Development of High-Vigour Oat Varieties in Australia

PATRICK MARTIN GUERIN

Copyright © Patrick Martin Guerin 2005 This book is copyright. Apart from any fair dealing for the purpose of study, research, criticism, review, or as otherwise permitted under the Copyright Act, no part may be reproduced by any process without written permission. Inquiries should be made to the publisher. National Library of Australia Cataloguing-in-Publication Development of High-Vigour Oat Varieties in Australia Patrick Martin Guerin

ABOUT THIS BOOK AND

THE AUTHOR’S OAT VARIETIES

In reviewing “Development of High-Vigour Oat Varieties in Australia”, I considered that it was a very full and well documented account of oat breeding and testing in New South Wales in the latter half of the twentieth century. In correspondence with a research agronomist, who also runs a property in Northern NSW, she stated “---- Blackbutt (bred by the Author, P. M. Guerin) has stayed a very strong variety for a very long time. The quality of the breeding is reflected in its longevity as a preferred variety---”. I think that everyone interested in the oat crop whether researcher, advisory officer, producer or plant breeder should read, study and learn from this important book.

Professor Haydn Lloyd-Davies Former Professor of Pastoral Science in the School of Wool and Pastoral Science,

University of New SouthWales Past President of the Australian Society for Animal Production (NSW Branch)

& Author of “Animal Production” This book discusses the evolution of oats as a crop in Australia, emphasising its versatility and value to farmers and to the agricultural sector. In particular, it sets out the importance of dual-purpose oat varieties in Australian agriculture, which are of significant value to this day. As the prime lamb industry in Australia continues to expand, and in a sense is about to “take-off”, dual-purpose oat varieties will play a significant part in the expansion of this industry. This is a book that had to be written. I believe the book illustrates clearly what a visionary the Author was in developing the oat varieties and lines he did.

Norm Markham

Former District Agronomist (25 years), New South Wales Department of Primary Industries

(formerly NSW Department of Agricutlure), & Independent Agricultural Consultant currently based in West Wyalong, NSW, Australia

“Reading Patrick Guerin’s book reminded me of times, as a research agronomist on the Southern Tablelands. There I conducted numerous experiments on the effects of winter grazing of cereals and other crops, on vegetative yield, animal production and subsequent grain yields. This work was published in the Australian Journal of Experimental Agriculture, its predecessor and elsewhere. Guerin’s variety Blackbutt was always the stand-out crop for maximum combined forage and grain production, particularly in the severe winter environment of the Tablelands” “Congratulations, Paddy, on an important publication”

Paul Dann Former Research Agronomist, NSW Agriculture

Having witnessed oats being successfully grazed on a family property at Lyndhurst in Central West NSW, and having already been convinced of the health benefits of oats as a human breakfast staple, it was a great pleasure for me to study each absoprbing, and very readable chapter. Without closing the door on gene technology as a way forward, his highlighting of the challenges, Patrick is convincing in his support of the Isolection Medelian plant breeding system.

Bob Fozzard, Sydney, Australia

Member of the Australian Institue of Agricultural Science and Technology “I started growing Carbeen at our property in mid 1980’s. We like to start sowing in February. We often have a dry March – April and on these occasions, other varieties will run to head, whereas Carbeen doesn’t and it recovers well. With the quantity of leaf material and its prostrate growth, our sheep can keep grazing for a longer time compared to erect growing oat varieties. If Carbeen is eaten out early, its recovery is good. When we grow Carbeen for grain, then a sowing rate of 30 kg/ha will yield up to 2.9 t/ha, and that is under continuous grazing”. “Carbeen is a variety well suited for growing in the Tamworth region of New South Wales”.

John McQueen Farmer

“Colindale”, Loombenah, New South Wales, Australia

“I grew Blackbutt in the very early 1980s, switching over from Cooba which my father had grown for years. My early memories of harvesting Cooba were with a comb front harvester, sometimes with croplifters, which was nightmarish! We noticed straight away that with Blackbutt that it had more tillers and the plants were very hardy, able to withstand the often dry autumns that we endure on our farming area. Blackbutt was fairly prostrate in early stages but with late autumn rain, it emerged into a massive bulk of feed in the winter months”. “We continued to grow Blackbutt until we heard of another one, of related breeding, called Carbeen. A friend had been growing it for a few years. We gave it a go and have been growing it ever since. Some of the characteristics of Carbeen have been its ability to withstand lodging in all but the very lushest of seasons, its ability to put out lots of tillers when sown early, and when these become erect later, they have a very nutritious broad leaf that our stock thrive on. “I am so happy with the production levels from Carbeen that I have not even tried the new varieties or even winter wheats as I really don’t think they could be any better than what I have seen with Carbeen”

Paul McCulloch Farmer

“Danibe”, Tamworth New South Wales, Australia

ABOUT THE AUTHOR

The Author was born in Chile in 1928, of an Irish father and an English mother. He graduated with a Bachelor of Agricultural Science from National University of Ireland, Dublin, in 1952. From 1950 he became an amateur beekeeper and from 1952 to 1955, he was a Milk Costings Officer for the Irish Department of Agriculture. He then became Lecturer in Chemistry and other agricultural subjects at Warrenstown Agricultural College, Co. Meath, Ireland. He was then an Abstractor for Herbage and Field Crop Abstracts for the Commonwealth Agricultural Bureau (CAB) at Maidenhead, England from 1955-1956. In 1956 he was appointed Plant Breeder for NSW and permanently moved to Australia. In Australia,

the Author was an oat and linseed breeder from 1956 to 1964, stationed at Glen Innes with NSW Agriculture. Using both established and original techniques, the Author bred Australia’s most frost resistant and productive winter grazing cereal variety, Blackbutt oats. The Author developed the Isolection system of plant breeding, a technique for producing High-vigour oat varieties. Using this system, he made a High-vigour cross in 1957, from which he bred and selected P4315, as well as Blackbutt oats, as well as numerous other oat verities. Blackbutt and P4315 both broke world records for yield in 1973. From 1972 until 1985, he engaged in farming near Temora, NSW, giving his 7 children experience of a farming lifestyle. He produced wheat and forage crops and managed sheep, cattle, and high quality pigs for bacon. He then retired to the Sydney region of NSW to return to study and writing. The subjects he studied included plant breeding, genetic engineering, languages, philosophy, physical anthropology, prehistory, celtic and religious studies, theology, and history. Patrick currently lives in Lithgow, NSW.

CONTENTS

List of Tables 1

List of Figures 3

Preface 6

Acknowledgments 8

Chapter 1 Introduction 9

Chapter 2 Australian Oat Varieties And A Germplasm Inventory For Breeding

37

Chapter 3 The New Isolection Plant Breeding System 62

Chapter 4 Breeding Oats For Irrigation In Australia 104

Chapter 5 The Influence Of Environment On Oat Grain Quality

115

Chapter 6 Plant Breeding Methods And Technologies For Increasing Oat Crop Yields

124

Glossary 137

Appendix A Australian Oat Statistics 150

Appendix B Plots From A Heavy Grazing Trial 154

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LIST OF TABLES

Table 1.1 Chemical composition of oats 10

Table 1.2 Comparative feed grain values of oats, barley, wheat and maize 11

Table 1.3 Percentages of nutritive values in oats, barley, wheat and maize 16

Table 1.4 Species of Avena genus, the 3 karyotypes and their genomes 18

Table 1.5 World population densities 22

Table 1.6 Food production and population growth 23

Table 1.7 Changes in total grain yields and reduction in total crop growing area 24

Table 1.8 Annual rate of change (%) of increase in production of farm products 25

Table 1.9 World land utilisation 27

Table 1.10 Oat yields, growing days, population density and agricultural policy 28

Table 1.11 Stocking capacity of oats compared with other pastures 29

Table 2.1 Effect of grazing oats twice, versus no grazing, on grain yield of various cultivars

39

Table 2.2 Summer rainfall germplasm 47

Table 2.3 Uniform rainfall germplasm 49

Table 2.4 Winter rainfall cultivars 50

Table 2.5 Crosses combining rust resistance with agronomic value 51

Table 2.6 Segregation in landraces for juvenile growth habit, Glen Innes 1958 51

Table 2.7 Resistances for various environments 52

Table 2.8 Origin and description of genotypes developed from the High-vigour cross, 28 X 23

53

Table 3.1 Rapid method of breeding oats for large biomass yields 64

Table 3.2 Morphology and pathology of parents of the High-vigour cross 65

Table 3.3 The effect of grazing intensity on a range of cereal genotypes sown late March in a cool, moist, summer rainfall climate: F6 generation trial of High-vigour lines (1962)

69

Table 3.4 The effect of grazing intensity on a range of cereal genotypes sown in early March in a cool, moist summer rainfall climate: F7 generation trial of High-vigour bulk oats (1963) at Glen Innes

70

Table 3.5 Second testing of High-vigour bulk oats in north-west NSW, contrasting cooler elevated site (Tamworth) with warmer plains site (Narrabri): F5 generation trial (1961)

76

Table 3.6 A comparison of southern and northern NSW bred cultivars under intensive grazing and hay recovery: F10 generation testing of High-vigour lines at Richmond (1966)

77

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Table 3.7 Heavy (4 P cuts) and lenient (2 P cuts) grazing and grain recovery (pG) at Cowra: F10 generation testing of the High-vigour lines (1966)

83

Table 3.8 Lenient grazing and grain trial: F17 generation testing of High-vigour bulk oats

84

Table 3.9 Effect of multiple grazing cuts on grain and pasture yields on a range of oat cultivars

85

Table 3.10 Grain and pasture yields from 1955 competing crop trials 86

Table 3.11 Effect of a single grazing, grain recovery, total yield and grain protein (%) on a range of oat cultivars

89

Table 3.12 Effect of two grazing cuts on grain recovery and pasture yields on a range of oat cultivars

90

Table 3.13 Dry matter of pasture and grain recovery trial, Gunning, NSW (1999) 91

Table 3.14 Grain yields from competing a cereal crop trial conducted in New England

91

Table 3.15 A continuous grazing (P) and grain recovery (pG) trial in Central NSW; F34 generation testing of High-vigour varities (1990) at Blayney

92

Table 3.16 Effect of two grazing cuts and grain recovery (Site 1) and grain only (Site 2) on a range of oat cultivars

94

Table 4.1 NSW cereal crop yields under dryland and irrigation (t/ha) 105

Table 4.2 Comparisons of early and late maturing cultivars under Irrigation in the Riverina for a 10 year period for grain only (G), and grain recovery (pG) (1963-1973)

109

Table 4.3 Comparisons of grain only (G) yields and grain recovery (pG) in the Dryland Riverina for a 10 year period (1963-1973)

110

Table 4.4 Year 24 of testing Blackbutt oats under irrigation versus dryland (1985): F29 generation trial

111

Table 5.1 Grain quality, as groat (gt %), of Australian cultivars and accession lines 117

Table 5.2 Oat grain quality, as weight in grams of 1000 seeds (groat + hull) and as groat %, of cultivars at four sites in Northern NSW

118

Table 5.3 Grain quality of mainly High-vigour oats (Cross C and Cross A, 28 x 23), F7 generation testing

119

Table 6.1 Isolection-bred versus conventionally-bred oat varieties (Richmond, NSW)

130

Table 6.2 Isolection-bred versus conventionally-bred oat varieties (Southern Highlands NSW)

131

Table 6.3 Yield ratios of Isolection-bred to conventionally-bred oat variety (Cooba) across climatic zones from statistically analysed trials over a 29 year period

132

Table 6.4 Comparing features of GM crops with conventional crops 134

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LIST OF FIGURES

Figure 1.1 Statistical yields of the major cereals grown in NSW 32

Figure 2.1 Straw strength of various lines and cultivars. Fulghum (F) showing lodging; weak strawed Belar; strong strawed Garry, VRBke.F (W4598); strong strawed Fulmark

42

Figure 2.2 Climatic regions of Australia 45

Figure 2.3 A transect of NSW showing the Northern, Central and Southern regions in NSW approximating summer rainfall, uniform rainfall and winter rainfall zones, respectively

46

Figure 2.4 BLACKBUTT variety with medium panicle shape and light brown grains: the longest grazing season cultivar of Australian winter cereals. Photograph is from Australian Oat Varieties by R.W. Fitzsimmons et al. CSIRO (1983)

47

Figure 2.5 COOLABAH is an early grazing and grain variety with medium panicle shape and cream coloured grains. It is too frost susceptible for the summer rainfall germplasm list. Photo is from Australian Oat Varieties by R.W. Fitzsimmons et al. CSIRO (1983)

49

Figure 2.6 ORIENT is an erect early midseason variety for grain only, with medium to open panicle and dark brown grains. It is too frost susceptible for the summer rainfall germplasm list. Photograph from Australian Oat Varieties by R.W. Fitzsimmons et al. CSIRO (1983)

50

Figure 2.7 ALGERIAN variety with open panicle and mid-brown grains from Australian Oat Varieties by R.W. Fitzsimmons et al. CSIRO (1983). Table 6.5 shows that both Algerian and Fulghum were segregating for juvenile habit of growth in the 1958 F2 summer rust nursery, confirming Coffman’s claim that the related varieties of Red Rustproof and Kanota in the USA could not be fixed

51

Figure 2.8 COOBA is a mid-season grazing and grain variety with open panicle and mid brown grains from Australian Oat Varieties by R.W. Fitzsimmons et al. CSIRO (1983). Cooba is inferior to Blackbutt and Carbeen for grazing and frost resistance in the summer rainfall zone

52

Figure 2.9 CARBEEN variety with condensed panicle shape and medium brown grains. A mid-season variety with prostrate early habit of growth, the most adaptable to the 3 rainfall zones. Photograph from Australian Oat Varieties by R.W. Fitzsimmons et al. CSIRO (1983)

53

Figure 2.10 FULGHUM spiklets and florets from Oat Identification and Classification by T.R. Stanton (1955) US Department of Agriculture Technical Bulletin No. 1100. Fulghum is a semi-winter type and appears to be of hybrid origin, with many traits intermediate between the northern common oats, A. sativa, and the southern red oats, A. byzantina, as judged by observers in the US

54

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Figure 2.11 FLORET SEPARATION distinguishes the 2 types of cultivated oats which are Avena sativa, separating by distal fracture, and A.byzantina, usually separating by basal fracture. Photograph from F.A. Coffman, Inheritance of Morphological Characters in Avena, Technical Bulletin No. 1308, Agricultural Research Service, United States Department of Agriculture

54

Figure 2.12 MORPHOLOGICAL CHARACTERISTICS OF THE OAT PLANT, showing panicle and spikelet, main rachis and panicle branches; rachilla and basal hairs of mature grain; spikelet showing pedicel, glumes, rachilla, primary grain and secondary grain and awn on the primary grain; culm nodes and nodal hairs and leaf margins and leaf sheaths, both hairy and glabrous. From Anonymous (1962)

55

Figure 2.13 The Author assessing mature oat crop stands. Avon x VRFB; Garry x VRBke (2056) and Fulghum (see summer rainfall germplasm inventory)

56

Figure 2.14 Mature oat crop stands. The Author with a tall strong strawed line; Avon (see the uniform rainfall germplasm inventory) and taller W4477

57

Figure 3.1 Results of heavy grazing by sheep; The pasture cut technique using manual shears at Glen Innes, NSW

74

Figure 3.2 Tall strong straw of Fulghum (F) x Garry (Ga) (F.Ga or W4595), typical of the F.Ga cross; Close up of the panicles of F.Ga, the female parent of the High-vigour cross

75

Figure 3.3 Non-stress growing environment. A plastic covered frame for establishing rust infected plants, transplanted from the subtropical station at Grafton, and designed to spread rust and determine rust resistant plants; Inspection of individual oat plants

78

Figure 3.4 Non-stress growing environment Fulghum x Garry (female parent of the High-vigour cross showing) showing its strong straw; Wide spacing of individual oat plants

79

Figure 3.5 Crossing of a rust resistant line, of oat, 0600 and VRAF (W4890) 80

Figure 3.6 A typical Western Australian bred cultivar, Swan, showing poor dry matter recovery under a 5 grazing cut regime at Temora, New South Wales, 1969, in comparison with moderately frost-hardy Cooba and very frost-hardy P4315

82

Figure 3.7 The Author shows greater damage to Algerian from a combination of frost and grazing pressure than that to Klein 69B the Argentine oat, which showed excellent frost resistance and grazing recovery almost equal to Blackbutt; The Author shows poorer performance of Algerian compared to High-vigour line P4314. Further images of the grazed plots at Hawkesbury Agricultural College trials in Richmond NSW in 1966, are presented in Appendix B

95

Figure 3.8 A comparison of the five selections from the High-vigour cross for total biomass yield (P + pH) with conventionally bred cultivars at Hawkesbury Agricultural College, Richmond, NSW (1966)

96

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Figure 3.9 A comparison of a standard cultivar, Algerian, with five selections from the High-vigour cross, with 5 separate pasture cuts at Hawkesbury Agricultural College, Richmond, NSW (1966). The extent of the grazing is shown in individual plots within the trial presented in Appendix B

97

Figure 3.10 The Author at Temora Agricultural Research Station taking notes near seed increase blocks. Selecting hardy, productive and rust resistant dual-purpose oats by wide spacing of plants in the Author’s Isolection breeding system, produced P4315, Blackbutt both from the same High-vigour cross; Mugga, also bred by the Author, is the hardiest of the oats tested in Glen Innes NSW, equivalent hardiness to winter wheat. Mugga was selected from VRBop x Belar

99

Figure 4.1 A diagrammatic comparison of eight cultivars under irrigation (grain only) and cool dryland (grain recovery) in t/ha (Colleambally, NSW in 1985)

112

Figure 5.1 Grain shape and sizes of the parents of the High-vigour cross 120

Figure 5.2 Grain shape and sizes of the High-vigour varieties and lines (Blackbutt, Carbeen and P4315) alongside conventionally bred cultivars

121

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PREFACE

“An economic oat breeding program will only succeed in proportion to the breeder's skill and knowledge in being able to consider the project as a whole, rather than as separate entities” J.G. Carroll (1951)1 This book describes the importance of the oat crop to sustainable farming and the pivotal role that oat breeders have in this. It describes the outcomes from the author’s contribution to the oat breeding program run by the New South Wales (NSW) Department of Agriculture from 1957 to 1974, including oat line and variety breeding and testing. The book covers the development of high yielding, dual-purpose grazing and grain oat varieties, and the methods used to breed and test these varieties, including trial results up to the present day. Some of this work was published in 1961, 1965, 1966, 1992 and in 2003. Chapter One introduce the role that oats play as an important role in human and livestock nutrition, and as such, an understanding of the genetics of oats is significant in world agriculture and economics. Oats provides grain for humans and livestock, a grazing or forage crop for livestock, as well as the ability to provide combined grazing and grain production. While the significance of the oat grain in benefiting human health has received considerable attention in the past decade, relatively little attention has been given to this important attribute of combined grazing, grain production and total crop value in the research and extension literature. This reflects a lack of awareness of the full potential of the oat crop. Based on the recent findings of FAO studies, the world supply of agricultural produce is meeting the demands of the current world population. The total world production of cereals increased at an annual rate of 1.45% over the period of 1981-1990, while total meat production increased at an annual rate of 2.87%. These trends suggest that increased cereal crop yields have allowed for an increase in the area available for pasture and hence livestock production. Improving the total quantity and quality of world pasture production is therefore becoming increasingly important for meeting the corresponding increases in global food demands. The oat crop has a sigficant role to play in this increase in pasture production. Chapter Two describes how oat breeding has led to the development of oat varieties for the 3 main climatic regions of Australia. These three climatic regions or zones also exist in the state of New South Wales (NSW). These three regions are as follows: The sub-tropical climate zone, also referred to as the summer rainfall zone, and also occurs on the coastal areas of Southern Queensland and Northern NSW (including Grafton, where a crown rust nursery is located). The uniform rainfall climate zone which covers the inland area of NSW from as north as Dubbo to Temora in southern NSW. The winter rainfall climate occurs south of Temora and includes the Australian states of Victoria, Tasmania, South Australia and Western Australia. An inventory of oat cultivars and their pedigrees is presented in relation to the climatic regions in which oats are grown in Australia. The inventory tables list the name or accessional line of the oats, their pedigrees and breeder. A description of the Austalian oat ideotype is also proposed. Chapter Three described the results of 34 years of oat breeding and testing of dual-purpose varieties (for grazing and grain recovery) by the NSW Department of Agriculture are summarised in this chapter. A High-vigour cross (HvII 57-75) is identified which led to the 1 From a NSW Department of Agriculture Internal Report. James Carroll was a plant breeder dedicated to oats, potatoes and gladioli.

