
relation to his hobbies, most notably on trout fishing. Major recent
reviews by him have covered horizontal resistance to diseases on crops,
an interest originally fired by John Niederhauser almost 30 years before
in Mexico; potato propagation by seed as distinct from clonal propaga-
tion by tubers (a recently fashionable topic); tropical crops and their
improvement; and, most recently, an informal history of statistics for
Plant Breeding Reviews, Volume 17. His interest in statistics goes back
to the start of his academic training at University and has been demon-
strated throughout much of his published work, indicating both his lik-
ing for the subject and his growing appreciation of its importance for
biology and agriculture. Although largely self-taught, he has always
been able to focus on the principles and build upon these to produce a
clearer understanding of the importance and applicability of statistics
in plant genetics breeding and agriculture for the benefit of students and
practitioners.

THE MAN

Norman Simmonds is a unique and stimulating individual, with diverse


interests. He is an iconoclast, a gifted scientist and profound thinker, a
stimulating teacher, a loyal colleague and friend, and a kind and gener-
ous man who has been ably supported in all his endeavors by his lov-
ing wife, Christa. As a result of his teaching duties he has profoundly
affected hundreds of students and has directly influenced many people
now in senior positions in agricultural research and administration. His
many travels and consultancies have made him a well known and well
regarded figure in tropical agriculture and in many developing countries
of the world. He is a world authority in many areas and crops. As a result
3499 P-01 9/26/2000 1:07 PM Page 5

1. DEDICATION 5

of his skill as a writer and editor he has become one of the most respected
names in plant breeding, stimulating interest and endeavor in others.
Over the years he has received many accolades. Recognition by the
University of Edinburgh for his contributions in research and education
resulted in a well deserved Honorary Professorship in 1975, and simi-
larly he was elected to be a fellow of the Royal Society of Edinburgh in
1970. One international award which gave him considerable pleasure
was that of Distinguished Economic Botanist by the American Society
of Economic Botany in 1991, the only non-American then to be so
awarded. Norman Simmonds’s career, which has demonstrated clear
commitment to improving knowledge, has been an inspiration to us all
and we proudly dedicate this volume of Plant Breeding Reviews to him.

PUBLICATIONS OF NORMAN SIMMONDS

Books
Simmonds, N. W. 1959. Bananas. Longman, London.
Simmonds, N. W. 1962. The evolution of the bananas. Longman, London.
Simmonds, N. W. 1966. Bananas, 2nd edition. Longman, London.
Simmonds, N. W. 1976. The evolution of crop plants (ed.). Longman. London.
Simmonds, N. W. 1979. Principles of crop improvement. Longman, London.
Simmonds, N. W., and R. H. Stover. 1987. Bananas, 3rd edition. Longman, London.
Smartt, J., and N. W. Simmonds. 1995. Evolution of crop plants, 2nd edition. Longman,
London.
Simmonds, N. W., and J. Smartt. 1999. Principles of crop improvement, 2nd edition.
Blackwell Science, Oxford.

Papers
Bananas
Dodds, K. S., and N. W. Simmonds. 1946. Genetical and cytological studies of Musa VIII.
The formation of polyploid spores. J Genet. 47:233–241.
Simmonds, N. W. 1946. The relative yields of bananas and potatoes. Trop. Agr. (Trinidad)
23:226–228.
Simmonds, N. W., and K. S. Dodds. 1947. Persistence of a nucleolar remnant during
meiosis in a diploid banana. Ann. Bot. (London) 11:370–374.
Simmonds, N. W., and K. S. Dodds. 1948. Genetical and cytological studies of Musa IX.
The origin of an edible diploid and the significance of interspecific hybridisation in the
banana complex. J. Genet. 48:285–296.
Simmonds, N. W. 1948. Genetical and cytological studies of Musa X. Stomatal size and
plant vigour in relation to polyploidy. J. Genet. 49:57–68.
Simmonds, N. W., and K. S. Dodds. 1948. Sterility and parthenocarpy in diploid hybrids
of Musa. Heredity 2:101–117.
3499 P-01 9/26/2000 1:07 PM Page 6