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release of Blackbutt (an F4 directed bulk type) in 1975, and Carbeen (an F6 plant progeny of the normal pedigree system) in 1981. This High-vigour cross also produced a number of high yielding F4 directed bulk types and F2 plant progenies bulked in the F3 as a result of their relatively high phenotypic uniformity. The highest yielding F3 bulk was numbered P4315, which although classed as an early oat, out-yielded all other varieties, including Blackbutt, for total biomass, following early sowings, and over a wide range of soils and climates and a great many seasons. The success of these oats was due to the Isolection plant breeding system pioneered by the Author at Glen Innes from 1957 to 1964. Other F4 directed bulks were P4314 (high-yielding both as a winter oat and a spring oat at Glen Innes) and P4318, both of which had large grains and, together with Blackbutt and P4315, were significantly superior over 5 grazing cuts (including the mid-winter cut) to Coolabah and all other advanced lines submitted by plant breeders from Temora NSW, that were using conventional breeding methods during the same period. Selection of lines at the F2 generation has been demonstrated as a simple way of forecasting wider adaptability of early generation material. Chapter Four describes why Glen Innes, on the New England Tablelands in NSW, has proven to be the best centre for breeding oats for the heavy soils of the Riverina at Leeton, southwestern NSW. Plant selections made on the black self-mulching soils of the Glen Innes Research Station of northern NSW have resulted in the varieties Acacia, Bundy, and Mugga; all now replaced by Blackbutt. Both areas require resistance or tolerance to stem rust, water logging, red-legged earth mites, BYDV, lodging, shattering and second growth. Although frost damage is less of a problem in the irrigation areas than on the northern tablelands of NSW, the frost resistant bulks from the cross F.Ga x VRAF.VRSF demonstrated good tolerance to water logging on heavy soils. Blackbutt also excelled as both a dual-purpose and a grain only variety and is recommended for both northern and southern irrigation areas. Chapter Five describes the important influence of the oat growing environment on oat grain quality. Oat grain quality (grain weights per 1000 seeds and groat percentages) was found to be an effective measure of the environmental stress imposed on an oat variety at a particular geographical and climatic centre. The results of various oat trials conducted across NSW show that the environment has an effect on the maturation and filling of the oat grain. The results compiled by the Author suggest that northern NSW (i.e. the summer rainfall zone) could be further sub-divided into 5 climatic regions, from east to west, for the purpose of recommending oat varieties. Glen Innes, at an elevation of 1,128m and latitude 29° 42” S. on the New England Tablelands, proved to be the ideal climate for developing high groat percentage and large grain size. A sixth climatic zone, located in Leeton, NSW (uniform rainfall zone), at elevation at 152m and latitude 34° 33” S., was the second most favourable centre, but required irrigation for full grain development. Chapter Six discusses plant breeding methods and technologies and their potential for increasing oat crop yields and oat crop improvement. It specifically introduces the importance of hybrid vigour and a non-stress environment for higher percentage heritability selection and therefore providing a more productive conventional plant breeding method for the improvement of crops. This chapter draws together the results from trials presented in Chapters Three and Four to show the superiority of the Isolection method over the conventional oat breeding method for development of high yielding, multi-purpose oat varieties. GM technology and crops derived from cloning, a process devoid of hybrid vigour, are compared with proven plant breeding methods.

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ACKNOWLEDGMENTS

The Author is indebted to his colleagues from the NSW Department of Agriculture; the late Dr F. Mengersen, oat breeder, for generously sharing his knowledge; Farm Manager Messrs. Eric Powell, the Farm Manager at Glen Innes, who made the Author welcome and at ease during the first years of his settling in Australia. He fondly remembers fellow agronomists, Martin Bellert of Queensland and the late Milton Walker, both of whom were very supportive in this work; Ern Tindale and Jack Loveridge for providing the fertile seed-beds so necessary for genetic gains in the oat crop; Mr. I. Cole for pure seed production; Mr. Bill Uppsdell and Mr. Jack Stapleton for maintaining the link of pedigree records; the oat germplasm and uniform techniques from pioneer oat breeders, the late Dr S. L. Macindoe and Mr. J. C. Carroll, previously of Glen Innes Station; District Agronomists, Agronomists-in-training and all personnel, past and present, involved in the NSW Agricutlure oat breeding and testing program. The Author also acknowledges other Australian agronomists, completely unknown to him, who carried out complex grazing experiments with his Blackbutt, P4315 and Carbeen (selected by Mr. Glenn Roberts of Temora Research Station), all cultivars from the High-vigour Cross, and had them published in scientific journals. Some of these people included Mr. Muldoon, who found Blackbutt to be the best variety, physiologically, for total grazing and grain of any winter cereal, under irrigation at Trangie; Mr. McLeod and Mr. Ramsey, who found Blackbutt and Carbeen were the highest producers of total grazing and grain after 4 grazings at Bendigo, Victoria; Mr. Craig and Mr. Potter found that Carbeen was the only variety at Kybybolite, South Australia, which increased its grain yield after 2 grazings. The Author also acknowledges the efforts of Mr. G. Hennessy, who obtained a world grain yield record for P4315 oats (an early maturing sister-line to Blackbutt) of 20 tonnes per ha, following two grazings, at Tamworth Research Station in 1973. The Author is grateful to his son, Dr. Turlough Guerin, for co-authoring and organizing two articles for the 1992 International Oat Conference at Adelaide, as well as for criticism, co-ordinating reviews, editing and assistance in putting this book together. The Author is also grateful to Mr. Roger Fitzsimmons, who retired as Assistant Principal Agronomist of Cereals in the NSW Department of Agriculture, for collating trial results, reading and correcting the manuscript and for valuable advice from his long experience. The critical reviews provided by several agricultural scientists are kindly acknowledged including those provided by Professor Haydn Lloyd-Davies, Professor Peter Ruckenbaur, Professor Frank Crofts, Paul Dann, Norm Markham, Wayne Vertigan, Bob Fozzard and Andy Roberts. Dr. W. Jim Althom is also acknowledged for his review of a final draft of the manuscript.

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CHAPTER ONE

INTRODUCTION

Oats play an important role in human and livestock nutrition, and as such an understanding of the genetics of oats is significant in world agriculture and economics. Oats provides grain for humans and livestock, a grazing or forage crop for livestock, as well as the ability to provide combined grazing and grain production. While the significance of the oat grain in benefiting human health has received considerable attention in the past decade, relatively little attention has been given to this important attribute of combined grazing, grain production and total crop value in the research and extension literature. This reflects a lack of awareness of the full potential of the oat crop. Based on the recent findings of FAO studies, the world supply of agricultural produce is meeting the demands of the current world population. The total world production of cereals increased at an annual rate of 1.45% over the period of 1981-1990, while total meat production increased at an annual rate of 2.87%. These trends suggest that increased cereal crop yields have allowed for an increase in the area available for pasture and hence livestock production. Improving the total quantity and quality of world pasture production is therefore becoming increasingly important for meeting the corresponding increases in global food demands. The oat crop has a sigficant role to play in this increase in pasture production. INTRODUCTION The importance of the oat crop in human and animal nutrition has been established only relatively recently. Studies demonstrating the cholesterol lowering effect of oat bran and other oat products in laboratory animals and humans have been known since the early 1990s. This has been attributed to the high soluble fibre (β-glucan) content of oats and oaten bran, confirming traditional beliefs in the value of oats, relative to all other cereals. Knowledge of β-glucan, oil and protein contents of oat varieties, in various germplasm collections, will enable breeders to add value to all agronomically useful varieties for optimum human and animal nutrition. The nutritional and health qualities of the oat crop are of considerable importance in establishing the context for this book and these attributes are discussed in this chapter. A brief introduction to oat genetics and the origin of the oat crop is also provided as well as a discussion of the broader agricultural and economic significance of this crop globally.

OAT GRAIN QUALITY AND HEALTH Overview Research has identified oats as the health grain for humans and animals (McDonald et al. 1992). There are active components in oats which lower blood lipids, regulate blood glucose and protect against tumour development in the colon.

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The cholesterol-lowering benefits of oats have been attributed to ß-glucan in the oat fibre fraction. Oat bran and oatmeal supplementation studies show a more favourable effect on blood glucose and insulin responses than other cereal grains like wheat and maize. Oat soluble fibre should delay the onset of fatigue and enhance athletic performance. Besides addressing major diseases of wealthy nations, like coronary heart disease, cancer and diabetes, oats could provide benefits for blood pressure and weight reduction. Oats also contain a high proportion of monounsaturated fat, antioxidants such as tocotrienols, and an amino acid composition rich in arginine relative to lysine. Antioxidants have been linked to reduced risk of cancer, heart disease and degenerative changes in the eye as well as to increased immune function (Bunce et al. 1990; Diplock 1991). Coeliac disease in human individuals, sensitive to gluten and unable to eat wheat, barley or rye (all high in gluten) can, usually, safely eat oats which contains only trace quantities of the gluten protein (Welsh, 1995). Table 1.1 shows the marked superiority of oat bran over rolled oats both in protein and in dietary fibre, contrary to popular uninformed opinion which formerly regarded oat bran as less valuable. Table 1.2 compares feed grain values of the 3 winter cereals and maize. It should be noted that high fibre in oats goes hand-in-hand with a high oil content. The oil composition in oats is high in linoleic acid and low in linolenic acid. Oat hulls are very effective in inhibiting the development of dental caries in animals at dietary levels of 3 to 25%. Phenolic compounds in the hulls may involve antioxidant or antimicrobial activity (Madsen, 1981). Table 1.1 Chemical composition of oatsa. Nutrients Rolled Oats Oat Bran

Energy 1600 kJ 1030kJ Protein 10.5 17.3 Fat, total 8.0 7.0 Fat, saturated 1.5 1.2 Carbohydrates 61 50.3 Sugars 0.0 2.6 Dietary fibre 10.0 15.9 Sodium < 5.0 < 5.0 Thiamine (B1) - 1.2 mg

a From BiLo (2004). Values given grams per 100 grams.

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Table 1.2 Comparative feed grain values of oats, barley, wheat and maizea. Nutrients Oats Barley Wheat Maize

Protein (%N x 6.25) 10.5 11.0 12.5 10.4

Oil (%) 5 2 2 4

Crude fibre (%) 10 5 2 2

GEb (MJ/kg DMd) 19.6 18.5 18.7 18.9

MEc (ruminants) 12.0 12.9 13.5 13.8 a From Welsh (1986); bGE. = Gross energy; cME. = Metabolisable energy in ruminants; d = dry matter. Genes encoding the oat kernel storage proteins, avenins and globulins, have been isolated and characterised. Oat globulins, which make up 50-80% of the kernel protein, resemble legume globulins in amino acid composition thus explaining the nutritionally balanced amino acid content of oat proteins. The protein of oats is unique among temperate cereals because of this high content of globulin, which closely resembles a major seed legume protein, glycinin (Peterson and Brinegar, 1986). Therefore oat and legume proteins may have similar hypocholesterolaemic properties. These properties or cholesterol reducing effects are higher in oat bran than in rolled oats. Thus, Ripsin et al. (1992) took 3g of soluble fibre to be equivalent to 42g of oat bran or 84g of oatmeal. This superior effect of oat bran suggests that there is a role for both the gum and the protein since both these components are higher in the bran. The major component of oat gum is β-glucan.

Oats have long been the breakfast cereal of the Celtic people of Ireland, Scotland, Wales and the cooler, wetter northern counties of England, the North American, Scandinavian, North European and Slavonic peoples. More recently, the crop has spread to West Africa and is likely to become universally important with its unique value for the health of humans and animals, relative to other grains. Composition of oat grain Oats have a slightly sweet, slightly sour taste, which does not require the addition of sugar or honey. Oats can also be blended with a variety of other health-giving foods. To understand the nutritional significance of oats, it is necessary to look at the various constituents of oats. Protein. Oats have a higher concentration of well-balanced protein than other cereals and therefore a greater potential value to provide a substantial proportion of protein requirements than other cereals. Among the essential amino acids that make up protein quality, cereals are generally limiting in lysine. Oat protein is higher in lysine than that of other cereals. Lipids. Lipids are a concentrated source of energy, being higher in energy value than carbohydrate. The lipid concentration of oats is higher than that of other cereals. The lipid composition of oats is favourable because of the high proportion of unsaturated fatty acids. Oats are high in linoleic acid, an essential fatty acid for human nutrition. Linoleic acid is used in the synthesis of prostaglandins that are found in all tissues and regulates smooth muscles.

Minerals. Oats are a good source of manganese (Mn), magnesium (Mg), iron (Fe), calcium (Ca), zinc (Zn) and copper (Cu). The major proportion (58%) of the phosphorus in the oat

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kernel occurs as phytic acid. Phytic acid may bind minerals, making them unavailable in nutrients. Rolled oats, however, did not decrease the absorption of Fe any more than did milk, which contains no phytic acid.

Vitamins. Oats contain little or no vitamin A, C or D but it does contain small yet significant quantities of thiamine, folic acid, biotin and pantothenic acid.

Starch. The starch concentration of oats on a whole grain basis is lower than that of rye, barley or wheat, reflecting the relatively thick hull of oats.

Soluble sugars. Total free sugar concentration of oats is low, relative to barley, wheat and rye, but similar to maize (Table 1.1). Fibre. Dietary fibre is defined as plant polysaccharides and lignin, substances resistant to human digestive enzymes. Starch is the only plant polysaccharide that is digestible by humans. Therefore, dietary fibre includes all non-starchy polysaccharides (NSP) plus lignin. NSP include cellulose, hemicellulose and lignin (all water insoluble), whereas other fibre components are classified as soluble. The solubility factor is important for understanding the importance of oats for human nutrition (Shinnick et al. 1988). The significance of oat fibre and human health Whole oats, before processing, have 20-37% fibre. After processing, the oatmeal has about 12% fibre, while the oat bran, the coarse milling fraction, contains about 18% dietary fibre. The dietary fibre of oats is a mixture of soluble and insoluble fractions and the soluble fraction is high relative to other cereals due to the high concentration of ß-glucans in oats. The irregular configuration of these polymers makes them partially water soluble and functionally different from cellulose in the human digestive system. Only barley exceeds oats in concentration of ß-glucans, but a higher proportion of oat ß-glucan is soluble. There is a wide range of ß-glucan concentration among diploid oat species but narrower ranges among tetraploid and hexaploid oat species. The highest values are found in the hexaploid or cultivated species of oats.

Oat ß-glucans are especially abundant in the bran fraction that contains the outer layer of the caryopsis and thick cell walls of the sub-aleurone region (Henry 1987).

Cholesterol lowering properties The cholesterol lowering properties (or hypocholesterolemic) effects of oats have been proven in both animal and human studies. These are discussed in the following sections.

Animal studies. As early as 1963, rolled oats were found to decrease serum cholesterol level of rats fed a semi purified diet with 10g per kg cholesterol and 2g per kg cholic acid (De Groot et al. 1963). The hypocholesterolemic effect of oats was greater than that of other grains tested. In other experiments, the soluble gum fraction of oat bran was the most effective in lowering serum and liver cholesterol.

Human studies. Studies with experimental animals have been confirmed in human feeding trials. When hypercholesterolemic male subjects were fed diets containing 140g of rolled oats daily, their cholesterol levels were significantly lowered 11% in 3 weeks, and levels rose again when the oat-containing diets were discontinued (De Groot et al. 1963). In another

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study, an 8% drop in the serum cholesterol levels of 17 hypercholesterolemic individuals was observed after 4 weeks of a diet containing <35% of energy from fat (Turnbull and Leeds 1987). Subsequently, the inclusion of 150g of rolled oats resulted in a further reduction of serum cholesterol levels by 5%, whereas wheat supplementation produced no further reduction in cholesterol. Further, in a study involving 236 subjects with normal cholesterol levels, total cholesterol decreased 6.6% with a fat-modified diet alone and 8.3% with a fat-modified diet plus 56g of oatmeal daily (Van Horn et al. 1988). This study showed that oatmeal or oat bran ingestion may enhance serum cholesterol reduction induced by dietary fat modification both in individuals with high cholesterol and healthy levels.

Hypotheses to explain cholesterol lowering by oats. There are several theories as to how oats lead to lowering of cholesterol. The value of oat soluble fibre has been explained by dietary cholesterol absorption, bile acid reabsorption, production of lipoproteins in the liver and removal of lipoproteins in peripheral tissues (Anderson and Gustafson 1988). The presence of oat products in the small intestine increases the viscosity of the intestinal contents, leading to a slower rate of dietary cholesterol absorption, thus reducing its availability and increasing faecal excretion (Lund et al. 1989). In the same study, oat bran diets increased the faecal excretion of bile acids in human subjects. Lower amounts of bile acids returned to the liver may divert liver cholesterol from lipoproteins to bile acids. However, not all soluble fibre sources that lower plasma cholesterol increase bile acid excretion as does oat bran fibre, and the magnitude of the increase from those that do, is small. A further hypothesis is that oat fibre-induced short chain fatty acids inhibit cholesterol synthesis in peripheral tissues. This would result in a surplus of low-density lipoprotein (LDL) receptors, to increase the rate of LDL clearance. No single mechanism will explain the effects on cholesterol concentrations of oat bran soluble fibre (Marshall and Sorrells 1992). Further, the effects of a high-fat meal (50g fat) on healthy individuals have been shown to be alleviated by oats. Endothelial dysfunction induced by acute fat ingestion is prevented by concomitant ingestion of oats or vitamin E, but not wheat. As a result, Katz et al. (2001) concluded that oats are better than wheat for cardiovascular health. The glycemic effects of oats Soluble dietary fibre in the diet slows the increase in blood glucose that normally follows a meal and is important in the treatment of Type II diabetes. Ingestion of oatmeal or oat bran decreased the glycemic index (blood glucose response relative to that induced by white bread) and insulin response in healthy and diabetic individuals (Heaton et al. 1988). In no insulin-requiring diabetics, oat bran and oat gum at levels of 8g of soluble fibre slowed the rate of increase in blood glucose (Braaten et al. 1988). At 40 min, blood glucose concentrations were significantly lower for both treatments, compared to a control (cream of wheat), and peak glucose concentrations were delayed 30 to 40 min by both treatments. Similar results were obtained with healthy individuals. Oat gum was as effective as guar gum, but oat gum was tolerated better by most subjects. Oat bran with 15% ß-glucan lowered blood glucose by 40%.