6 W. SPOOR AND F. ENGLAND

Simmonds, N. W. 1948. The effects of ploidy upon the leaf of Musa. Ann. Bot. (London)
12:441–453.
Simmonds, N. W., and K. S. Dodds. 1949. Meiosis in seeded diploids of Musa. J. Genet.
49:221–251.
Simmonds, N. W. 1950. Polyploidy in bananas. Proc. VII Int. Bot. Congr. (Stockholm), 335.
Baker, R. E. D., and N. W. Simmonds. 1951. Bananas in East Africa I. Emp. J. Expl. Agr.
19:283–290.
Baker, R. E. D., and N. W. Simmonds. 1951. Banana research: Changes in outlook. Trop.
Agr. (Trinidad) 28:43–45.
Simmonds, N. W. 1952. Experiments on the pollination of seeded diploid bananas. J.
Genet. 51:32–40.
Simmonds, N. W. 1952. The strength of banana petioles in relation to ploidy. Ann. Bot.
(London) 16:341–347.
Simmonds, N. W. 1952. The germination of banana seeds. Trop. Agr. (Trinidad) 29:2–16.
Simmonds, N. W., and K. Shepherd. 1952. An Asian banana (Musa acuminata) in Pemba,
Zanzibar Protectorate. Nature (London) 169:507.
Simmonds, N. W., and R. E. D. Baker. 1952. Bananas in East Africa II. Emp. J. Expl. Agr.
20:66–76.
Simmonds, N. W. 1952. La banane à la Guadeloupe et à la Martinique. Fruits d’Outre Mer
7:67–69.
Simmonds, N. W. 1953. Segregations in some diploid bananas. J. Genet. 51:458–469.
Simmonds, N. W. 1953. The development of the banana fruit. J. Expt. Bot. 4:87–105.
Baker, R. E. D., and N. W. Simmonds. 1953. The genus Ensete in Africa. Kew Bul.
1953:405–416.
Simmonds, N. W. 1953. Classification of the bananas. II. T. Musa erecta U. Musa
angustigemma. Kew Bul. 1953:571–574.
Simmonds, N. W. 1953. Notes on the banana bunch. Trop. Agr. (Trinidad) 30:54–59.
Simmonds, N. W., and F. J. Simmonds. 1953. Experiments on the banana borer, Cos-
mopolites sordidus, in Trinidad. B.W.I. Trop. Agr. (Trinidad) 30:216–223.
Simmonds, N. W. 1954. Isolation in Musa section Eumusa and Rhodochlamys. Evolution
8:65–74.
Simmonds, N. W. 1954. Mutations in the Cavendish banana group. Trop. Agr. (Trinidad)
31:131–132.
Simmonds, N. W. 1954. Anthocyanins in bananas. Nature (London) 173:402.
Simmonds, N. W. 1954. Anthocyanins in bananas. Ann. Bot. 18:471–82.
Steward, F. C., and N. W. Simmonds. 1954. Growth promoting substances in the ovary and
immature fruit of the banana. Nature (London) 43:1083–1084.
Simmonds, N. W. 1954. A survey of the Cavendish group of bananas. Trop. Agr. (Trinidad)
31:126–130.
Simmonds, N. W. 1954. Varietal identification in the Cavendish group of bananas. J. Hort.
Sci. 29:81–88.
Simmonds, N. W. 1954. Notes on banana varieties in Hawaii. Pacific Sci. 8:226–229.
Simmonds, N. W. 1955. Wild bananas in Malaya. Malayan Nat. J. 10:1–8.
Simmonds, N. W., and K. Shepherd. 1955. The taxonomy and origins of the cultivated
bananas. J. Linn. Soc. Bot. 55:302–312.
Simmonds, N. W. 1956. Botanical results of the banana collecting expedition 1954–5. Kew
Bul. 1956:463–489.
Simmonds, N. W. 1956. A banana collecting expedition to South East Asia and the Pacific.
Trop. Agr. (Trinidad) 33:251–271.
Simmonds, N. W. 1958. Ensete cultivation in the southern Highland of Ethiopia: A review.
Trop. Agr. (Trinidad) 35:302–307.
3499 P-01 9/26/2000 1:07 PM Page 7

1. DEDICATION 7

Simmonds, N. W. 1959. Experiments on the germination of banana seeds. Trop. Agr.


(Trinidad) 36:259–273.
Simmonds, N. W. 1960. Megasporogenesis and female fertility in three edible triploid
bananas. J. Genet. 57:269–278.
Simmonds, N. W. 1960. Notes on banana taxonomy. Kew Bul. 1960:212.
Simmonds, N. W. 1960. Experiments on banana fruit development. Ann. Bot. (London)
24:212–222.
Simmonds, N. W. 1960. The growth of post-war West Indian banana trades. Trop. Agr.
(Trinidad) 37:279–283.
Simmonds, N. W. 1962. The classification and nomenclature of the bananas and potatoes:
Some implications. Proc. Linn. Soc. (London) 173:11–13.
Simmonds, N. W. 1985. Classification and breeding of the bananas. p. 62–73. In:
IRAZ/IDRC, Workshop on Banana Production in Eastern and Central Africa, Burundi.
IDRC, Ottawa, Canada.
Simmonds, N. W. 1985. Bananas, Musa cvs. p. 17–24. In: FAO Workshop on Disease Resis-
tance in Tropical Perennial Crops. FAO, Rome.
Simmonds, N. W. 1987. Classification and breeding of bananas. p. 69–73. In: G. J. Persley,
and E. A. DeLanghe, Banana and plantain breeding strategies, ACIAR/INIBAP, Canberra.
Simmonds, N. W., and S. T. C. Weatherup. 1990. Numerical taxonomy of the wild bananas.
New Phytol. 115:567–571.
Simmonds, N. W., and S. T. C. Weatherup. 1990. Numerical taxonomy of cultivated
bananas. Trop. Agr. (Trinidad) 67, 90–92.

Potatoes
Simmonds, N. W. 1963. Correlated seed and tuber dormancy in potatoes. Nature (London)
19:720–721.
Simmonds, N. W. 1963. Experiments on the germination of potato seeds. Eur. Potato J.
6:45–60, 69–76.
Simmonds, N. W. 1963. Abbreviations of potato names. Eur. Potato J. 6:186–190.
Simmonds, N. W. 1963. Studies of the tetraploid potatoes. I. J. Linn. Soc. Bot. 58:461–474.
Simmonds, N. W. 1964. Studies of the tetraploid potatoes. II. J. Linn. Soc. Bot. 59:43–56.
Simmonds, N. W. 1964. The genetics of seed and tuber dormancy in the cultivated pota-
toes. Heredity 19:489–504.
Simmonds, N. W. 1964. Observations on potato callus and adventitious shoot formation.
Am. Potato J. 41:129–136.
Simmonds, N. W. 1965. Some experimental techniques with potatoes. Eur. Potato J.
8:125–132.
Simmonds, N. W. 1965. Attempted graft-transmission of potato tuber dormancy. Eur.
Potato J. 8:197–199.
Simmonds, N. W. 1965. Seed size in the cultivated potatoes. Euphytica 14:143–152.
Simmonds, N. W. 1965. Chimeral potato mutants. J. Hered. 56:139–142.
Simmonds, N. W., and J. B. Harborne. 1965. Control of malvidin synthesis in the cultivated
potatoes. Heredity 20:315–318.
Simmonds, N. W. 1965. Somatic segregation of the spectacle pattern on potato tubers.
Heredity 20:277–288.
Simmonds, N. W. 1965. Mutant expression in diploid potatoes. Heredity 20:65–72.
Simmonds, N. W. 1966. Studies of the tetraploid potatoes III. J. Linn. Soc. Bot. 59:279–288.
Simmonds, N. W. 1966. Linkage to the S-locus in diploid potatoes. Heredity 21:473–479.
Simmonds, N. W., and P. E. Waggoner. 1966. Stomata and transpiration of droopy pota-
toes. Plant Physiol. 41:1268–1271.
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8 W. SPOOR AND F. ENGLAND