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Prevention of colon cancer The antioxidant properties of the tocotrienols and phenolic compounds in oats should inhibit colon tumour development. In countries where the diet is associated with a low prevalence of coronary heart disease, prevalence of colon cancer is also low (McDonald et al. 1992). Relevance of the oat health factors to agriculture The oat health factors are also of significance to agriculture more broadly and these are discussed in the following sections. Relevance to plant breeding. This book stresses the importance of dual-purpose oat breeding, that is oats used for grazing and grain production (described in Chapters 2-6), for which the most successful centre for NSW was at Glen Innes. There is no longer a need to grow oats only for ease of milling. Heavily grazed oats may or may not (depending on good summer rain) recover grain with a higher proportion of oat bran, now the most sought after health component of the oat crop, both for humans and animals. Oat bran is of significance because it contains a high proportion of ß-glucan. Composition analysis shows 4.3-4.6% ß-glucan in rolled oats and 7.3-8.9% in oat bran (Welch 1995). The lignin (insoluble fibre) component of total fibre in oat bran was 20% while that of oatmeal was 27%, indicating total lignin contents of 3.8% and 3.3% respectively (Shinnick et al. 1988). Within a given oat cultivar, increasing nitrogen fertility levels increased groat (i.e. seed minus the husk) protein and groat ß-glucan (Welch et al. 1991). There are also genotypic differences in groat ß-glucan and this can be selected for without undesirable correlated responses (Peterson 1991). In oats, the β-glucan is found within the bran (or the outer portion) of the groat. Oats, belonging to the Aveneae family, have higher levels of all the essential amino acids, namely cysteine, histidine, isoleucine, leucine, lysine, methionine, phenylalanine, threonine, tryptophan, tyrosine and valine, than the Triticeae family, which includes wheat, rye and barley. Lysine tends to be the first limiting essential amino acid in cereals and lysine deficiency is exacerbated if the protein content of a cereal is increased by nitrogen fertilizer application. The decline in protein quality at higher protein levels is less pronounced in oats than in other cereals. This is associated with the relative contribution of the various protein solubility fractions to oat total protein. The prolamine fraction, which is low in essential amino acids, is chiefly increased in other cereals, as protein is raised by genetic or environmental changes. This accounts for the marked decline in protein quality observed in wheat, barley or maize as protein is increased (Mossé 1968). Globulin was found to be the major protein fraction in oats. Since the globulin fraction has a similar amino acid composition to the total protein, the relative increase in globulins with increasing total protein accounts for the relative stability of the amino acid composition of oats over a wide range of protein contents (Peterson 1976). Thus, globulins account for 70-80% of the total oat groat protein, with glutelins accounting for less than 5-10% of the total oat groat proteins. Relevance to milk production. Friesian-cross cows were fed ad libitum on grain-based diets, comparing barley, wheat and oats, all rolled. Although the oat-based diet had the lowest content of dietary metabolisable energy (MJ/kg), it produced the greatest yield of milk and milk fat. Replacing barley with oats changed the fatty acid composition of the milk, by significantly reducing the saturated fatty acids and significantly increasing the content of stearic and oleic acids (Moran 1986). The oat-based diet in dairy cattle therefore increases the appeal of milk and milk products to the health-conscious consumer.

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Relevance to animal feeding and production. Oats may be successfully fed to pigs, cattle, sheep, poultry and horses. By crushing oats and feeding them to pigs, the Author, farming at Temora from 1972 to 1985, obtained a price premium from Gilbertsons of Melbourne (meat processors), due to the reduction in back-fat, which was high when he was feeding crushed wheat to his pigs (both lots receiving a similar protein supplement). Oats in cow rations produced milk fat with an increased proportion of polyunsaturated fatty acids (Martin and Thomas 1988). The high fibre content of whole oats limits their use to ruminants and horses, both capable of digesting fibre. Oats is the only cereal that does not need to be processed (as by rolling) prior to feeding to horses. For highly productive animals, naked oats have a superior nutrient content to wheat and barley and for this reason they justify a price premium on the basis of least-cost formulation (Valentine 1990). Oats are the preferred grain for horses: other cereals like wheat pack too tightly in the gut, whereas oats remain in a loose mass that can be easily digested by the horse. The two facets of the oat crop are (1) herbage that becomes richer in protein the more frequently it is grazed and (2) grain, which is the safest of the cereals for ruminants and farm work horses (Whittemore and Elsley, 1976). In varying proportions, any nutritive ratio such as 10:1 (starch:protein equivalent) for dry and resting stock or 4:1 (starch:protein equivalent) for breeders, lactators or growers, can be easily attained. Even oaten straw, which is nutritionally superior to that of wheat and barley, can be used for maintenance (Welch, 1986). Oat straw is softer and more acceptable to stock than other cereal straws and has a higher metabolisable energy content than other cereals in terms of available energy. Oats pasture is superior to native grass-subterranean clover pasture for ewe live-weight gain (Dann et. al. 1974). For finishing prime lambs, daily live-weight gains of 400 g and stocking rates of 60 lambs per ha were reported on oat pastures (Archer and Swain 1977). A Canadian study investigated growth performance, carcase and meat quality of pigs fed oat-based diets containing four levels of β-glucans. No evidence of detrimental effect of ß-glucans in oat-based diets, particularly at levels below 4%, was detected, lending support for the inclusion of oats in finisher diets (Fortin et al. 2003). The high quality of oat grain and especially the biological value of its protein content and higher calcium content are both important for humans and young growing animals including pigs (Table 1.3). For pigs, however, barley has the ideal fibre content for fattening quickly. In summer rainfall regions, however, there is no shortage of damaged wheat. This can be mixed with oats in a proportion to give about 5% crude fibre. Oats should be crushed prior to preparing a balanced food ration, which should include meat offals, or some other source of protein, for pigs.

Oats can also enhance resistance of animals to bacterial and parasitic infections. In one study, the oral or parenteral oat β-glucan treatment enhanced the resistance to Staphylococcus aureus and Eimeria vermiformis infection in mice. The β-glucan, extracted from oats, significantly enhanced phagocytic activity (Yun et al. 2003).

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Table 1.3 Percentages of nutritive values in oats, barley, wheat and maize.

Valuea Oats Barley Wheat Maize

Dry matter Crude fibre Gross energyb Digestible energy Digestible protein Lysine Methionine + Cystine Calcium Protein qualityc

86 10.0 16.9 11.4 7.7 0.37

0.40 0.07 73

86 4.8 16.0 12.7 7.7 0.32

0.27 0.04 69

86 3.0 16.2 14.0 8.3 0.28

0.38 0.03 63

86 2.0 16.2 14.5 7.3 0.26

0.25 0.02 58

a Values based on all being 86% dry matter; b expressed as megajoules (MJ) per kilogram of feed; c biological value is highest in oats due to favourable amino-acid ratios (Whittemore and Elsley 1976). In another livestock nutrition study, Flinn and Foot (1992) found oat grains samples ranged from 7-12% protein. Oats with low protein were shown to inhibit microbial activity in the rumen of grazing sheep and needed either green oat pasture or a protein supplement. All of the varietal samples determined by Craig and Potter (1983) were 12% protein or over in a South Australian study study assessing the effects of grazing on various oat varieties. Carbeen, a prostrate growing variety, tested 14.2% protein. Such oat grain would be ideal for drought feeding, when no ‘green feed’ is available. ORIGIN AND GENETICS OF OATS The oat genome All species of oats have originated in the northern hemisphere. The cultivation of oats is not very old. Neither the Egyptians nor the early Europeans grew oats. De Candolle (1883) ascribed a European origin to our cultivated oats, leaning on historical and philological facts. The European group of Avena sativa were typical of north-western Europe. A large Mediterranean group, sharply isolated from A. sativa, were A. sterilis and A. byzantina. These three species belong to the hexaploid oats (2n = 42), and can be easily be crossed together or with A. chinensis (2n = 42), the low yielding large naked oat from the Chinese centre of origin. These are our most important oats for breeding, testing and extension. “2n” represents the number of chromosomes in the sex cells (gametes) after fertilization. The number of chromosomes before fertilization is represented by n = 21. These chromosomes, however, each consist of 3 basal groups of chromosomes, each of which has 7 chromosomes. By dividing 7 into 42 we obtain 6, hence the term hexaploid. This represents the genome, the complete complement of genetic material in a cell of this species. Here the genome is written as AACCDD.

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The weedy species of A. fatua, or black oats, also belongs to this group. North-western Europe, including Wales, was also a centre for A. strigose and A. brevis, half weedy, part cultivated diploid species (2n = 14), neither of which can be hybridised with A. sativa or A. byzantina. These wild diploids were noted for resistance to smut, Ustilago avenae, crown rust, Puccinia coronata, as well as mildew, Erysiphe graminis. Vavilov found many varieties of A. sativa in Mongolia and northern China as well as in Georgia and Armenia, together with A. fatua and A. ludoviciana (Vavilov 1920-1940). These latter two were widely distributed all over south-western Asia. Vavilov found China to be the centre for the large and naked-grain oats, A. chinensis, genetically related to the European oats with chromosome numbers (2n = 42) and easily hybridising with each other. They were first brought from China to Europe in the 5th century AD (Breitschneider 1881). The Author calls this A. chinensis to distinguish it from the small naked oat, A. nuda, which is a diploid (2n = 14), like A. strigose and A. brevis. Most of the diploid species cannot cross with one another, the exceptions being clauda x eriantha, wiestii x hirtula, wiestii x strigose, and hirtula x strigose as listed in Table 1.4. Within the tetraploids, only the following crosses are possible: barbata x vaviloviana, barbata x abyssinica, and vaviloviana x abyssinica. This shows that possession of the same genomes, or sets of chromosomes, does not guarantee interfertility within the diploid or tetraploid species. Ethiopia, or the highlands of Abyssinia, is the centre of origin for A. abyssinica. Centre of origin is a better term than Vavilov’s centre of type formation, which may have been influenced by Darwin’s term, natural selection. In Table 1.4, the large naked oat created for China, A. chinensis, is on the same line as the small naked oat, A. nuda. This is to show that in the middle column for the tetraploids, a naked oat tetraploid has yet to be found. Many investigators, as reviewed by Legget and Thomas (1995), thought that the cultivated hexaploid oat had been derived by a simple trichotomy from a common progenitor. This was found to be improbable. If all 3 groups came from a single basic species, the polyploid species (the hexaploids) would have to be autopolyploids but they are not. Autopolyploids are derived by the doubling of the constituent genomes, as by the conversion of AA, the single genome, to AAAA. There is only one oat species like this, A. macrostachya, whose genomic constitution is unclear (Legget and Thomas 1995). This is the only outbreeding and perennial species of Avena and the only one that is autotetraploid. All other oat species are allopolyploids. One fact is certain: the cultivated hexaploid oats did not evolve from any of the known diploid or tetraploid species, because the donor of the DD genome is unknown. The same is true for the hexaploid wheat genome, which also has an unknown donor. The origin of the third or D genome of the hexaploid (2n = 42) species in cultivated oats varieties is completely unknown. This makes Rajhathy and Thomas’ (1974) theory of oat evolution purely speculative. The discovery of A. canariensis and the magna-murphyi complex in the tetraploid group (2n = 28) of oats is said to realize Vavilov’s law of homologous variation. This is said to be a structural analogy but this does not explain anything. The missing D-genome has never been found in the diploid oat species, which have AA or CC genomes, or in the tetraploid species which have AABB or AACC genomes. Our cultivated hexaploid oats are designated by the AACCDD genome complex. Therefore, on evidence, hexaploid oats cannot have originated from diploids or tetraploids, certainly not by natural crossing, or ‘fusion of distinct genomes,” as postulated by Rajhathy and Thomas (1974).

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Table 1.4 Species of Avena genus, the 3 karyotypes and their genomesa. Diploid = 7 Genome Tetraploid n= 14 Genome Hexaploid n = 21 Genome

clauda CC barbata AABB fatua AACCDD eriantha CC vaviloviana AABB sterilis: ventricosa CC abyssinica AABB ludoviciana AACCDD prostrata CC maroccano AACC maxima AACCDD damascena AA murphyi AACC macrocarpa AACCDD longiglumis AA sativa: hiemis AACCDD canariensis AA orientalis AACCDD wiestii AA diffusa AACCDD hirtula AA bizantina: strigose AA hiemis AACCDD brevis AA verna AACCDD nuda AA chinensis AACCDD

a From Guerin (2003). Note that there is a large diagram on p.154 of the Author’s self-published book referred to here, showing all the crosses that are possible within the Avena genus.

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The explanation given is that the donor is either extinct or has evolved into a different species. This depends entirely on the value of the hypothesis itself, that differentiation is a function of time. The possibility that the genome donor may never have existed is not even stated, let alone the alternative which that possibility would imply: separate origin of species in the various Vavilovian centres. This applies both to oats and wheat, Triticum aestivum L., which has a genome complex of AABBDD, in which the donor of the B-genome is unknown. Much ingenious effort and thinking have gone into this work, but we have not yet exploited a fraction of the cultivated oat gene pool. The significance of multigenic traits The history of the science of genetics has been a stormy one. The first of the great hybridisers was Joseph Kölreuter, 1733-1806 (see glossary). He described over 500 experiments, including Nicotiana rustica x N. paniculate, which gave a very vigorous hybrid, which was sterile when self-fertilised. It was Gregor Mendel (1822-1884), the father of genetics, who explained the continuous variation in height in Kölreuter’s second generation (F2) tobacco plants after crossing a dwarf with a tall parent. This was the green light or impetus for multigenic plant breeding for traits requiring quantitative or cumulative effects, as for high yielding oats from the Author’s Isolection system. Between 1900 (when Mendel’s paper was discovered) and 1910, most geneticists could see Mendel’s work as showing only discontinuous variation, looking only at his pea crosses. Mendel, however, had also discovered continuous variation, when he crossed white-flowered and purple-red-flowered beans. This gave an intermediate flower colour (pink) in the F1 progeny and a continuous spread from white to purple-red in the second generation. Geneticists then began to see that alleles (pairs of a gene) had small but cumulative effects with semi-dominance rather than complete dominance, which were behaving in a Mendelian fashion. This gave rise to the multiple-gene hypothesis. This is now one of the most important principles of genetics (Gardner and Snustad, 1984). This principle has been greatly strengthened by the use of statistical methods by R.A. Fisher in England. Fisher laid the foundation for the analysis of variance and the beginnings of experimental design and success in comparing oat variety yields in biometrically designed trials in Australia (Fisher 1925). These trials proved to other plant breeders that yield differences were or were not significant. The economically significant groups of oats There are various classifications for oats. These include those based on grain morphology. One can look at the grains after threshing or harvesting and see if the rachilla or stalk remains with the primary grain (Avena sativa) or with the secondary grain (Avena byzantina). Although the varieties Blackbutt and Carbeen derive from the same cross, the latter’s grain articulation is typically A. byzantina, while that of Blackbutt is a 50-50 mixture suggesting its own hybrid origin. Similarly Swan has the hybrid morphology of its sister-line, West. Swan and West belong to the specialised grain oats and therefore another mark of identity is required. The photograph of floret separation in Figure 2.11 of Chapter Two shows this. The early habit of growth is, however, the best indicator of economic significance. The habit of growth has a significant impact on the economic significance of oats and this is further described under the latter sections in this book that address dual-purpose oat varieties. These groups are prostrate and erect growing varieties and these are further sub-divided into intermediate, semi-erect, erect and very erect (described below). The most reliable mark of identity is whether the juvenile stage has a prostrate (Blackbutt and Carbeen), intermediate (Cooba), semi-erect (Coolabah), erect (Avon, Cassia, Stout and Swan) or a very erect (Moore

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and West) habit of growth. The prostrate varieties bury their growing point, tiller profusely, resist frost and grazing damage and are therefore dual-purpose varieties, suitable for both for both grazing and grain production. These two groups of oats, based on growth habit, have been compared in South Australia (Craig and Potter, 1983). This trial was evenly grazed by sheep, which also provided fertiliser and an even grazing. Comparing 0, 1 and 2 grazings, the erect varieties yielded more grain after one grazing than after 0 or 2. The most prostrate variety, Carbeen, was the only variety to yield more grain after 2 grazings than after 0 or 1 grazing in this trial. In this trial, plants were grazed by 100 sheep for 3 days to a uniform height of 2.5 cm above ground level. The prostrate variety, Carbeen, significantly outyielded all other varieties in grain recovery. The variety Carbeen, and the details of this trial, are further described in Chapter Three. The erect varieties from Western Australia have larger grains than the prostrate varieties from the Glen Innes breeders, and are usually accepted for milling for this reason, and the fact that they are grown in a drier finishing season, which does not discolour the grain. The Glen Innes bred varieties are smaller grained but are higher in groat percentage than varieties bred at Temora, Southern NSW, South Australia and Western Australia. Avena strigosa, cultivar Saia, has very small grains which give it a high volumetric weight. Saia is crown rust resistant and sown in Southern Queensland and Northern NSW coastal areas for cattle grazing. The grains possess up to 20% protein but belong to the diploid species of Avena that cannot be crossed (or only with great difficulties) with the normal cultivated hexaploid species of oats. GLOBAL AND ECONOMIC ASPECTS OF THE OAT CROP Overview Economic factors, and to some extent political factors, determine the motivation to grow a particular crop or pasture. These factors encompass global agricultural land potential, world population and comparative crop and pasture yields. Some of the data for this study has been taken from the Food and Agriculture Organisation (FAO) of the United Nations annual reports from 1948 to 1992. This data has been assembled and critically evaluated by Sassone (1994) and further elaborated by the Author in the remainder of this chapter. Oats, mainly Avena sativa and A. byzantina, have an important role in world pasture production. Considerable research and developments have been conducted on this crop in Australia and overseas. The application of research findings in agriculture has contributed to overcoming world food shortages (Sassone, 1994). Such has been the impact of improved practices in agriculture that countries in Asia, for example, are now demanding more milk products and meat in the diet as compared with traditional foods, in particular, rice. United Nations Yearbooks show that even with population increases of about 20%, the number of telephones, refrigerators and other amenities in third world countries have approximately doubled. Average real incomes have more than doubled (UNICEF, 1993). This demand for milk and meat products now increases the need for more efficient means of their production, including improved pastures and grain production. The role of dual-purpose grain types, oats grown for both grazing and grain production, to assist these developing countries to meet their demands in the Southern Hemisphere, including Australia, has been identified as being important (Guerin, 1961; Guerin and Guerin, 1992). Oats is not a coarse grain only, or a source of carbohydrate only such as wheat and rice (Whittemore and Elsley, 1976). Oats, however, possesses other characteristics which make it unique as a food source

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for both humans and livestock and these have been described earlier in this chapter. Due to favourable amino-acid ratios, oats have a higher biological value than barley, wheat or maize. Along with the cholesterol lowering attributes of the grains, oats can be considered as “the health crop”. Global trends in population, food supplies and diets Population growth or rate of increases is defined as the birth rate minus the death rate. As of 1990, this value for the world was 1.7%. Africa has the highest rate of increase in the world at 3.0% (Table 1.5). Asia and Latin America also have the second and third fastest rates of increase. The first world continents, Europe, North America and the former Soviet Union are increasing at less than 1%. These latter regions have fertility rates below 2.2 children per female, which represents the replacement or zero population rate. Sassone (1994) predicts that world populations may begin to decrease by the year 2050. Based on the FAO data, it is evident that world food production has increased, regardless of world population growth. Over the period studied by Sassone (1994), food production, especially rice and meat, quadrupled, while world population has little more than doubled. It is apparent that farmers in the developing world have adopted many of the new technologies and developments in agricultural science, which have dramatically improved crop yields. Intense effort by agricultural extension practitioners in developing countries have improved the rates of adoption of appropriate technologies for both grain and pasture production in these countries. These adoptions have included improved understanding of the need for fertilisers, pesticides, herbicides, soil tillage practices, and the growing of improved crop and pasture varieties. In India for example, on average, the population consumes the 2,200 calories recommended by the Food and Nutrition Board (Sassone, 1994). Developing countries in the Far East increased grain production by 12% while in Africa, grain production increased by 47% (Sassone, 1994). Meat production consistently increased from 1981 to 199 (Table 1.6). Total cereal tonnages, including wheat and rice, declined in volume over the same period. An inference that can be made from this data is that the area of land under pasture is likely to be increasing. World grain prices fell after 1981, while stocks of grain rapidly increased (Sassoon 1994). Supply controls were applied in most countries in the form of acreage reduction measures. During 1981-85, difficulties stemmed from depressed agricultural exports, high interest rates, and supply surpluses. In Australia this effect was particularly marked where meat production increased 22% from 1984-91. In the developed nations, farmers reduced grain production after 1984 because of massive grain surpluses. Farmers of the Near East and of Africa, on the other hand, increased grain production by over 40% between 1984 and 1991. The expansion of land areas used for rice and meat production has broader implications for integrating more balanced diets into the households of developing countries. Furthermore, the increased area of land devoted to pastures reflects the potential of increased crop rotations and ley farming and, therefore, general soil improvement.