Simmonds, N. W., and J. F. Malcolmson. 1967. Resistance to late blight in Andigena pota-
toes. Eur. Potato J. 10:161–166.
Simmonds, N. W. 1968. Prolonged storage of potato seeds. Eur. Potato J. 11:150–156.
Simmonds, N. W. 1968. Change of leaf size in the evolution of the Tuberosum potatoes.
Euphytica 17:504–506.
Simmonds, N. W. 1969. Variegated mutant plastid chimeras of potatoes. Heredity 24:
303–306.
Simmonds, N. W. 1969. The genetics of spectacle in diploid potatoes. Heredity 24:487–490.
Simmonds, N. W. 1971. The potential of potatoes in the tropics. Trop. Agr. (Trinidad)
48:291–299.
Simmonds, N. W. 1973. A note on somatic segregation of the spectacle pattern in potatoes.
Heredity 31:405–407.
Simmonds, N. W., and R. J. Killick. 1974. Specific gravity of potato tubers as a character
showing small genotype environment interactions. Heredity 32:109–112.
Simmonds, N. W. 1974. Dry matter content of potatoes in relation to country of origin.
Potato Res. 17:178–186.
Simmonds, N. W. 1976. Neotuberosum and the genetic base in potato breeding. ARC Res.
Rev. 2:9–11.
Simmonds, N. W. 1977. Relations between specific gravity, dry matter content and starch
content in potatoes. Potato Res. 20:137–140.
Simmonds, N. W. 1980. Comparisons of the yields of four potato varieties in trials and in
agriculture. Expl. Agr. 16:393–398.
Simmonds, N. W., and R. L. Wastie. 1987. Assessment of horizontal resistance to late blight
of potatoes. Ann. Appl. Biol. 11:213–221.
Simmonds, N. W. 1998. A review of potato propagation by seed as distinct from clonal
propagation by tubers. Potato Res. 40:191–214.

Sugarcane
Simmonds, N. W. 1967. Potato breeding ideas applied to sugarcane. SCBNL 19:1–8.
Simmonds, N. W. 1967. Botanical nomenclature of sugarcane. SCBNL 9–11.
Simmonds, N. W. 1969. Radiation-induced non-flowering mutants. SCBNL 24:4–7.
Simmonds, N. W. 1971. More generations? SCBNL 26:4–6.
Simmonds, N. W., and R. J. Killick. 1972. More generations? SCBNL 29:4–6.
Simmonds, N. W. 1972. Profitability selection in relation to trials economy. SCBNL
29:20–23.
Simmonds, N. W., and D. I. T. Walker. 1972. Rate of turnover of cane varieties. SCBNL
30:8–10.
Simmonds, N. W. 1973. Optimal replanting time for sugarcane. Int. Sugar J. 75:107–108.
Simmonds, N. W. 1976. Towards a strategy for sugarcane smut control in the West Indies.
Int. Sugar J. 78:329–330.
Simmonds, N. W. 1976. A slide rule for sequential Brix sampling. SCBNL 38:701.
Simmonds, N. W. 1976. Two points about selection intensity. SCBNL 38:73–74.
Simmonds, N. W. 1976. Progress in sugarcane breeding. SCBNL 38:75–78.
Simmonds, N. W. 1979. The impact of plant breeding on sugarcane yields in Barbados.
Trop. Agr. (Trinidad) 56:289–300.
Simmonds, N. W. 1981. Comparisons of the performance of sugarcane varieties in trials
and in agriculture. Expl. Agr. 17:137–144.
Simmonds, N. W. 1983. Contingency and correlation: Applications to sugarcane breeding.
SCBNL 45:20–25.
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1. DEDICATION 9

Simmonds, N. W. 1983. Variety trials. SCBNL 45:26–29.