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Table 1.5 World population densitiesa.

Location Population (P x 106) 1980

Population (P x 106) 1990

Population Increase (%)

Land Area (km2 x 103)

Population Densityb (P/km2)

World 4,448 5,292 1.7 136,255 39

Africa 477 642 3.0 30,305 22

Asia 2,583 3,171 1.9 27,582 115

Latin America 363 448 2.1 20,535 22

North America 252 278 0.8 21,962 13

Europe 484 498 0.2 4,933 101

Oceania 22.8 26.5 1.5 8,536 3

Former USSR 266 289 0.8 22,402 13 a Sassoon (1994); bP = Individual persons.

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Table 1.6 Food production and population growtha.

World Tonnes (x 106) Year

1948-52 1960 1970 1980 1990

Total cereals NAb NA 1215 1566 1952

Wheat 155 222 318 446 597

Rice 111 158 316 399 518

Total meat production 40 60 107 133 175

Population (x106) 2516 3020 3698 4448 5292

a Sassoon (1994) from Food and Agriculture Organization of the UN annual reports; b NA = not available.

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Table 1.7 Changes in total grain yields and reduction in total crop growing areaa.

Cropping Years Cropping Area (ha x 106)

Total Grain Production (x106)

Yield (t/ha)

1975/76 707.7 1236.8 1.75

1980/81 721.8 1427.2 1.98

1985/86 715.0 1645.6 2.30

1989/90 693.3 1665.8 2.40

1992/93 687.7 1758.5 2.56 a From World Grain Situation and Outlook (USDA, 1993).

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Table 1.8 Annual rate of change (%) of increase in production of farm productsa.

World Tonnes (x 106) Produce

1981 1986 1990 1991 Change (%)

Total cereals 1646 1854 1971 1887 1.45

Wheat 454 535 601 553 1.97

Paddy rice 412 471 522 518 2.26

Total meat 138 157 176 179 2.87 a Sassoon (1994).

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International increases in cereal grain yields World cereal grain yields have increased marginally in the period 1979 to 1994. The greatest increase was observed with the wheat crop. The increases in barley and oat yields were lower over the same period. While the total tonnage of wheat and barley have increased over this time, world oat yields “appear” to have decreased slightly. However, the accuracy of Australian oat yield statistics do not reflect actual yields because the oat crop is typically grazed throughout the growing season prior to harvest and considerably less stringent agronomic management is applied to this crop than to other cereals, in particular wheat and barley (Simmons, 1987). World grain production multiplied by 2.6 from 1950 to 1984 at the same time that the world human population less than doubled. The price of grain decreased over the same period and removed farmer’s incentives to grow more. In the 1950s and 1960s, a bushel of grain was worth the equivalent in dollar value of a barrel of crude oil. In the 1970s and 1980s, the price of grain in real terms was approximately 20% of the 1950 price, allowing for inflation, or 10% of the price of a barrel of crude oil. Farmers in the developed world therefore reduced grain production after 1984. Total world grain yields increased substantially over the period of 1976 to 1993 from 1.75 to 2.56 t/ha (Table 1.7). Over the same period, the total area of land used for grain production decreased from 7.08 x 108 ha to 6.88 x 108 ha. However, there was a temporary increase during this period to 7.22 x 108 and 7.15 x108 during the years 1981/82 and 1985/86, respectively. As a result of this increased productivity in grain production, approximately 30x106 ha have been made available for other agricultural activities including the growing of improved pastures. This increased availability of land for pasture production has increased the potential for dual-purpose grazing cereals, including oats. There is evidence that this has occurred from the increase in total meat production worldwide (Tables 1.5 and 1.8). This total increase in meat production, however, does not include increases due to the increased total number of lot-fed livestock and other intensive livestock industries. World-wide, approximately 22% of the land area has potential to be used for pasture. This does not include the 11% of arable land or the 30% estimated to be utilised in forestry. India utilises 92% of its agricultural potential without using the 18% of its total area that is devoted to forestry. The arable area of Australia of 48 million ha includes 17 million ha of crops and 31 million ha of sown pasture and grasses. Forestry includes 41 million ha of native forest, 1 million of plantation forestry and 36.5 million ha of protected wilderness areas, national parks and conservation areas (Table 1.9). The greatest part of Australia’s agriculturally potential land area of 419 million ha is used for grazing. In no other country or continent has livestock production dominated agriculture as in Australia. Hence Australia has played a leading role in the development of pasture improvement and development of dual-purpose oat varieties. Furthermore, Australia has a large potential to improve its dual-purpose oat and hence livestock production.

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Table 1.9 World land utilisationa.

Region Total Area (T) (ha x 106)

Agricultural Potential (ha x 106)

Arable Area (ha x 106) (% of T)

Pasture Potential (ha x 106)

Forestry Area

(ha x 106) (% of T)

Unusable e.g. Desert (ha x 106)

World 13,392 4,407 1,406 (11) 3,001 4,068 (30) 4,917

Africa 3,030 1,047 204 (7) 843 629 (21) 1,354

North America 2,241 627 253 (12) 374 815 (36) 799

South America 1,784 497 89 (5) 408 927 (52) 360

Asia 2,753 893 444 (16) 449 565 (21) 1,295

China 956 287 109 (11) 178 77 (8) 592

India 328 178 164 (50) 14 61 (18) 89

Indonesia 190 13 13 (7) 0 152 (80) 25

Japan 37 7 7 (19) 0 25 (68) 5

Europe 493 240 149 (30) 91 140 (28) 113

Holland 3.6 2.2 0.91 (25) 1.3 0.3 (8) 1.1

UK 24.4 19.4 7.4 (28) 12 1.9 (8) 3.1

Australia 768 467 48 (6) 419 77 (10) 224

a Sassoon (1994).

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While there is a link between oat crop yields and the length of the growing season, on a world basis (Table 1.10) (Forsberg 1986), there also appears to be a link between high oat yields and high population density, but a closer correlation exists between high oat yields and the “nurturing” or constructive policies of the mixed economy (private and socialist enterprises) of the European Union. The small-scale, isolated nature of agricultural production, relative to urban industries in the European countries makes state aid essential. O'Brien (1929) showed how Germany led the world in this respect, followed by Denmark and France. England was indifferent to rural problems, due to her espousal of free trade (see glossary), except in wartime when feverish efforts were made to increase crop yields. Australia has inherited the same predilection for free trade with disastrous repercussions on both agriculture and manufacturing industries. Cribb (1982) portrays in detail this state of Australian agriculture. Free trade is stated to be an “optimal” policy for a small, open and competitive economy. A small economy is defined as having negligible market power and one that cannot influence the equilibrium prices in world markets by its trade policy. Protection may create sheltered markets and monopolies with little incentive for producers to be efficient (Parikh et al. 1988). Fair trade, however, is necessary to protect farm families as is being done in the European Union under the revamped Common Agricultural Policy. Countries will become more self-supporting and trade-restrictive in the future, as free trade inflicts further economic damage on countries like Australia. Table 1.10 Oat yields, growing days, population density and agricultural policy.

Country Yields (t/ha)a Growing daysa People/km2 b Policyc

Ireland Netherlands UK Germany France USA China Soviet Union Australia Spain

4.68 4.52 4.31 3.44 3.32 1.88 1.78 1.28 1.18 1.01

142 155 165 140 150 93

n.a.d n.a.d

200-340 n.a.d

49 421 229 220 99 24 105 12 2 75

Nurturing Nurturing Nurturing Nurturing Nurturing Nurturing Socialist Socialist Capitalist Capitalist

a 1983 data from Forsberg (1985); b Russell and Coupe (1987); c O'Brien (1929) for background; d not available.

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Advantages of grazing the oat crop Craig and Potter (1983), however, point out advantages of grazing the oat crop: (A) Stimulating tillering and increased number of grain producing lateral shoots; (B) Reducing the incidence of fungal diseases common in ungrazed crops; and (C) Reduction in lodging by promoting stronger shoots and removing excess leaf area. Craig and Potter (1983) also found that nearby annual pasture carried 8 ewes/ha giving 1450 kg/ha and 1830 kg/ha feed in early August and early September respectively. Another advantage of oats was the ‘saved pasture’ on which Crofts (1966) carried 7.4 ewes/ha at Orange, NSW. This was twice the rate of grazed ryegrass-clover pastures yielding 4.5 kg dry matter/ha/day and only one-fourth that of heavily seeded oats given N fertiliser (Table 1.11). Table 1.11 Stocking capacity of oats compared with other pasturesa.

Dry Matter Yields Treatment

kg/ha kg/ha/day

Stocking Rate (ewes/ha)

Ryegrass-clover (A) 448 4.5 3.7

Saved Pasture (B) 840 8.4 7.4

Oats @ 90kg/ha seeding rate (C) 1680 16.8 14.8

Oats @ 179kg/ha seeding rate + 67 kg/ha N (D)

3360 33.6 29.6

Ratio of D:A Treatments 7.5:1 7.5:1 8:1 a From the trial conducted in Orange and reported by Crofts (1966). Records for a 100 day winter (late May to August). Clover-grass pastures grow abundantly in early summer but very slowly in winter in comparison with winter or dual-purpose oats. Oats grow 4 to 8 times as rapidly as pasture (Crofts, 1966) during the 100-day winter at Orange NSW, Australia. This result was achieved with the old variety, Algerian, which gave at Richmond NSW, in a separate trial, less than one-seventh the July yield of the High-vigour oat Blackbutt. Therefore oats and pasture are both necessary for good livestock husbandry. To further demonstrate the significance of the oat crop for grazing, even without N fertiliser added, oats gave 4 times the stocking rate given by ryegrass-clover pastures and the dry matter recorded 18% crude protein (Crofts 1966). Crofts (1966) also found that oats should be planted when the mean daily temperature approaches 18°C (or 65 Fahrenheit), which at Orange is about early March, and early April for lower elevations locations. By shutting up large areas of oats in early September, soon after grazing, Crofts (1966) in no time could still recover one tonne of grain per ha. Although drier winters are better for pasturing sheep and cattle on an annual pasture like oats, there is considerable potential untapped in the southern areas of NSW. In the winter rainfall zone at Orange, NSW, Crofts (1966) obtained a remarkable response with Algerian (liable to frost damage in the severe winters in New England), and to heavy rates of seeding and

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nitrogen in 1962 and 1963. Algerian produced 3.4 tonnes per ha, 7.5 times the yield of improved pastures and carried 29.6 sheep per ha, 8 times as many ewes as the clover-ryegrass pasture during the 100-day winter period and carried 30 ewes per ha. The extra yield from nitrogen was less expensive than quality pasture hay. By excluding sheep from the crop from early September onwards, the 1 tonne per ha grain recovery obtained could be sold to offset total costs, including 179kg of seed and 67kg elemental nitrogen per ha. Alternatively, the grain could be kept as a drought reserve (Crofts 1966). Algerian, however, could not produce such yields in a year like 1961 on the New England Tablelands, due to frost damage. Forsberg and Reeves (1995) found that oats, next to rye (Secale cereale L.) are the most versatile of the cereals regarding suitable soil type. Maximum oat yields require soil pH of 5.3-5.7 but can tolerate acid soils with a soil pH of 4.5. Nutrient (NPK) needs for oats are less than those for wheat (Triticum spp.) or corn (Zea mays L) and can be tailored to the desired yield level. Oat production statistics and limitations to their interpretation Overcoming the limitations of statistics is critical in maximising the productivity of the oat crop. Government or industry compiled statistics typically have little value in guiding the direction of research or funding of oat breeding programs. It is more profitable to study a few statistically designed, well-managed and executed yield trials, as described in the latter chapters of this book. The world statistics (Table 1.10) given do not record the 1-2 tonnes of herbage dry matter produced on many farms during the long growing season such as in NSW, Australia. This is not recorded in the state and national yield statistics. This claim is supported by Mengersen (1963) who at that time, estimated that at least 70% of the NSW oat crop is used for dual-purposes production, that is, one or more grazings and then grain recovery. This is a very important aspect of oat production that unfortunately is not reflected in state and national yield statistics. Other examples include the reported statistical oat yields in Ireland. Ireland, although possessing the world's highest average yield, result from late spring sowings and could be boosted further by winter or dual-purpose oats that utilise a longer growing season. The same applies to China as well as to NSW, where late sowings result in low yielding crops that are far behind actual Government research findings in NSW, as recorded here. Oat yields at Cowra, NSW, have been higher than any values reported in world statistics. The research plots at Cowra were not irrigated and yet out-yielded the irrigated trial at Coleambally. At Richmond on the eastern coast of NSW, a dual-purpose oat line, P4315, produced over 10 tonnes of biomass per ha in a dry season (50% of the normal rainfall) without irrigation. Other examples are described in the following chapters. Statistics on oat production in NSW have been documented for many years. These figures give little indication of oat yield potential, however, are of minor value only because they do not always report which varieties were sown, soil types, sowing dates or if the crop was grazed and for how long. This has been the case at least in NSW. NSW oat production statistics, which are composite data compiled by the NSW Government, and grouped from all regions of the state, are criticised because of their failure to show the differences between the summer rainfall northern zone and the winter rainfall southern zone. Furthermore, such oat production statistics do not reflect the total biomass yield or the total value that the oat crop contributes to farming systems. The application of these statistics has led to a frost susceptible

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oat variety being used for pasture research at Armidale (1977), after the frost resistant Blackbutt had already been released to farmers in 1975. Shortcomings in government statistics result in poor funding of oat research. Political economy, sociology and lack of knowledge of agronomy and the value of oats for human and livestock nutrition, are also inhibiting the further exploitation of the oat crop. Figure 1.1 shows cereal production statistics since 1981. In 1990, only Scone and Windouran (both in NSW) showed higher yields for oats than for wheat and barley (Fitzsimmons, 1990). Before 1982, even in 8 irrigation Shires, wheat and barley outyielded oats in production of grain. Beginning in 1982 at Leeton, NSW, oats yields started to rise above those of wheat and barley (Table 4.1, Chapter Four). During the previous 52 years from 1930 to 1982, the wheat and barley yields were always higher than those of oats, and barley was the highest yielder until 1960. From 1960 onwards, wheat took over as the highest yielder of the three cereals and continued so into the 1983 - 1990 period. Shires in NSW in which oat grain yields are superior to those of wheat grain are those in which irrigation is available. In these areas excessive grazing is physically impossible and less likely than the production of crops for grain and this is described in detail in Chapter Four. The Australian statistics (Cribb 1991) are also misleading because oats yield higher than wheat and barley in the medium to higher rainfall zones (>500 mm) and oat breeders have been forced to create early maturing lines like Cooba and Coolabah to be harvestable even earlier than the early wheats. There are other limitations to the value of oat and cereal grain yield statistics. Even when the Bureau of Statistics gives the areas of oats that are grazed, it is important to know the name and characteristics of the actual variety of oats that was grown. In many cases specialised grain varieties, usually from Western Australia, have been grazed in the colder, drier winters of NSW and because of their frost and grazing susceptibility have not recovered much, if any, grain. The comparative statistical cereal yield data presented in Figure 2.1 may suggest that a shorter season variety like Cooba or Coolabah is invariably chosen to avoid a clash with wheat harvest, with the result being a reduced yield. This is counter intuitive because in a wet harvest period, wheat takes much longer to dry out than oats, and in a very short time after rain, the oats can be harvested. This enables better use of expensive harvesting equipment during the harvest season, even if grain storage facilities have to be increased, or organised more efficiently. Longer season cultivars can be safely harvested in the higher rainfall wheat areas, using Glen Innes to breed rust resistance. Another important factor in statistics is the length of the growing season, which can vary from year to year, and the response of the oat variety. This can be broken down in to the following traits: (A) the length of the vegetative period of the variety, (B) its response to frost or vernalisation, and (C) how rapidly it can flower and set grain after the last day of grazing. The “statistical” Australian oat yield is a fraction of the actual yield, due to the grazing tonnage not being recorded. As a result, oat research receives only a fraction of the funds due to it in Australia where every other crop is funded on the basis of its total tonnage delivered to a government agency or entering commercial channels. In turn, Australian farmers are denied the benefits of more thorough and long-term research and plant breeding in oats. Appendix A contains further statistical data on oat yields in Australia.

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Figure 1.1 Statistical yields of the major cereals grown in NSW. Extension of research to oat growers With so much research and extension of information to farmers on other cash crops, relatively little advice appears to be extended to the farmer’s oat crop, which experiences more environmental stress than any other because of its versatility. While plant breeders in the winter rainfall areas of Australia have produced dual-purpose oats with particular emphasis on grain yields, with at least 70% of the total area sown to oats during the 1960s was for dual-purpose grain and grazing (Mengersen, 1963), there is a gap between oat breeding research, which has created new dual-purpose varieties, and farm practice. This is due to a lack of knowledge and skill relative to the oat crop for a particular location, soil and climate. A similar lack of skill in wheat growing would result in uneconomic yields of wheat. Prevention of Ophiobolus attacks on wheat requires that wheat should follow a crop of early sown, well-grazed oats, to kill susceptible grasses that spread Take-all disease (Lazenby and Matheson 1975). Heavy grazing of oats in cool winters will also destroy black oats (A. fatua and others). It is anticipated that the research presented throughout this book on the value of the oat crop to famers will help fill the gap between scientific research into the further development of oats as a pasture and grain crop and farming practice.