Simmonds, N. W., and D. I. T. Walker. 1984. Breeding, selection and trials. p. 124–135.
In: F. H Blackburn, Sugar cane. Longman, London.
Simmonds, N. W. 1984. A note on the strategy of breeding clonal crops. Heredity 53:
397–410.
Simmonds, N. W. 1984. Decentralised selection. Sugar Cane 6:8–10.
Simmonds, N. W., and D. I. T. Walker. 1986. An economic selection index for sugar cane
breeding. Euphytica 35:311–317.
Simmonds, N. W. 1986. Sugarcane, Saccharum. p. 115–124. In: FAO Workshop on Dis-
ease Resistance in Tropical Perennial Crops. FAO, Rome.
Simmonds, N. W. 1987. The leading features of cane breeding. Sugar Cane 1987
(Suppl.):8–11.
Simmonds, N. W., and D. A. Elston. 1988. Models of sugar cane smut disease and their
implications for testing variety resistance. J. Appl. Ecol. 23:319–329.
Simmonds, N. W. 1994. Some speculative calculations on the dispersal of sugar cane smut
disease. Sugar Cane 1994/1:2–5.
Simmonds, N. W. 1995. Reflections on sugar cane. Int. Sugar J. 97:24–26.

Other Plants
Simmonds, N. W. 1945. Meiosis in tropical Rhoeo discolor. Nature (London) 155:731.
Simmonds, N. W. 1945. Biological flora of the British Isles. Polygonum, P. persicaria, P.
lapathifolium, P. petecticale. J. Ecol. 33:117–143.
Simmonds, N. W., and K. S. Dodds. 1946. A cytological basis of sterility in Tripsacum
laxum. Ann. Bot. (London) 9:109–116.
Simmonds, N. W. 1946. Biological flora of the British Isles. Gentiana pneumonanthe. J.
Ecol., 33:295–307.
Simmonds, N. W. 1949. Notes on the biology of the Aracaea of Trinidad. J. Ecol.
38:277–291.
Simmonds, N. W. 1950. The Aracaea of Trinidad and Tobago, B.W.I. Kew Bul.
1950:391–406.
Baker, R. E. D., and N. W. Simmonds. 1953. Solanaceae. Flora Trin. Tob. 2(4):241–272.
Simmonds, N. W. 1954. Chromosome behaviours in some tropical plants. Heredity
8:139–146.
Simmonds, N. W., and E. Harrison. 1959. Genetics of reaction to pepper vein-banding
virus. Genetics 44:1281–1289.
Simmonds, N. W. 1960. Flower colour in Lochnera rosea. Heredity 14:253–261.
Simmonds, N. W. 1964. Nyctaginaceae—Batidaceae. Flora Trin. Tob. 2(7):439–480.
Simmonds, N. W. 1966. Plant and seed colours in Chenopodium pallidicaule. Heredity
21:316–317.
Simmonds, N. W. 1967. Hydrocharitaceae—Burmanniaceae. Flora Trin. Tob. 3(1):4–12.
Simmonds, N. W. 1967. Marantaceae. Flora Trin. Tob. 3(2):4–34.
Simmonds, N. W. 1971. The breeding system of Chenopodium quinoa. I. Male sterility.
Heredity 27:73–82.