1

1.25

1.5

1.75

2

1981 1984 1987 1990

Year

Yie

ld (t

/ha)

wheat barley oats

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CONCLUSIONS There is now ample evidence to demonstrate that oats play an important role in human and livestock nutrition. Oats provides grain for humans and livestock, a grazing or forage crop for livestock, and the health benefits of oats are predominantly the cardiovascular disease prevention properties of oats (through the cholesterol lowering effects of β-glucan) and the glycemic effects of oats which is to reduce the increase in human blood glucose levels. While the significance of the oat grain in benefiting human health has received considerable attention, however, relatively little attention has been given to the important attribute of combined grazing, grain production and total crop value in global agriculture. The oat crop, because of its potential for grain recovery after grazing, has a sigficant role to play increasing global pasture production. Particular oat groups, specifically those with prostrate growth habit, have the required attributes for inclusion in breeding programs to develop suitable oat varieties for dual-purpose capability and therefore for increasing pasture production. The application of oat statistics by government agencies is holding back the attainment of higher oat yields in Australia. Governments should review how oat production statistics are used and recognise that these typically underestimate the overall value of the oat crop in achieving sustainable farming systems.

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CHAPTER TWO

AUSTRALIAN OAT VARIETIES AND A GERMPLASM

INVENTORY FOR BREEDING

In Australia, oat breeding has led to the development of oat varieties for the 3 main climatic regions of this continent. These three climatic regions or zones also exist in the state of NSW. These three regions are as follows: The sub-tropical zone, referred to throughout this book as the summer rainfall zone, also occurs on the coastal areas of Southern Queensland and Northern NSW. The uniform rainfall zone approximates the inland area of NSW from as north as Coonabarabran/Dubbo to West Wyalong/Temora in southern NSW. The winter rainfall climate occurs south of West Wyalong/Temora and includes the Australian states of Victoria, Tasmania, South Australia and Western Australia. An inventory of oat cultivars and their pedigrees is presented in relation to the above climatic regions in which oats are grown in Australia. The inventory tables list the name or accessional line of the oats, their pedigrees and breeder. A description of the Australian oat ideotype is also proposed. INTRODUCTION The objectives of oat breeding in Australia have varied depending on the climate, and the needs of farmers and customers in the regions in which they are grown. The flexibility of the oat plant for grazing as well as for hay and grain production has been an important means of increasing farm profitability. It is important for the understanding of the material presented in this chapter, and the subsequent chapters, that there are three major climatic regions in Australia for the breeding of oats. These are introduced in the following sections and the oat breeders who have developed oat varieties in each of these climatic regions are included. An inventory of oat cultivars, lines and crossbreds are presented and which are arbitrarily grouped into the three broad rainfall patterns that occur in Australia. Identification numbers are given for each cultivar and crossbred and these appear in brackets when referenced in the text of this chapter. Where available, the names of the varieties are provided and the breeder. DEFINING OBJECTIVES IN AUSTRALIAN OAT BREEDING The breeding of specialised oats for grain Western Australia and South Australia have reliable winter rainfall climates which enable oat breeders to select oat varieties for high yield of both grain and hay for export. The cultivar Mortlock (208) from West Australia possesses grains with excellent milling quality and could therefore be useful for crossing with the Glen Innes cultivars. Vertigan (1979) described his varieties Esk and Nile as high yielding and suitable for Tasmania, although one of the parents is the frost susceptible West Australian oat, Avon.

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Apart from Tasmania, the winter rainfall zone of Southern Australia has concentrated mainly on the production of grain only oats. These are not therefore exhaustively dealt with here. Andrew Barr, a prolific plant breeder at Adelaide (South Australia), has created Echidna (206), a dwarf oat, less suitable for milling, as well as a more recent release, Bandicoot, a naked oat (A. sativa var nuda) with a view to human food, pig and poultry formulations (Farrell et al. 1992) and possible value for racehorses (Valentine and Clothier, 1992). Barr has also bred Potaroo for Cereal Cyst Nematode resistance and tolerance. This is a significant disease in South Australia and Victoria. Potaroo, however is a dwarf feed grain oat to replace Echidna. Potaroo is not expected to be a suitable dual-purpose oat for grazing and grain (see following sub-section), even though its disease resistance comes from New Zealand Cape and a wild oat (Avena sterilis).

The breeding of dual purpose oat varieties The flexibility of the oat plant for grazing as well as for hay and grain production has been an important means of increasing farm profitability. The initial breeding and testing work in this field of dual-purpose oats was carried out by the NSW Department of Agriculture. This work started in 1921, and was mostly conducted at Glen Innes, NSW. Glen Innes, at an elevation of 1,128 m. and latitude 29°42’S, is central to, and representative of, the New England Tablelands of NSW. It has a cool, temperate climate with a higher incidence of summer than of winter rainfall and is the best natural environment for the testing of oat stem rust (Puccinia graminis var. avenae) in Australia. The Glen Innes Research Station is also close to the Grafton Research Station, which is naturally and regularly visited by oat crown rust (Puccinia coronata). It is also close to the warmer climates of Inverell, Tamworth, Gunnedah and Narrabri, each providing progressively warmer climates as one travels westwards and into areas of lower elevation. Glen Innes is also close to the fertile Darling Downs region of Southern Queensland. A trial by Dwyer (1934) at Bathurst on the Central Tablelands showed that early sown oats, grazed twice, yielded an average of 6% more grain than ungrazed oats sown at the normal time for grain production (Table 2.1). This research showed that grain yields following pasture can actually exceed grain only yields. After World War II most trials have been lenient grazing until about 1960, when the Author began conducting heavy grazing trials. Vrome, a later maturing and frost resistant variety, demonstrated an excellent response to grazing (Dwyer, 1934). However, two grazings represent what is here defined as lenient grazing. Heavy grazing, involving four grazings and usually a February or early March sowing, is now required to obtain maximum production from the oat crop and to test a new variety.

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Table 2.1 Effect of grazing oats twice, versus no grazing, on grain yield of various cultivarsa.

Cultivar A (sown May) grain

(g) B (sown early March) grain after two

grazings (g) B/A (%)

Algerian 3063 Abruzzes x Victory Fulghum W987 Fulghum x Gidgee Fulghum x Belar Vrome W835 Kareala W998 Total

3,095 2,556 2,953 3,038 2,854 2,925 3,081 20,505

3,266 2,797 2,726 2,782 3,138 3,663 3,394 21,669

105.5 109.4 92.3 91.5 110 125 107

105.7% a Dwyer (1934) at Bathurst, a cool climate. The area of the grain samples harvested could not be located. The Author has therefore calculated the average increase in grain yield as a result of 2 lenient grazings (sown in early March) as approximately 6%. However, the frost resistant variety, Vrome, increased its grain yield by 25%, as a result of the 2 lenient grazings. Subsequent trials, especially at Glen Innes and Tamworth, demonstrate that an important principle of oat plant breeding is here involved and can be utilised by the breeder.

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An objection to the use of grazing oats to supplement pasture production in the winter rainfall areas has been made (Craig and Potter, 1983). In their experiment, they refer to stands of subterranean clover and ryegrass seedlings of 3,500 – 4,000 plants/m2, in comparison to only 230 oat plants/m2. Craig and Potter (1983) do admit, however, that (A) such plant density differences are not comparable, (B) the late May sowing is too late for establishment of the oat crop and (C) they suggest that earlier or even dry sowing of oats might be better. Here it should be stressed, however, that the oats provided some grazing and much grain, over 3 tonnes per ha in the case of Carbeen. Breeding a triple-purpose oat The triple purpose capacity, a concept used here for the first time, is the ability of an oat crop to produce grain after grazing as well as being able to produce grain only. A crop can be considered triple-purpose if it can be grazed and then harvested for grain or grown for grain only. In Finland where only specialised grain oats are being bred, because of the marginal climate, attention to strong vegetative growth, crop architecture and source-sink relations is considered extremely important (Peltonen-Sainio 1992). The old linkage between early prostrate habit of growth and weak straw no longer applies to the new range of oat cultivars bred in northern NSW. In a grain only trial2, under irrigation, the dual-purpose cultivar bred in northern NSW, Blackbutt, yielded over 5 tonnes per ha in the Riverina area at Coleambally. Blackbutt outyielded all the specialised “grain only” oat varieties from West Australia, South Australia and Victoria. This triple purpose capacity was displayed by Blackbutt, due to its strong straw or general resistance to lodging, derived from the wide diameter culms of Garry (one of its parents). The trial at Colleambally included the dwarf oat Echidna, which appeared to lack drought resistance in the dryland trial site at Adelong, NSW (notwithstanding the cool climate). Coleambally and Adelong yields are compared in Chapter Four (see Figure 4.1). This highlights the dangers of breeding for specialised grain only oats. Figure 2.1 shows the strong straw of Garry. The variety Blackbutt topped the grazing and grain yields under dryland conditions at Cowra, NSW as well as under irrigation for grain only in comparison with specialised grain oats bred for the Mediterranean climates of Australia (see Chapter Three). Blackbutt can therefore be considered as a triple-purpose oat variety. This dual-purpose and grain only character is largely due to its fairly short and very strong straw. It is recommended for irrigated farms, where the soil is often too wet to bear the traffic of grazing animals, and thus where grazing is impossible. Grazing reduces the height of the grain crop and so prevents lodging. This eliminates the necessity to use chemicals to shorten crop height. The variety Blackbutt increases this flexibility of management and enables it to be grazed by cattle, sheep or horses. Blackbutt, Carbeen and many other proven high yielding lines were derived from the cross F.Ga (2113 E 57) X VRAF.VRSF (1309 G 57)3, which is the High-vigour line and is described in further detail in Chapter Three. The above male parent (always written on the right hand side) is a more frost resistant and larger grained sister line to Cooba. Cooba is an excellent dual-purpose, but not triple-purpose, oat for less frost liable areas, which was 2 The results of this trial are discussed in further detail in Chapter Four. 3 F = Fulghum, Ga = Garry, VR = Victoria x Richland, A = Algerian, S = Sunrise, which is an early maturing selection derived from a natural cross with Algerian.

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discovered and promoted by Dr Mengersen (Mengersen 1963) at Temora Agricultural Research Station. Cooba is much less resistant to frost damage and water-logging than Acacia and the High-vigour selections, Blackbutt and Carbeen. Cooba’s weak straw results from narrower diameter culms than those of Blackbutt. It should be noted that the Glen Innes oat breeding program had always aimed primarily for high quality feeding grains (see Chapter Five). If these are suitable for oatmeal milling, that is a bonus but in the Author’s view this is secondary to storage and stock feeding quality. AN OAT GERMPLASM INVENTORY All cultivars, lines and crossbreds referred to throughout this book have been given an inventory number. An inventory of oat cultivars, lines and crossbreds are presented and which are arbitrarily grouped into the three broad rainfall patterns that occur in Australia. The reason for this The first of these groups is the summer rainfall areas, namely the eastern coastlands of NSW and Queensland; the Darling Downs of Queensland, the New England Tablelands and the North-West Slopes and Plains. The second group consists of the central and southern inland NSW from Dubbo to the Murray River. This is considered a uniform rainfall area. The third group is the winter rainfall zone which includes the Riverine Plains of NSW, Victoria and South Australia. Tasmania and south-west of Western Australia are also included. Identification numbers are given for each cultivar and crossbred and these appear in brackets when referenced in the text of this chapter. For pedigrees of the older Australian oat varieties, readers are referred to Lazenby and Matheson (1975). It is necessary to have an inventory of germplasm to help distinguish particular selections within crosses and even within old landraces like Algerian and Fulghum. Although both of these lines have been “bulk typed” for many years in the Department of Agriculture foundation seed areas, the Author recorded segregation in both lines for juvenile habit of growth in the F2 summer rust nursery of 1958 (Table 2.2). Similar variability has been reported by Coffman for the related varieties of Red Rustproof and Kanota in the Southern United States (Coffman 1961). Mediterranean-derived landraces that have been very successful in Australia and in the U.S.A. This concept of an assemblage of genotypes has been transferred to the F3 and F4 bulks and the F4 bulk types of the High-vigour cross (Guerin and Guerin 1992). By contrast, Carbeen (Roberts, 1981) is an F6 selection from this cross. Carbeen was the only cultivar tested by Craig and Potter (1983) in the winter rainfall zone of South Australia which increased its grain yield following two grazings. This inventory does not include naked oats within this primary gene pool of A. sativa and A. byzantina. Naked oats are susceptible to threshing damage and reduced germination, as well as discoloration of the groat, caused by Alternaria alternata and A. tenuissima. Also, more animal feeding experiments are needed to justify it. It is difficult to store in bulk and has been intensively studied (Stanton, 1923; Valentine 1990).

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Figure 2.1 Straw strength of various lines and cultivars. (Top Left) Fulghum (F) showing lodging; (Top Right) weak strawed Belar; (Bottom Left) strong strawed Garry, VRBke.F (W4598); (Bottom Right) strong strawed Fulmark.

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CLIMATIC ZONES FOR OAT BREEDING AND PRODUCTION Overview Australia is classed as a continent and therefore it contains most climatic zones found throughout the world (Figure 2.2). For the purposes of oat breeding and production in Australia, 3 main climatic zones of this continent have been identified and in the following sections, the oat varieties discussed in this chapter and book have been assigned to these zone. This assignment of cultivar or line to a zone has been based on the zone in which the line or cultivar was bred. These are: The sub-tropical climate zone, also referred to as the summer rainfall zone, and also occurs on the coastal areas of Southern Queensland and Northern NSW (Table 2.2). The uniform rainfall climate zone approximates the inland area of NSW from as north as Coonabarabran/Dubbo to West Wyalong/Temora in southern NSW (Table 2.3). The winter rainfall climate occurs south of West Wyalong/Temora and includes the Australian states of Victoria, Tasmania, South Australia and Western Australia (Table 2.4). These three climatic regions or zones also exist in the state of New South Wales (NSW) (Figure 2.3). The degrees of resistance for each zone and cultivar requirements are listed (Table 2.5). Figues 2.4-2.14 illustrate examples of the morphology of oat varieties and lines bred in various climatic zones. Summer rainfall zone - Northern NSW and Southern Queensland The large inventory of germplasm for the summer rainfall zone at Glen Innes (Table 2.2) is due to the greater complexity of breeding objectives for dual-purpose oats than in the other two zones. A wider germplasm pool was required for this zone to incorporate all the resistances required. Tribute must be paid to the pioneer oat breeders of Australia, many of whom bred cultivars in this zone. Pridham selected Belar, an excellent milling oat, as well as Sunrise, Lachlan, Mulga and Guyra (Mengersen 1960). Dwyer created Lampton, a spring oat resistant to rust, and various crosses giving great yield potential. Steve Macindoe created the famous high grain quality oats of Acacia, Blythe (good yielder in Tasmania); the female parent of Cooba and inspired all succeeding agronomists with his spartan but humane expounding of the low technology plant breeding equipment at Glen Innes. Carroll created Cooba, 1309 G 57 (line 28) and Fulghum x Garry, 2113 E 57 (line 23); Two older varieties formerly recommended for irrigation have been documented in this inventory. These are Bundy (15) (Guerin, 1965), a midseason dual-purpose oat suitable for milling and with resistance to red-legged earth mite derived from Burke (17) and Mugga (58) (Guerin, 1966), the most winter-hardy oat, equal to Winglen winter wheat in this respect. Mugga has high quality grains and is an excellent late-maturing Tableland oat. If allowed to finish properly, that is grown in a suitable climate, it may prove suitable for milling. Both of these varieties are resistant to smut, but are not resistant to crown or stem rusts. Bundy is also dual-purpose oat with high grain yields in the 1957 drought and Mugga, still the most winter-hardy Australian oat and one of the most frost resistant oats with high grain quality and hectolitre weight, were developed by Guerin (1965, 1966). Mugga is as hardy as the winter wheats, and significantly superior in July grazing production to Windebri and Coolabah. It is, however, later-maturing than Blackbutt and lacked the stem rust resistance and superior yields of Blackbutt. Winter production and frost resistance are two separate traits, genetically. Bundy and Mugga are more resistant than Cooba to frost and waterlogging. The latter

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characteristic made them suitable for the irrigated Riverina trials, until they were replaced by Blackbutt (Komoll and Fitzsimmons 1974) in 1975. A High-vigour cross was identified by its F1 vigour and the continuing “heterosis” and uniformity of its F2 plant progenies (Guerin, 1961). This cross is written as HvII 57-75, referred to as A in Table 2.7. These became “F3 bulks” in 1959 and in the F4 were first “drill-tested” in 1960. They continued to be stable high yielding bulks over time. This observation was similar to Suneson's barley composites (Allard and Handshe, 1964; Suneson, 1956). P4315 (38), one of the High-vigour selections, was outstanding for long-season high biomass yields and was rapid in ripening. Blackbutt, Carbeen and P4315 resulted from this cross.the cross, line 28 x line 23, made by Guerin (Guerin 1961). The large number of useful high yielding selections from this cross suggested that it should be referred to as the High-vigour cross or HvII. P4315 broke the world oat grain yield record of 10.6 t/ha by almost 90% (Fageria 1992) and the UK 1982 world wheat yield record of 15.7 t/ha by 26% at Tamworth Agricultural Research Institute in 1973 (Evans 1996). Where the late sown F3 selection (from F2 residue seed) segregated markedly for maturity, rust resistance or grain type, “bulk typing” was applied. The “ideal type” in both F3 and F4 is represented by “n”, a number of single plants exhibiting phenotypic similarity to one another. The final result was a small number of F4 “directed bulks”, including Blackbutt, which were first drill tested in 1962. Gammie (1990), many generations after its release, describes Blackbutt as among the best dual-purpose (grazing and grain) cultivars in oats, wheat, barley and triticale trials and these dual-purpose attributes are described in detail in the following chapter. Carbeen, also the result of the High-vigour cross, is also classified as a summer rainfall variety although its final selection and promotion was done by Roberts at Temora. Newer varieties, bred by Roberts (1989 a,b) at Temora, include Hakea and Yarran. The origin and description of the High-vigour cross from Glen Innes are detailed in Table 2.6. Garry, a Canadian spring oat, provided rust resistance to the High-vigour cross. Many selections from the High-vigour cross were equal and even superior in grazing yields to that of Fulghum on the black soil of the Liverpool Plains (Komoll 1989). This was hailed by NSW Agriculture, Regional Supervisor, Jim O’Reilly, as a big advance in oat breeding (Kommoll 1989), as the new selections are greatly superior to Fulghum in grain recovery yields, and resistance to lodging, shattering, smut (Ustilago sp.) and stem rust (Puccinia graminis avenae). Cooba, although classified as a summer rainfall variety, because the crosses and early selections were made by Macindoe and Carroll at Glen Innes, the final selection and promotion/extension was done by Mengersen at Temora in NSW (the uniform rainfall zone). Uniform rainfall zone - Southern NSW This uniform rainfall zone has received intensive attention by Dr. F. Mengersen and Mr. G. Roberts. The former developed Cooba and bred Coolabah and Cassia for more lenient grazing and less frosty conditions. The latter developed Carbeen from the High-vigour cross, mentioned above and discussed in detail in Chapter Three, and bred Yarran, Hakea and Eurabbie for less frosty conditions.

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Figure 2.2 Climatic regions of Australia.

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Figure 2.3 A transect of NSW showing the Northern, Central and Southern regions in NSW approximating summer rainfall, uniform rainfall and winter rainfall zones, respectively.

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Table 2.2 Summer rainfall germplasma.