Plant Breeding
Simmonds, N. W. 1961. Mating systems and the breeding of perennial crops. Advancem.
Sci. (London) 18:183–186.
Simmonds, N. W. 1962. Variability in crop plants, its use and conservation. Biol. Rev.
37:442–465.
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10 W. SPOOR AND F. ENGLAND

Simmonds, N. W. 1969. Genetical bases of plant breeding. J. Rubb. Res. Inst. Malaya
21:1–10.
Simmonds, N. W. 1973. Plant breeding. Phil. Trans. Royal Soc. Lond., B. 267:145–156.
Simmonds, N. W. 1974. Costs and benefits of an agricultural research institute. R&D Man-
agem. 5:23–28.
Simmonds, N. W. 1977. Breeding perennial crops. TAA Newslett. 17(4):27–29.
Simmonds, N. W. 1980. Polyploidy in plant breeding. SPAN 23:73–75.
Simmonds, N. W. 1981. Genotype (G), Environment (E) and GE components of crop yields.
Expl. Agr. 17:355–362.
Simmonds, N. W. 1983. Plant breeding: The state of the art. p. 5–25. In: T. Kosuge, C. P.
Meredith and A. Hollaender (eds.), Genetic engineering of plants. Plenum Press, New
York.
Simmonds, N. W. 1983. Strategy of disease resistance breeding. p. 637–654. In: J. P.
Gustafson (ed.), Gene manipulation and crop improvement, Columbia, MO (16th Stadler
Symp.).
Simmonds, N. W. 1985. A plant breeder’s perspective of durable resistance. FAO Plant.
Prot. Bul. 33:13–17.
Simmonds, N. W. 1985. Perspectives on the evolutionary history of tree crops. p. 3–12.
In: M. G. R. Cannell and J. E. Jackson (eds.), Attributes of trees as crop plants, ITE.
Simmonds, N. W. 1985. Two stage selection strategy in plant breeding. Heredity
55:393–399.
Simmonds, N. W. 1986. Strategies for disease resistance breeding in tropical perennial
crops. p. 3–16. In: FAO Workshop on Disease Resistance in Tropical Perennial Crops.
FAO, Rome.
Simmonds, N. W. 1986. The strategy of rubber breeding. Int. Rubber Conf., Kuala Lumpur,
1985 3:115–126.
Simmonds, N. W. 1986. Theoretical aspects of synthetic/polycross populations of rubber
seedlings. J. Nat. Rubb. Res. 1:1–15.
Simmonds, N. W. 1986. Oil palm, Elaeis guineensis. p. 79–84. In: Workshop on Disease
Resistance in Tropical Perennial Crops. FAO, Rome.
Kennedy, A. J., G. Lockwood, G. Mossu, N. W. Simmonds, and G. Y. Tan. 1987. Cocoa
breeding, past, present and future. Cocoa Growers Bul. 38:5–22.
Simmonds, N. W. and S. Rajaram. (eds.) 1988. Breeding strategies for resistance to the rusts
of wheat. CIMMYT, Mexico.
Simmonds, N. W. 1988. Synthesis: The strategy of rust resistance breeding. p. 119–136.
In: N. W. Simmonds, and S. Rajaram (eds.), Breeding strategies for resistance to the rusts
of wheat, CIMMYT, Mexico.
Simmonds, N. W. 1988. Environmental features of plant breeding. p. 3–10. In: AAB, Envi-
ronmental aspects of applied biology, AAB, Warwick.
Simmonds, N. W. 1989. Rubber breeding. p. 85–124. In: C. C. Webster and W. J. Baulkwill
(eds.), Rubber, Longman, London.
Simmonds, N. W. 1989. Economic aspects of plant breeding, with special reference to eco-
nomic index selection. Res. and Dev. in Agr. 6:53–62.
Patterson, H. D., and N. W. Simmonds. 1989. Tables to calculate means in a doubly trun-
cated bivariate normal population. Euphytica 42:241–249.
Simmonds, N. W. 1989. A statistical vade-mecum for the itinerant agriculture/researcher.
Trop. Agr, (Trinidad) 67:9–15.
Simmonds, N. W. 1989. How frequent are superior genotypes in plant breeding popula-
tions? Biol. Rev. 64: 341–365.
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1. DEDICATION 11

Simmonds, N. W. 1990. The social context of plant breeding. Plant Breed. Abstr.
60:337–341.
Simmonds, N. W. 1990. Breeding horizontal resistance to South Americal leaf blight of rub-
ber. J. Nat. Rubb. Res. 5:102–113.
Simmonds, N. W. 1990. Plant breeding, seed technology and beyond. Proc. ISTA Conf.,
Edinburgh 1989, p. 81–88.
Simmonds, N. W. 1991. Genetics of horizontal resistance to diseases of crops. Biol. Rev.
Camb. Phil. Soc. 66:189–241.
Simmonds, N. W. 1991. Selection for local adaptation in a plant breeding programme.
Theor. Appl. Genet. 82:363–367.
Simmonds, N. W. 1991. Uniformity and yield in hybrid crop cultivars. Trop. Agr.
(Trinidad) 68: 198–199.
Simmonds, N. W., and M. Talbot. 1992. Analysis of on-farm rice yield data from India.
Expl. Agr. 28:325–329.
Simmonds, N. W. 1992. Cocoa breeding in Sabah. Trop. Agr. Assoc. Newslett. 12(2):9.
Spoor, W., and N. W. Simmonds. 1993. Pot trials as an adjunct to cereal breeding and eval-
uation of genetic resources. Food Crops Res. 35:205–213.
Simmonds, N. W. 1993. Introgression and incorporation. Strategies for the use of crop
genetic resources. Biol. Rev. Camb. Phil. Soc., 68:539–562.
Simmonds, N. W. 1993. Tropical plant breeding: success or failure or a bit of each? Trop.
Agr. Assoc. Newslett. 13:3–5.
Simmonds, N. W. 1994. The breeding of perennial crops. Int. Workshop on Conservation
Characterization and Utilization of Cocoa Genetic Resources into the 21st Century.
CRU/UWI, Trinidad, 1992, p. 156–162.
Simmonds, N. W. 1994. Horizontal resistance to cocoa diseases. Cocoa Growers’ Bul.
47:42–52.
Simmonds, N. W. 1994. Yield and sugar content in sugar beet. Int. Sugar J. 96:414–416.
Simmonds, N. W. 1995. Plant breeding. Biol. Sci. Rev., Jan. 1995:32–34.
Simmonds, N. W. 1996. Family selection in plant breeding. Euphytica 90:201–208.
Simmonds, N. W. 1996. Bias in the estimation of horizontal resistance to airborne fungi
in plant breeding. Bot. J. Scotl. 48:275–284.
Simmonds, N. W. 1996. Profits, projects and plant breeding. Trop. Agr. Assoc. Newslett.
16(1):31–32.
Simmonds, N. W. 1997. Gough and Chittenden, two rubber pioneers. Planter, Kuala
Lumpur.
Simmonds, N. W. 1997. Tropical crops and their improvement. p. 257–293. In: C. C. Web-
ster and P. N. Wilson. Agriculture in the tropics, 3rd ed. Blackwell, Oxford.

Phenolics
Forsyth, W. G. C., and N. W. Simmonds. 1954. A survey of the anthocyanins of some trop-
ical plants. Proc. Roy. Soc. B. 142:549–564.
Simmonds, N. W., and R. Stevens. 1956. Occurrence of the methylene-dioxy bridge in the
phenolic components of plants. Nature (London) 178:752–753.
Forsyth, W. G. C., and N. W. Simmonds. 1957. Anthocyanidins of Lochnera rosea. Nature
(London) 180:247.
Harborne, J. B., and N. W. Simmonds. 1964. The natural distribution of the phenolic agly-
cones. Ch. 3. In: J. B. Harborne (ed.), Biochemistry of phenolic compounds. Academic
Press, London.
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