Name/no. (Abbrev.) Pedigree Breeder/Source

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19

Acacia (Ac.) Advocate (Adv.) Algerian (A) A x Lag, 2208G58 A x Lag, W4576 Andrew (And.) Arkwin, W4518 Asia Minor, W2270 Belar (B) Berger (Bg) Blackbutt (Bbt) Blythe (Bly.) Bond (Bo) Boppy (Bop) Bundy (Bdy) Brigalow (Blw) Burke (Bke) Camellia (Cel) Carbeen (Cbn)

VRAF Sunrise x Reid New A = original A x Red Rustproof F8, G0-262-0-0-0-0-1 F8, G0-262-0-0-0-0-2 Bond x Rainbow - - A x Sunrise natural cross ex Alber, Uruguay cv. 28 x 23, pedigree at 47 VRAF ex Tasmania Red A x Golden Rain (White Ligowa x A) x (White Ligowa x A) VR Bke F.WF A x Lampton ex Kherson (Russia) Bo x Alber 28 x 23, pedigree at 41

Macindoe Dwyer (ex Algeria) Carroll Carroll - (ex USA) - Pridham Boerger Guerin Macindoe Pridham - Carroll Dwyer Pridham (ex USA) Guerin

a This section of the inventory is continued on the next page. Figure 2.4 BLACKBUTT variety with medium panicle shape and light brown grains: the longest grazing season cultivar of Australian winter cereals. Photograph is from Australian Oat Varieties by R.W. Fitzsimmons et al. CSIRO (1983).

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Table 2.2 Summer rainfall germplasm (continued)a.

Name/no. (Abbrev.) Pedigree Breeder/Source

20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70

Cedar (Ced) Clintafe (Cf) Cooba (Cba) 1309 G57, Hv II parent Dasix Delair (Del) Delta Red (DR) Fulghum (F) 1183 G57, HvII parent 400 G59 W4584 W4597 W4605 2116 E57 Garry (Ga) Guyra (G) High-vigour cross (HvII) 843 G59 P4315 (851 G59) P4314 (856 G59) P4317(856/1G59) Carbeen(869 G59) P4322 (871 G59) P4318 (871/1 G59) P6093 (871-80-11) P6094 (871-80-16) P4316 (886 G59) Blackbutt (886/1 G59) 898 G59, Improved Algerian (I.A.) Iogold (Io) Kanota (Kan) Kent (K) Klein 69B (K69B) K69B G.R. (W4571) Laggan (Lag) Lampton (L) Landhafer (Lf) Mugga (Mga) Rodney (Rdy) Saia Santa Fé (SF) Seminole (Sem) Sual Ukraine 0545 Victoria.Richland (VR) Victory (Vy) Winter Fulghum (WF) Winter Palestine (WP) Yarran (Y) Yates Algerian (YA)

VR, Victoria x Richland Clinton x Santa Fé VRAF.VRSF, F7 G0-102-H0-G0-0-4 VRAF.VRSF, F8 G0-102-H0-G0-0-0-2 ex Sixty-Day cv. Fulghum x Bond ex Red Rustproof ex USA, segregating F x Garry, G0-56-0-10 Fx Garry, G0-98-0-1. Large grained Fx Garry G0-56-0-5. High grain yield Fx Garry G0-56-0-6. High test wt. F x Garry G0-56-0-3 F x Garry G0-81-0-1 High test wt. Vy x (V.Hajira-Banner) ex Canada White Ligowa x A (Same as Lachlan) 28 x 23 28 x 23, G6-0, F3 bulk 28 x 23, G13-0, F3 bulk 28 x 23, G17-0, F3 bulk 28 x 23, G17-n-n, F4 bulk type 28 x 23, G29-90-3-0-3,F6 plant 28 x 23, G31-0, F3 bulk 28 x 23, G31-n-n, F4 bulk type 28 x 23, G31-80-11, F4 plant 28 x 23, G31-80-16, F4 plant 28 x 23, G44-0, F3 bulk 28 x 23, G44-n-n, F4 bulk type 28 x 23, G55-0, F3 bulk Largest grained A cultivar ex Kherson, like Richland ex Fulghum Ballidu x (Mulga x Laggan) ex Argentina G0-0-R27, F4 plant ex Kelsalls cv. Abruzzes.Vy. Reid ex Germany VR Bop B W4527, (V. Hajira-Banner) x Roxton A. Strigosa sp. ex South America Appler x Clinton-Santa Fé Rust resistant A. (ex-University of Sydney) - ex USDA ex Milton cv., Baltic/German landrace ex Fulghum cv. ex Palestine cv. White Ligowa x A Latest A cv., only A since 1959

(ex USDA) (ex USDA) Carroll Carroll (ex Canada) - (ex Mississippi) Fulghum Carroll Carroll Carroll Carroll Carroll Carroll Welsh Pridham Guerin Guerin Guerin Guerin Guerin Guerin Guerin Guerin Guerin Guerin Guerin Guerin Guerin - Burnett (ex Kansas) Bateman Klein Carroll Hurst Dwyer Gassner Carroll Welsh (ex Brazil) - - Baker (ex Russia) Murphy Nilson (ex USA) Palestine Pridham ex A(3)

a 0 = a bulk of all plants derived from a single F2 plant where the succeeding long F3 row appeared uniform and distinct from other rows; n = a bulk of plants in F3 and F4 selected for phenotypic similarly in rows segregating for rust resistance, grain type, maturity, and height; G = Glen Innes, H = Hawkesbury Agricultural College, R = Grafton.

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Table 2.3 Uniform rainfall germplasm.

Name/no. (Abbrev.) Pedigree Breeder

101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123

Avon Ballidu (Bl) Cassia (Cia) Coker's Fulgrain (CF) Coolabah (Cab) Dale (Dl) Eurrabie Fulmark (Fk) Gidgee (Gid) Hakea Markton (Mt.) Mulga (Mlg) Osage (Os) Sunrise (S) Victorgrain (Vic) Yarran (MA3236) M1179 M1255 M1305 W4465 M1300 M1309 M1301

Bl x Mlg x Lag Mlg x Burt's Early CF/2 x Osage V x Fulgrain (ex USA) Fk.Bl.Dl.F. Mlg x Burt's Early Echidna x M3227 (Involves crossing of Avon, Fulmark, Ballidu, Kent, Cooba) (F x F x Gid) x Mt White Ligowa x A Kent/O//Cba/3/Cab - ex Sunrise (S) cv. Fulton x VR Segregating ex A. V x Fulgrain M127/Radar 2//M1335/ M1345/3/Tamo 312 Avon x Fk Avon x Os Avon x W4465 VR Bke F.WF F x Avon F x Avon F x Vic.

Bateman Bateman Mengersen Wilds Mengersen Bateman Roberts Kitamura Pridham Roberts (ex Turkey) Pridham Coffman Pridham Wilds - Roberts Mengersen Mengersen Mengersen Carroll Mengersen Mengersen Mengersen

Figure 2.5 COOLABAH is an early grazing and grain variety with medium panicle shape and cream coloured grains. It is too frost susceptible for the summer rainfall germplasm list. Photo is from Australian Oat Varieties by R.W. Fitzsimmons et al. CSIRO (1983).

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Table 2.4 Winter rainfall cultivars.

Name/no. (Abbrev.) Pedigree Breeder/Source

201 202 203 204 205 206 207 208 209 210 211 212 213 214

Algeribee (Agb) Barmah Bulban Bundalong Dolphin Echidna Esk Mortlock Nile Orient (O) Palestine (P) Stout West Moore

ex New Algerian (Agb.Ga) x Avon (Agb.Ga) x Avon Cayuse x Avon West x OT207a West x OT207a Bly x Avon Elan 6161/3/(66Q01-63)Fk/ Newton/Swan Bly x Avon P x Dawn - - K/Bl(M127)/Radar 2 Fulmark/Newton/Swan

(ex Werribee) Brouwer Brouwer Brouwer Barr Barr Vertigan Portman Vertigan Raw (ex Palestine) (ex USA) Portman -b

a OT207 has dwarf mutant gene W6; bWest Australian Dept Agriculture. Figure 2.6 ORIENT is an erect early midseason variety for grain only, with medium to open panicle and dark brown grains. It is too frost susceptible for the summer rainfall germplasm list. Photograph from Australian Oat Varieties by R.W. Fitzsimmons et al. CSIRO (1983).

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Table 2.5 Crosses combining rust resistance with agronomic value.

Cross Parents Female Male Genotypes Phenotype

A B C D E F

28 x 23 0600a x F 0614b x B 1.A. x 0600 0615c x W4477d 1244e x B

SRRf SRR SRR SRS MSRR SRR

SRSg SRS SRS SRR SRS SRS

dissimilarh " " " " "

similar dissimilar " " " "

a 0600 = (Bo x Rainbow) x (Hajira-Joanette) x Lf, ex Minnesota, USA b 0614 = R + Canadian + White Tartarian, SRR, ex Minnesota, USA; c 0615 = Kubanca + White Tartarian, MSRR, ex Minnesota, USA d W4477 = VR Bke F.WF (sister to Bundy) e 1244 G57 = F.Ga, F5 bulk, very early, very large grains f SRR = stem rust resistant; M = moderate; R = resistant; g SRS = stem rust susceptible; S = susceptible h All crosses are genetically dissimilar (A. sativa x A. byzantina) but only cross A is phenotypically similar to some extent with dual-purpose qualities in both parents, although morphologically different. Cross A produced only 147 plants in the F2, out of which 62 progenies were sown in F3 long rows. Table 2.6 Segregation in landraces for juvenile growth habit, Glen Innes 1958.

Landrace TPa Erect Semi-erect Prostrate

Algerian Fulghum

25 68

15 48

7 14

3 6

aTP = Total number of plants observed in row.

Figure 2.7 ALGERIAN variety with open panicle and mid-brown grains from Australian Oat Varieties by R.W. Fitzsimmons et al. CSIRO (1983). Table 6.5 shows that both Algerian and Fulghum were segregating for juvenile habit of growth in the 1958 F2 summer rust nursery, confirming Coffman’s claim that the related varieties of Red Rustproof and Kanota in the USA could not be fixed.

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Table 2.7 Resistances for various environmentsa.

Rainfall zones Cultivar Resistance or Requirement Summer Uniform Winter

Frost Smut (Ustilago) Crown rust (grazed) Crown rust (ungrazed) Stem rust BYDV Waterlogging Drought Lodging Heavy grazingd Lenient grazinge Grain only grain recoveryf Long season Short season

H H L H H H H Hb H H L L H H L

M H L M L M M Hc M

M-H L M H M M

L H L L L L M Hc M L H H

L-M L H

a H = high; M = medium; L = low (levels of resistance or requirement); b regular autumn drought; c regular summer drought; d 4-5 grazings; e 1-2 grazings; f grain yield after grazing.

Figure 2.8 COOBA is a mid-season grazing and grain variety with open panicle and mid brown grains from Australian Oat Varieties by R.W. Fitzsimmons et al. CSIRO (1983). Cooba is inferior to Blackbutt and Carbeen for grazing and frost resistance in the summer rainfall zone.

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Table 2.8 Origin and description of genotypes developed from the High-vigour cross, 28 X 23.

F3 No 1959a

F2 single plant

yield of seed Crown rust

(1-6) Stem rust

(1-6) Smut

0/seg. Final result (Name/No)

838 1460 segb seg 0 F3 bulk (too early-discontinued)

843 360 seg 1- seg F3 bulk (frosted-discontinued) 851 600 1- 1- 0 P4315, F3 bulk (top yielder) 856 300 seg seg seg P4314; F4 bulk type is P4317 869 420 2+ seg 0 Carbeen, F6 plant progeny 871 360 seg seg seg P4322; F4 bulk type is P4318,

BYDV tolerantc 886 240 seg 1- 0 P4316; F4 bulk type is P4319 or

886/1 (Blackbutt) 898 420 6 1 0 F3 bulk (discontinued)

a Out of 62 single plants selected in F2, Blackbutt was released in 1975 and Carbeen in 1981; b Segregation required bulk typing while fixing disease resistance; c Tolerant to BYDV.

Figure 2.9 CARBEEN variety with condensed panicle shape and medium brown grains. A mid-season variety with prostrate early habit of growth, the most adaptable to the 3 rainfall zones. Photograph from Australian Oat Varieties by R.W. Fitzsimmons et al. CSIRO (1983).

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Figure 2.10 FULGHUM spiklets and florets from Oat Identification and Classification by T.R. Stanton (1955) US Department of Agriculture Technical Bulletin No. 1100. Fulghum is a semi-winter type and appears to be of hybrid origin, with many traits intermediate between the northern common oats, A. sativa, and the southern red oats, A. byzantina, as judged by observers in the US.

Figure 2.11 FLORET SEPARATION distinguishes the 2 types of cultivated oats which are Avena sativa (on the left), separating by distal fracture, and A.byzantina (on the right), usually separating by basal fracture. Photograph from F.A. Coffman, Inheritance of Morphological Characters in Avena, Technical Bulletin No. 1308, Agricultural Research Service, United States Department of Agriculture.

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Figure 2.12 MORPHOLOGICAL CHARACTERISTICS OF THE OAT PLANT, showing (1) panicle and spikelet, main rachis and panicle branches; (2) rachilla and basal hairs of mature grain; (3) spikelet showing pedicel, glumes, rachilla, primary grain and secondary grain and awn on the primary grain; (4) culm nodes and nodal hairs and (5) leaf margins and leaf sheaths, both hairy and glabrous. From Anonymous (1962).

2.

1.

4.

5.

3.

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Figure 2.13 The Author assessing mature oat crop stands. (Top) Avon x VRFB; (Bottom) Garry x VRBke (2056) on the right and Fulghum (see summer rainfall germplasm inventory) on the left.

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Figure 2.14 Mature oat crop stands. (Top) The Author with a tall strong strawed line; (Bottom) Avon (see the uniform rainfall germplasm inventory) on the left and taller W4477 on the right.

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For the uniform rainfall zone, Mengersen (1960, 1963 and 1968) has described Algerian (3), Cooba (22) and Coolabah (105), respectively, as high yielding dual-purpose oats. Mengersen’s variety Cassia (103) is a high yielding feed grain oat described by Fitzsimmons et al. (1983). These varieties have not been used for crossing at Glen Innes because of their frost susceptibilities. Such exclusion is not always a logical plan as transgressive segregation for frost resistance is common in oats where only one parent is frost resistant. Winter rainfall zone Apart from Nile, Bass and Esk, three high-yielding dual-purpose cultivars bred by Mr. W. Vertigan in Tasmania, most varieties from this southern rainfall zone have tended to be grain only varieties. This is suggested by trials carried out at Coleambally, Adelong and Temora (all in NSW) and described in Chapter Three. Mr. A Barr of Adelaide has created unique varieties in this grain only category, the dwarf oats, Dolphin and Echidna, with the mutant gene W6; Potoroo with resistance and tolerance to cereal cyst nematode (CCN) a problem in South Australia and Victoria; and Bandicoot, a naked oat with exciting possibilities for animal diet formulations for pigs, poultry and racehorses. More recent varieties bred in South Australia are described by Zwer (2005). Varieties from Victoria were Palestine, Dawn, Algeribee, Alpha and Orient, the largest-grained Australian oat, as tested at Glen Innes by the Author. The breeders of many of these Victorian varities, along with varieties from other locations, are provided in Tables 2.3. AN AUSTRALIAN OAT IDEOTYPE The concept of an ideal Australian oat variety (or Ideotype) is one that is dual-purpose and preferably sown in February or March (August to September in the Northern Hemisphere if winter is not excessively severe or if the growing location is not too far north). An effective program of breeding and testing dual-purpose oat varieties would include the following goals: (A) Selection for resistance to frost, smut, stem rust, and BYDV; (B) Grain quality should be considered as more important than millability. Grain quality should therefore be assessed by its weight in kilograms per hectolitre, at least 50, and the groat percentage (1000 dehulled grain over the 1000 whole grains tested) should be at least 70; (C) Total yield or biomass of herbage and high quality grain. Hay yields, straw height and lodging should be recorded; (D) requires strong vegetative growth and strong straw; and (E) strong competitive habit of growth compared with that of weeds. Instead of herbicides, which involve both extra cost and canlead to potential adverse environmental impacts, oats can better utilise the long fallow as they can be sown four months earlier than wheat or barley and therefore with less risk of soil erosion.

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CONCLUSIONS The oat genetic material developed at Glen Innes, NSW has not been fully exploited. The greatest potential for oat expansion in Australia is in the northern half of NSW and Southern Queensland, the eastern coastal regions, and the irrigated and dryland Riverina areas of NSW and Victoria. In contrast to these areas are the marked winter rainfall climates and soils of Western and South Australia. These are states which produce specialised grain oats and are aiming to supply the export market for milling and feed oats. This type of specialization should be supplemented by the breeding of triple purpose cultivars at Glen Innes, NSW, and adjacent regions. This would enable the generation and exchange of new oat genotypes to take place, giving rise to greater adaptability in the crop and a more productive oat ideotype would result from the implementation of such action. On balance there seems no reason why pasture oat varieties cannot be grown in the southern rainfall zones of Australia. On the other hand, the frost susceptible specialised grain oats of Western Australia cannot be grazed in Central and Northern NSW. The agronomic advantages of grazing oats have many beneficial effects for “whole farm” management that they cannot be wisely neglected. As the winter rainfall climate of Western Australia is generally the most productive of oat grain of any Australian climate, it would benefit growers in NSW if seed of the frost and grazing resistant oat varieties could be grown in Western Australia, where seed growers could be encouraged with a premium price or bonus. This could also lead to the incorporation of dual-purpose oat varieties into farm management and rotation systems.

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REFERENCES Allard, R.W. and P.E. Hansche. 1964. Some parameters of population variability and their implications in plant breeding. Advances in Agronomy 16: 281-325. Anonymous. 1962. The Aberystwyth Varieties of Oats, University College Wales, Wales. Coffman, F.A. 1961. Oats and oat improvement. Madison: American Society of Agronomy. Craig, A.D. and T.D. Potter. 1983. The effect of grazing on the yield of ten oat cultivars in the south-east of Australia. Agricultural Record 10: 4-7. Dwyer, L. 1934. The effect of cutting back oats on the grain yield. NSW Agriculture Report, pp.1-2. Evans, L.T. 1996. Crop Evolution, Adaptation and Yield. Cambridge University Press. pp. 288-289. Fageria, N.K. 1992. Maximizing Crop Yields. Marcel Dekker, Inc. New York. Farrell, D.J., B.S. Takhar, E. Thomson and A.R. Barr. 1992. The nutritional value of naked oats in broiler, layer and duckling diets. In 4th International Oat Conference. Edited by A.R. Barr, R.G. McLean, J.D. Oates, G. Roberts, G. Rose, K. Saint, and S. Tasker. Fitzsimmons, R.W., G.L. Roberts, and C.W. Wrigley. 1983. Australian Oat Varieties, CSIRO. Gammie, R.L. 1990. Orange district crop trial results. NSW Agriculture and Fisheries, pp.2-3. Guerin, P.M. 1961. Breeding new oat varieties for northern NSW. Agricultural Gazette NSW 72: 1-7. Guerin, P.M. 1965. Bundy - The breeder's report: A new winter-hardy oat for the Northern Tablelands. Agricultural Gazette NSW 76: 667 - 669. Guerin, P.M. 1966. Mugga - The breeder's report: A new winter-hardy oat for the Northern Tablelands. Agricultural Gazette NSW 77: 675 - 678. Guerin, P.M. and T.F. Guerin. 1992. Breeding oats for irrigation in Australia. In Fourth International Oat Conference, Adelaide, Edited by A.R. Barr, R.G. McLean, J.D. Oates, G. Roberts, G. Rose, K. Saint, and S. Tasker, pp. 187–190. Komoll, R.F. 1989. The North West. The memoirs of Jim O'Reilly, p88. NSW Agriculture.

Komoll, R.F. and R.W. Fitzsimmons. 1974. A new variety, Blackbutt…released. Agricultural Gazette NSW 85, 6: 8. Lazenby, A. and E.M. Matheson. 1975. Australian Field Crops. I. Angus and Robertson, p.490. Mengersen, F. 1960. Oat improvement in NSW. Agricultural Gazette NSW 71: 449-461. Mengersen, F. 1963 Choosing oats for grazing and grain in Southern NSW. Agricultural Gazette NSW 74: 678-683. Mengersen, F.1968. Coolabah, a new dual-purpose oat for the central and southern districts. Agricultural Gazette NSW 78: 633-636. Peltonen-Sainio, P. 1992. Description of a productive oat ideotype characterised by morpho-physiological traits associated with high grain yield. University of Helsinki. Publication No. 34. Roberts, G. 1981. Oat variety-Carbeen. Agnote: NSW Agriculture, Agdex 113/33. Roberts, G. 1989a. Avena sativa cv. Yarran. Australian Journal of Experimental Agriculture 29, 1: 144-145. Roberts, G. 1989b. Avena sativa cv. Hakea. Australian Journal of Experimental Agriculture 29, 1: 146-147.

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Stanton, T.R. 1923. Naked Oats. Journal of Heredity 14, 4:177-183. Suneson, C.A. An evolutionary plant breeding method. Agronomy Journal 48: 188-191. Valentine, J. and R.B. Clothier. 1992. The development of naked oats in the UK. In Proceedings of the Fourth International Oat Conference, Adelaide. Edited by A.R. Barr, R.G. McLean, J.D. Oates, G. Roberts, G. Rose, K. Saint, and S. Tasker. Vertigan, W.A. 1979. What varietry of oats should you sow? Journal of Agriculture (Tasmania) 1979: 6-7. Zwer, P. 2005. Oat variety sowing guide. Primary Industries and Resources, South Australia.

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CHAPTER THREE

THE NEW ISOLECTION PLANT BREEDING SYSTEM

The result of 34 years of oat breeding and testing of dual-purpose varieties (for grazing and grain recovery) by the NSW Department of Agriculture are summarised in this chapter. A High-vigour cross (HvII 57-75) is identified which led to the release of Blackbutt (an F4 directed bulk type) in 1975, and Carbeen (an F6 plant progeny of the normal pedigree system) in 1981. This High-vigour cross also produced a number of high yielding F4 directed bulk types and F2 plant progenies bulked in the F3 as a result of their relatively high phenotypic uniformity. The highest yielding F3 bulk was numbered P4315, which although classed as an early oat, out-yielded all other varieties, including Blackbutt, for total biomass, following early sowings, and over a wide range of soils and climates and numerous seasons. The success of these oats was due to the Isolection plant breeding system pioneered by the Author at Glen Innes from 1957 to 1964. Other F4 directed bulks were P4314 (high-yielding both as a winter oat and a spring oat at Glen Innes) and P4318, both of which had large grains and, together with Blackbutt and P4315, were significantly superior over 5 grazing cuts (including the mid-winter cut) to Coolabah and all other advanced lines submitted by plant breeders from Temora NSW, that were using conventional breeding methods during the same period. Selection of lines at the F2 generation has been demonstrated as a simple way of forecasting wider adaptability of early generation genetic material.

INTRODUCTION The NSW Department of Agriculture (now the Department of Primary Industries) has been a world leader in the breeding of dual-purpose oat cultivars since 1921. This Department also has a strong tradition of releasing uniform oat varieties (meaning selection is in the sixth, F6, generation or later). These may take up to 15 years from making of the cross to release to farmers. With the present restriction of funds to Australian agricultural research, benefit/cost ratios in oat breeding research could be greatly improved by adopting a rapid “directed bulk” oat breeding method. This chapter describes the Isolection method which led to the breaking of world record yields. The following quote by Falconer (1975) sets the context for this chapter: “When large differences of environment, such as between different habitats, are under consideration, the presence of genotype-environment interaction becomes important in connection with the specialisation of breeds or varieties to local conditions…the important partition is into additive genetic variance versus all the rest…This partitioning is most conveniently expressed as the ratio of additive genetic to total phenotypic variance, Va/Vp, a ratio called heritability”.

Falconer (1975) notes that natural selection, takes no account of heritabilities or genetic correlations and is therefore less efficient in improving fitness than artificial selection. The standard Mendelian genetic ratios, found with the differences between blue-eyed and brown-eyed individuals, are not usually exhibited in quantitative genetics, which depends on

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gene differences at many loci, the effects of which are not individually distinguishable. Nevertheless, the inheritance of quantitative differences depends on genes subject to the same laws of transmission and with the same general properties as the genes whose transmission and properties are displayed by qualitative differences. Quantitative genetics, however, is simply an extension of Mendelian genetics. THE METHOD FOR SELECTING HIGH-VIGOUR OAT CROSSES Overview The scientific principles of Isolection plant breeding are essential for the breeding and testing of the oat crop. The process of breeding, however, must be separated from that of testing. This is dictated by the facts of Mendelian genetics and the danger of losing valuable material if testing is too severe and too early in the process of breeding. When the Author arrived at Glen Innes from Ireland in 1956, early breeding material was still being unevenly sown through the farm drill. This resulted in uneven stands, with individual plants unable to express and visually expose their yield potential or heritability, either for grazing or for grain production. In 1957, the Author spent a week with Dr. Fred Mengersen at Temora, learning to achieve a high percentage of success in crossing oats, before returning to Glen Innes, on the New England Tablelands, where spring came a little later and in time to make the necessary oat crosses. This was an excellent climatic centre for stem rust (Puccinia graminis avenae) and smut (Ustilago sp.) inoculated nurseries. Glen Innes was also close to the crown rust (Puccinia coronata) nursery at Grafton research station, where rust developed earlier in the season and supplied inoculum for the Glen Innes disease nursery. This was followed by a spaced plant rust test to study infection and segregation. The May sown F3 hand spaced rows were judged for uniformity and segregation of growth habit (from prostrate to erect) and for grain type and quality. Hand sown rows were opened up with the Planet Junior (a seeder manufactured by Powell Manufacturing) and were 2 foot (or 610 mm) apart. Plant density for the Isolection system is 3.66-5.38 plants/m2, as compared with 13.99-21.53 plants/m2 for the conventional system which it replaced during the development of the Isolection breeding method. More importantly, however, individual plants were far enough apart to express their potential yields relative to one another. The duplicate F3 sowing in August was for smut testing. The F4 test sown in early March was the first yield comparison, sown with the farm drill, and was subjected to the same rigours of winter grazing and frost as the normal biometrically designed and analysed small plot trials. Table 3.1 shows that the seeds of the hybrid plants were grown in the potato breeder’s glasshouse to ensure no loss of germplasm to grazing or frost.

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Table 3.1 Rapid method of breeding oats for large biomass yields.

Generation Date sowing Date harvest Growing season Testing sequence

F1 F2 F3 earlier (normal) F3 later duplicate F4

31.3.1958 19.11.1958 7.5.1959 11.8.1959 1.3.1960

9.10.1958 7.4.1959 31.12.1959 7.1.1960 17.12.1960

6 months 4.5 months 8 months 5 months 9.5 months

Glasshousea increase Spaced plant rust test Spaced plant grain test Smut and maturity test grazing and grain trialb

a Field cross 26 x 21 made on 15.10.1957; b2 years from cross.

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Table 3.2 Morphology and pathology of parents of the High-vigour crossa.

Cross A Female Male

Pedigree Homozygosity Early growth Mature height Harvest index Grazing-frost Foliage Seed wt. (g/1000 seeds) Groat % Hectolitre weight (kg/hectolitre) Milling suitability Grain articulation (specific trait) Ustilago (smut) Crown rustc Stem rust

F.Ga (1183 G57) G0-56-0-10 (F5) Fixity proven Prostrate Tall (122 cm) Medium Very hardy (1/6) Medium broad 32.30 70.50 51.13 Too small 50% A. sativa 50% A. byzantina Resistant MR (2/6) R (0/6)

VRAF.VRSF (1309 G57) G0-102-H0-G0-0-0-2 (F8) Fixity proven Prostrate Short (90 cm) High - 47.2%b Very hardy (1/6) Narrow leaves 44.00 74.00 54.24 Excellent 100% A. byzantina - Susceptible MR (3/6) S (6/6)

a Data obtained from row averages at Glen Innes 1959; b Harvest index determined subsequently at Tamworth 1961; c Crown rust is less important inland because grazing and cool winters discourage rank foliage growth.

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More importantly, however, yields were greatly improved where homozygosity was observed in the F2 and F3 rows, as in the case of P4315. In the case of Blackbutt, the F2 bulk (886 G59) was found to be segregating for habit of growth in the F4, where only the truly prostrate plants were harvested to constitute the F4 bulk type, P4319. An excellent team were available to determine grain quality on wet days and were happy to do hand spaced sowings (some of them had market gardens) and to cut the oat pasture to 1 inch (2.5 cm) above ground level using hand shears to avoid the inevitable loss of herbage involved with machine shears. This is consistent with more recent research on the Southern Tablelands of NSW (Dann et al. 1983), which found that grazing by sheep or cattle down to 2 cm did not reduce grain recovery yield any more than grazing down to 6 cm. The early generation plan in Table 3.1 enables F3 bulks and F4 directed bulks to enter drill-testing respectively at 2 years and 4 years after the cross is made. For example, P4315 (F3 bulk) entered trials in 1960 whereas Blackbutt (F4 directed bulk) entered trials in 1962. Only certain crosses (phenotypically not too dissimilar) described concurrently should be attempted. The morphology and pathology of the High-vigour parents are described in this chapter (Table 3.2), with the view to making the most effective cross or crosses. For example, a cross between a winter oat variety and a spring oat variety, like Fulghum x Garry, is only a preliminary cross to transfer disease resistance from Garry to Fulghum. In Cross A (Table 3.2), a line from this cross was selected which combined Fulghum’s hardiness with Garry’s disease resistance. This close crossing concentrates the multiple genes needed for each type (Guerin 2003). Features of the Isolection breeding method The features of Isolection breeding are: (A) High rate of success in crossing oats, achieved in 1957, before starting, in order to produce a large number of homozygous F2 plants; (B) The two parents to be phenotypically similar (as in a narrow cross) but genotypically different; (C) The F2 generation plants to be widely spaced by hand, 4.52 plants/m2, at Glen Innes, as against 17.76 plants/m2 plants for the conventional drill sowing at Temora Research Station (representing the southern wheat belt). Hence the name of Isolection system, to “isolate” pure breeding lines, like P4315, and “select” them for yield testing in F3. The F2 plant of P4315 produced 600 seeds; (D) Linkage assists the rapid breeding method, by telling us that a winter cereal has morphological features like prostrate habit of growth and deep root system, correlated with resistance to frost, drought and grazing damage.

In about 1960, the Author replaced the previous conventional trial system of only 2 grazings per trial with one of 4 to 5 grazings, the latter being followed by a grain recovery trial. This enabled identification of a deeper root system, resistance to more severe frost and drought, and medium size grain with high bushel weight and low husk percentage, compared to Algerian’s large husky grains (from the Mediterranean centre of origin). This benefit of quality proved that high total yields could be combined with high grain quality. Selection under low-stress conditions The 1966 trial at Richmond was a comparison of the conventional and isolection methods of breeding. The new Isolection breeding method developed at Glen Innes Research Station, requires all early generation plants to be hand spaced, in order to provide the non-stress conditions to facilitate selection for heritability (Figure 3.2 and 3.3). This is to increase the chances of obtaining progeny with the desired characteristics required by the oat breeder. The conventional breeding system requires the early generation plants to be sown with a farm or

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other type of drill. Even at low rates of seeding, it is impossible to ensure even spacing of oat seeds. This is certainly more difficult than in the case of wheat seeds. It was clear that population stress conditions (obvious in a dry period following the germination of masses of seeds released out of the farm drill at the end of sowing each variety of seed) were detrimental to all varieties in the trial. As a result of this technicality and lack of labour, conventional breeding usually means selecting plants only in a later generation, as in the F6, where the type selected is presumed to be fixed. However, it should be noted that Algerian and Fulghum are still segregating in the F6 (see Chapter Two) It is important to emphasise that non-stress selection will not reduce yields on the farm or in any trials where the optimum rate of seeding for maximum yields per unit area must be used, as the trials in this chapter demonstrate. As a reference point, in the Netherlands or Southern England, the density of field crops would be about 35 plants/m2. This concept of isolated selection, was later developed theoretically using a formula for heritability, h2, to obtain the additive breeding value, VA, giving: h2 = VA/VP (phenotypic value) (Falconer and Mackay, 1996) The total variance is the phenotypic (non-additive genetic and environmental) variance, VP, that needs to be reduced, in order to increase heritability percentage. Because of the true breeding nature of homozygotes, it is possible in the F2, to rapidly obtain a pure race with respect to any combination of parental factors provided that a large enough F2 generation was grown and tested. This concept is illustrated in the work conducted by the Author while breeding oats for NSW Agriculture at Glen Innes after 1956. His predecessor, James Carroll, had retired several years earlier from plant breeding and had already selected suitable lines from a moderately wide cross that he had made to incorporate crown and stem rust resistance from the Canadian oat Garry. A moderately wide cross, in this context, means a cross between different ecospecies like a winter oat, Avena byzantina var. Fulghum and a spring oat, A. sativa var. Garry, not a very wide cross like wheat x rye, which are different species. Nevertheless, a yield reduction is always involved but was easily overcome by only one cross in 1957, later referred to as the High-vigour cross (HvII 57-75): [F.Ga (1183 G57)], the female parent, x [VRAF x VRSF (1309 G57)], the male parent, where F = Fulghum, Ga = Garry, V = Victoria, R = Richland, A = Algerian and S = Sunrise were in the ancestry of the two 1957 rows at Glen Innes Research Station, NSW, Australia.

A number of other wider crosses were made to study linkage, but only this one close cross, the HvII, was necessary to accumulate the many genes for yield, frost resistance, drought resistance, tolerance to BYDV, resistance to smut, crown rust and stem rust. Figure 3.4 shows VRAF, one of the lines contributing to the genetic makeup of the male parent of this High-vigour cross. The Isolection system has since been proven to assist in the detection of heritability, by several other workers, including Professor Kenneth Frey, although the mechanism responsible was said to be unknown at the time (Frey 1964). The non-stress environment (that is, separate sowing by hand) makes it possible to select the highest possible yielding lines, while the close spacing of a drill sowing does not. Professor Frey found similar results with oats at Iowa. He found that non-stress conditions resulted in the retention of oat strains with a wide adaptation reaction, whereas the stress conditions did not (Frey 1964). He referred to a formula of Falconer on heritability that confirmed his oat results (Falconer 1952).

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In Japan, USA and the Netherlands, trials support the value of the non-stress conditions for the Isolection breeding system. Japanese wheat breeders also found that selection for heritability of yield was higher under high fertility than under low fertility conditions (Gotoh and Osanai 1959). Breeders of spring rye, Secale cereale L., in the Netherlands, concluded that single plant selection for yield at wide spacing gives a higher progress and allows a better identification of outstanding genotypes, due to freedom from interplant competition (Pasini and Bos 1990). Explaining “bulk” varieties A clear distinction must be made between blends or multilines versus varieties compounded in F3, F4 or F5. Acceptable uniformity was achieved with an F5 line of soybeans (Auckland 1967) and with Blackbutt oats compounded or bulked in the F4 (Guerin 2003). A blend is artificially compounded of unrelated homozygous subpopulations and unlikely to maintain equilibrium. By contrast the early generation compounded variety has aggregate homeostasis, or greater stability in yield. This results from making and comparing crosses within a particular crop ecospecies. Crossing between different ecospecies, like winter growing types x spring types, ignores the need for multigenic breeding and often results in the need for backcrossing and in lower yields. Early, not later, generation selection is suggested by the Mendelian diagram for a trihybrid or three factors (Guerin 2003). In summary, high yield requires many genes. Testing for dual-purpose capability in oat varieties The importance of dual-purpose oats, especially in Eastern Australia, is so great that special emphasis has been given to grazing production research in NSW since 1921. The nutritional value of grazing exceeds that of the grain, which in itself has a higher biological value than the other cereals, both winter and summer crops (see Chapter One). The value of grazing relative to grain also increases by heavy, as opposed to lenient grazing and as one travels from south to north into the drier, sunnier regions with lower winter rainfall, richer soils and greater incidence of frost damage. The standard grazing measurement techniques are as follows: (A) Continuous grazing, (B) Heavy grazing, (C) Lenient grazing, (D) No grazing, grain only, (E) Grain recovery after pasturing, (F) Total biomass, and (G) Pasture cut technique. (A) Continuous grazing. Variations of this such as “lax”, “put-and-take” and rotational have been described by Wheeler (1962). They are expensive methods involving animal weighing, wool or lamb weights and few have been planned to incorporate new dual-purpose oats, bred in Australia in the past 25 years. Spurway (1975) is an exception in this respect because a suitable Northern NSW Tableland oat cultivar, Acacia, and an early February 25) sowing were studied, giving valuable knowledge to the Northern NSW Tableland sheep and wool-grower. The variety Cooba is not a true dual-purpose one as was still assumed (Fitzsimmons, 1978), when Archer and Swain (1977) tested the early maturing Cooba with a late March sowing, two fixed and unsuitable parameters making valid results questionable, at least for best Northern NSW Tablelands practice. Farming experience, together with trial results, both prove the necessity for early sowing as against late sowing for achieving greater total yields. This is demonstrated by comparing yields from different NSW Tableland sowing dates (Tables 3.3 and 3.4).

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Table 3.3 The effect of grazing intensity on a range of cereal genotypes sown late March in a cool, moist, summer rainfall climate: F6 generation trial of High-vigour lines (1962)a.

Heavy grazing (t/ha) Lenient grazing (t/ha) Frost (0-6) Cultivar (no.)

T4Pb pGc TYd T2Pe pG TY 4P 2P

K69B.G.R, F4 (54) Blackbutt (11) 886 G59 (46) Blythe (12) 856 G59 (39) Wintok P4315 (38) 871-1 G59 (43) Acacia (1) 856-1 G59 (40) Winglen (W) Bundy (15) Klein 69B (53) Fulghum (27) Algerian (70) Cooba (22) Mugga (59) Dun/Grey Winter Abyssinian (B) NIAB (WR) Means SD CV (%)

2.60 2.24 2.15 2.47 2.07 2.43 2.26 1.96 2.07 1.87 1.77 1.98 2.11 2.32 1.55 2.13 1.62 1.70 1.53 0.89 1.99

- 33.2

0.60 0.90 0.95 0.47 0.82 0.36 0.47 0.69 0.57 0.69 0.78 0.57 0.43 0.21 0.90 0.31 0.44 0.47 0.50 0.50 0.58 0.30 35.0

3.20 3.14 3.10 2.94 2.89 2.79 2.73 2.65 2.64 2.56 2.55 2.55 2.54 2.53 2.45 2.44 2.06 2.17 2.03 1.39 2.57

- -

2.29 1.95 1.76 2.08 1.87 2.08 2.25 1.93 1.80 1.82 1.59 1.84 1.99 2.56 1.63 2.34 1.48 1.42 1.28 0.83 1.84

- 23.9

0.67 1.05 1.00 0.98 0.72 0.79 0.86 0.95 0.68 0.82 1.06 0.73 1.11 0.64 0.91 0.70 0.69 0.61 0.81 0.62 0.82 0.30 35.0

2.96 3.00 2.76 3.06 2.59 2.87 3.11 2.88 2.48 2.64 2.65 2.57 3.10 3.20 2.54 3.04 2.17 2.03 2.09 1.45 2.66

- -

1 0 0 0 0 0 1 1 0 0 0 1 1 2 2 2 0 0 0 0

0.6 - -

2 0 0 0 1 0 3 3 1 1 0 3 2 4 4 4 0 0 2 0

1.5 - -

a Trials were conducted at The New England Research Station, Glen Innes, NSW (cool moist climate). Date of sowing was 29.3.62. Season: Slow early growth; wet harvest weather. Date of harvest 30.1.63. This is not a comparison between heavy and lenient grazing due to the time difference in final grazing; b = total of 4 pasture cuts taken 15.6.62 (estimated); 7.8.62; 5.9.62; 2.10.62; c = grain recovered after pasturing; d = total yield of grazing and grain recovery = total biomass/energy; e = total of 2 pasture cuts taken on 15.6.62 (estimated) and 5.9.62.

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Table 3.4 The effect of grazing intensity on a range of cereal genotypes sown in early March in a cool, moist summer rainfall climate: F7 generation trial of High-vigour bulk oats (1963) at Glen Innesa.

Heavy grazing (t/ha) Lenient grazing (t/ha) Cultivar (Inventory Number) T4Pb pGc TYd T2Pe pG TY

Blackbutt (11) 871-G59 (43) Klein 69B (53) Cooba (22) Blythe 12) Fulghum (27) 856-1 G59 (40) K69B.G.R., F4 (54) 856 G59, (39) Winglen (W) P4315 (38) Wintok 886 G59 (46) Algerian (70) Bundy (15) Mugga (59) Acacia (1) Abyssinian (B) Dun (Grey Winter type) NIAB (WR) Means (excluding B & WR) SD CV (%)

3.85 3.88 3.20 3.23 2.79 2.89 2.53 2.38 2.35 2.31 2.18 2.10 1.80 1.83 2.09 1.84 1.66 1.95 1.05 0.15 2.44

- 30.5

0.47 0.42 0.70 0.29 0.47 0.18 0.35 0.49 0.50 0.26 0.28 0.35 0.59 0.42 0.13 0.36 0.35 NILf 0.45 0.03 0.39 0.19 36.1

4.32 4.30 3.90 3.52 3.26 3.07 2.88 2.87 2.85 2.57 2.46 2.45 2.39 2.26 2.22 2.20 2.01 1.95 1.50 0.18 2.84

-

2.80 2.38 3.29 2.95 2.15 2.06 2.49 2.95 2.21 2.39 3.40 1.86 2.58 1.74 1.98 2.08 1.75 1.59 1.79 0.14 2.38 1.08 33.2

0.48 0.25 0.75 0.17 0.43 0.16 0.48 0.44 0.31 0.30 0.19 0.38 0.50 0.34 0.22 0.39 0.39 0.03 0.43 0.09 0.37 0.21 43.2

3.28 2.63 4.04 3.12 2.58 2.22 2.97 3.39 2.52 2.69 3.59 2.24 3.08 2.08 2.20 2.47 2.14 1.62 2.22 0.23 2.75

-

a Date of sowing was 6.3.63 at the New England Agricultural Research Station, Glen Innes, NSW. Severe grazing was measured by 4 pasture cuts before each close even grazing by a large flock of sheep. Lenient grazing was similarly measured by 2 pasture cuts; b = total of 4 pasture cuts; c = recovery for grain harvest; d = total yield of grazing and grain recovery; e = total of 2 pasture cuts. W = wheat; B = barley; WR = winter rye; f = barley failed to recover after severe grazing. Grazing dates: 1.5.63; 24.5.63; 5.9.63; 3.10.63 (heavy series); 24.6.63; 18.9.63 (lenient series). Season: June very wet; dry after September grazings. Date of harvest 30.1.64.

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(B) Heavy grazing. This involves 4 or more grazings. This is synonymous with intensive grazing and is in between continuous (in all its variations) and lenient grazing. This reveals more accurately than (C) cold weather and other growth rates. A better term to describe this might be “close” or frequent grazing. (C) Lenient grazing. This involves only one or two grazings and is intermediate between heavy grazing and no grazing. The trials presented in this chapter show that heavy grazing gives a higher total yield than lenient grazing including Tables 3.5-3.8. Lenient grazing is incapable of revealing the true dual-purpose cultivars with high protein yields of herbage undamaged by severe frosts, and also likely to be of a higher digestibility (Pearce et al. 1987; Guerin 1966). Lenient grazing causes more frost and shade damage than heavy grazing as in Cooba and Fulghum (Table 3.3) and decreases digestibility (Fagan and Milton, 1931). A better term to describe this might be “lax” or infrequent grazing. (D) Grain only trial. They are a reference point for grain (or hay) yield potential, lodging resistance and susceptibility to diseases like crown and stem rusts, because ungrazed oats can grow tall and rank. (E) Grain recovery after pasturing. This is especially important in the case of lenient grazing but also in the case of heavy grazing, where the third and fourth cuts of a frosted variety are still yielding well, although much of the material has been “burnt” by frost and is probably low in digestibility but not showing a significant inferiority in grazing yields. This has been the case at Glen Innes, as recorded by Guerin (1966). In this particular case it is the grain after grazing (pG) yield that demonstrates the significant yield superiority of the true dual-purpose oat in conjunction with total yield (Tables 3.3 and 3.4). (F) Total yield or total biomass. This is the final arbiter of the value of the dual-purpose oat for energy purposes. On a weight-for-weight basis, however, the dry matter pasture yield (if not damaged by frost) has twice the protein content of the grain. This is important for growing, fattening, lactating and pregnant animals. On the other hand it is a waste of high quality feed to put dry adult stock on to oats. (G) Pasture cut technique. Only the “crash grazing” (Dann et al. 1977), or severe grazing technique is used, both for heavy and lenient grazing comparisons. This technique prevents selective and uneven grazing by sheep, who are “shy” grazers and must belong to a flock large enough to eat the experimental area down to a constant level of about 2 cm (within a few days) above ground level (Hodgson et al. 1981) (Figure 3.1). To avoid soil contamination of the cut herbage, the NSW Department of Agriculture at Glen Innes used hand sheep shears. This gives the operator more control of cutting height, and less noise to contend with, than in the case of power-driven equipment. Moreover, the work is done more accurately by this method. As most of the good varieties are prostrate growing, power-driven equipment would be less accurate. Even with 4 replications of each treatment, variation of grazed oats under the stress of the cold climate of the Tablelands in NSW, and sometimes waterlogging, is often high. The first grazing and cut are taken about 6-8 weeks after sowing. A fresh sample of the plot length is taken before each grazing, as sheep tend to over-graze these protein rich areas. This is due to a vigorous emergence of new shoots in the case of high tillering cultivars like Blackbutt. These new shoots are rapidly devoured by the sheep during the day or two they occupy the experimental area. It is for this reason that good grazing cultivars like Blackbutt actually produce much more grazing in practice than grazing

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trials (even those grazed four times) actually record. Because of the prostrate, low-growing habit of these cultivars, a motor mower cannot replace the hand cutting method. The converse is also true. Fertility is transferred by the sheep from the good grazing plots to the poor grazing plots which therefore record a higher yield than they would in farm practice. Sheep are therefore considered essential for testing the full potential of the oat crop. TESTING OF HIGH-VIGOUR LINES AND VARIETIES Yield increases from High-vigour lines and varieties Extensive yield testing of the High-vigour oat lines and varieties have been carried out since late 1950s. Table 3.4 shows the overall superiority of oats bred at Glen Innes compared with conventionally bred lines for frost resistance, cold weather growth to supplement pastures and long season intensive grazing with hay recovery on the richer type of soil in summer rainfall regions. This trial, conducted in Richmond NSW in 1966, also demonstrated the superiority of the Isolection breeding technique over conventional breeding and this is further discussed in Chapter Six Table 3.5 shows the value of grazing, by return of nutrients through the grazing animal (Wilson 1968) in boosting grain yields, an option not usually open to specialised grain oats, wheat or barley varieties. Here, P4315, Sual, Cassia and Blackbutt broke the world oat yield record (Fageria 1992), with P4315 proving significantly the highest yielder for the 2 pasture cuts. The oat varieties Cassia, Coolabah, Hakea (Roberts 1989a) and Yarran (Roberts 1989b), although suitable for lenient grazing, are not true grazing-hardy, dual-purpose oats as is the High-vigour line, P4315. Carbeen is only slightly handicapped by its straw being taller than Blackbutt, while Hakea is taller and weaker than Carbeen. Yarran is susceptible to BYDV. Tolerance to this disease is essential under high and summer rainfall conditions. Lenient oat grazing dry matter yields in South Australia (Craig and Potter 1983) were only 14% of those of continuously grazed pasture. By contrast, Algerian oats seeded at 90 kg/ha yielded 4 times that of ryegrass-clover pasture (Crofts 1966). The same variety seeded at 180 kg/ha with 67 kg/ha of nitrogen carried 30 ewes per ha or 8 times the carrying capacity of the pasture at Orange (Crofts 1966). At Armidale, which is similar in climate to that of Canberra and typical of much of eastern Australia, the ratio of oat to pasture growth in areas of dormant winter pastures (which contain no sub-clover) was 10:1 (Wheeler 1963). A comparison of the Isolection lines with conventionally bred oat lines from Temora Research Station (NSW) and other winter rainfall areas was made in 1966 at Hawkesbury Agricultural College, Richmond, NSW (Table 3.6). The highest yielding lines were all from the High-vigour cross and were identified as P4315, P4314, Blackbutt, 871-1 G59 and 871 G59, in that order, all significantly higher yielding than conventional lines, in 5 grazing yields and a hay recovery cut. All 5 lines produced grain of high test weight and low husk percentage, ideal for stock feeding. The top 5 yielders tested in this trial, by 5 grazings and a hay cut, were High-vigour lines. In this trial, P4315 produced over 10 tonnes of biomass per ha in a dry season (50% of the normal rainfall), with no irrigation. P4315 also produced the highest mid-winter growth of 1.45 tonnes in 33 days. This data is presented in Table 3.6 and

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the trial plots are shown in Appendix B. This daily production of 44 kg would support 35 sheep per ha, assuming that each sheep required 1.24 kg daily (Crofts 1966). Blackbutt was the highest yielder for total grazing, demonstrating its high recuperative ability. Blackbutt is a very high tillering variety with winter prostrate habit. P4314 was the highest yielder of hay. Blackbutt was only the sixth highest yielder of hay, indicating its short straw and lodging resistance. Cooba and Coolabah were found to be unsuitable for dry winters and were low yielders under heavy grazings. As these are the two most popular varieties in the state, their replacement by Carbeen and Blackbutt would, for most of the state, considerably increase livestock productivity and farm profitability. At Tamworth Agricultural Research Institute, in 1973, the early variety P4315 yielded significantly more than most varieties for 2 grazing cuts and recovered 19.83 tonnes of grain per ha (Table 3.8). In the late-maturing class, Blackbutt has yielded significantly more than all other oats, winter wheats and triticales, for grazing and grain recovery, from 1966 to 1999, on the Tablelands, Cootamundra and eastern Australia generally. It is still recommended in 2005 (McRae et al. 2005). The high grain after grazing yield of P4315 in the Tamworth trial (Table 3.8), broke the world grain oat yield record of 10.6 t/ha by 90% (Fageria 1992) and the world wheat yield record of 15.7 t/ha by 26% (Evans 1996). This can be explained not only by the excellence of the season of 1973 at Tamworth but also by consistent and sound soil management practises like contour cultivations and fallowing for weed control, ample soil water storage and early plantings over many years and, above all, the dedication of the Agronomist, Mr. Gerry Hennessy, who conducted the trial. Lenient grazing in this trial may have facilitated the high grain recovery yields. These ideal growing conditions enabled the full expression of the genetic potential of the varieties and lines tested in this trial. Varieties bred at Temora and research stations in the winter rainfall area of southern Australia are unlikely to be productive in the cold winters and summer rainfall of the northern New England Tablelands of NSW as set out in 2 statistically designed trials, typical of many tests, in 1969 (Table 3.9). The outstanding selection in these 2 trials and other cold winter areas, P4315, had an exceptionally high rate of growth in the coldest month of the year, July. It alone was significantly superior in this respect to Acacia and Klein 69B from Argentina, hitherto the best mid-winter varieties when livestock have their greatest food requirement. Seed of P4315 was not maintained in the germplasm collection transferred to Temora, contrary to the Author’s request for release of this High-vigour line to farmers in 1966. The Author recommended that P4315 should replace Cooba because of greater frost resistance and equally early maturity for the convenience of wheat growers, rust resistance, higher hectolitre weights and much greater straw strength. P4315 had proven to be the highest yielding oat line variety of all lines and varieties tested in NSW by the Author or in trials that the Author is aware of. Figure 3.2-3.5 and 3.10 illustrate some of the varieties and lines used and developed during the breeding of the High-vigour cross, their spacing in the field, their parents, and field testing.

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Figure 3.1 Results of heavy grazing by sheep (Top); The pasture cut technique using manual shears at Glen Innes, NSW (Bottom).

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Figure 3.2 (Left) Tall strong straw of Fulghum (F) x Garry (Ga) (F.Ga or W4595), typical of the F.Ga cross; (Right) Close up of the panicles of F.Ga, the female parent of the High-vigour cross.

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Table 3.5 Second testing of High-vigour bulk oats in north-west NSW, contrasting cooler elevated site (Tamworth) with warmer plains site (Narrabri): F5 generation trial (1961).

Tamworth (mild but frosty) Narrabri Cultivar (Inventory Number) CWGa

(t/ha) Frostb (0-6)

T4Pc (t/ha)

pGd (t/ha)

TYe (t/ha

pHf (t/ha)

G/Hg %

Gh t/ha

P4315 (38) F.Ga (30) Klein 69B (53) 871 G59 (42) A x Lag (4) 856 G59 (39) Fulghum (27) K69 B.G.R. (54) Acacia (1) 886G59 (46) Algerian (70) Cooba (22) Mugga (59) 1309 (23) Bundy (15) 843 G59 (37) Burke (17) Belar (9) Orient (210) SDj Date of sowing:

1.71 1.44 1.40 1.61 1.24 1.40 1.05 1.19 1.33 1.11 0.93 1.34 1.02 1.14 0.83 0.75 0.58 0.52 0.22

0.31

14.3.61

0 1 0 1 0 0 1 1- 0 1 4 3 0 1

2+ 4 4 3 6 -

6.97 6.06 6.45 6.56 5.37 6.15 6.03 5.29 5.66 5.35 4.91 5.99 4.86 5.20 4.58 4.76 4.11 3.82 4.18

1.61

1.63 2.48 1.78 1.60 2.31 1.46 1.49 2.15 1.59 1.79 2.09 0.86 1.91 1.05 1.59 1.29 1.24 1.14 0.43

0.55

8.60 8.54 8.23 8.16 7.68 7.61 7.52 7.44 7.25 7.14 7.00 6.85 6.77 6.25 6.17 6.05 5.35 4.96 4.61

-

3.61 -

4.15 3.51

- 2.88 3.12

- - - -

2.07 -

2.18 - - - - -

0.46

45.2 -

42.8 45.1

- 48.6 48.0

- - - -

44.8 -

47.2 - - - - - -

3.5.61

1.09 0.52i 2.69 1.19 0.24i 0.71

- -

0.50i 0.28i 1.05 0.82 0.05i 0.69 1.38 0.74 1.71 2.09 4.38

0.41

a Cold weather growth (CWG) measured by 3rd cut; high fertility soil; b Frost damage score of 3 and over lowers grain recovery yields; c Total of 4 pasture cuts, each immediately followed by sheep grazing trial area down close to ground level on 16.5.61, 10.7.61, 23.8.61 and 19. 9. 61; d Grain yields after grazing; e Total yield of c + d; f Hay recovered after grazing at grain ripeness stage; g Relative harvest index, grain as a percentage of hay; h Grain only trial, high fertility soil; i Specialised winter oats, too late maturing for the north-west plains. All yields in tonnes/ha, with pasture as dry matter; j

All trials are biometrically designed and analysed to give significant differences in yields between all the lines tested.

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Table 3.6 A comparison of southern and northern NSW bred cultivars under intensive grazing and hay recovery: F10 generation testing of High-vigour lines at Richmond (1966). Cultivar (Inventory Number)

5P (t/ha)

pH (t/ha)

5P+pH t/ha

Tillers killeda

F score (0-10)

July P (t/ha)

P4315 (38) 856 G59 (39) Blackbutt (11) 871-I G59 (43) 871 G59 (42) Klein 698 (53) Cooba (22) Fulghum (27) F x V (122) Coolabah (105) F x Avon (121) Avon x Fk (116) Avon x Os (117) F x Avon (120) Fulmark (107) M1305 (118) Yates Algerian (70) SD

6.55 6.21 6.67 5.66 5.60 5.01 5.18 4.87 4.21 4.09 3.89 3.96 4.04 3.45 3.78 3.36 3.38 0.90

3.62 3.70 2.86 2.97 2.99 3.37 2.21 2.20 2.47 2.08 2.23 1.93 1.81 2.11 1.70 1.48 0.60 0.99

10.17 9.91 9.53 8.64 8.59 8.38 7.39 7.07 6.68 6.17 6.12 5.90 5.85 5.57 5.48 4.85 3.98 1.54

4 0 4 0

3.6 0 14 30 19 76 43 40 157 106 165 132 35 -

1 1- 1 2 2

2+ 3+ 3

4+ 6+ 4+ 7+ 8 7 9 7 8 -

1.45 1.23 1.35 0.83 0.74 0.72 0.95 0.64 0.52 0.45 0.36 0.28 0.33 0.23 0.20 0.25 0.19 0.34

a No. per row; 5P = 5 pasture cuts as DM (dry matter); pH = air dried hay after pasture; F = frost (0 = no damage, 10 = extreme); Date of sowing 25.3.66 at Hawkesbury Agricultural College, rich alluvial soil, no irrigation; dry season (rainfall 50% of 86 year mean); seeding rate 72 kg/ha; rate of fertilizer, in seed drill only, 22.4 Kg N + 22.4 P2O5/ha. Cultivar inventory reference numbers with 3 digits were bred in Temora, a Conventional breeding centre, and those with 2 digits, excepting Klein 69B, Cooba, Fulghum and Algerian, were bred in Glen Innes using the Isolection breeding system; SD = significant difference, obtained by biometrical analysis performed by NSW Agriculture Biometricians at Rydalmere, NSW, Australia, during 1966-1967.

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Figure 3.3 Non-stress growing environment. A plastic covered frame (Top) for establishing rust infected plants, transplanted from the subtropical station at Grafton, and designed to spread rust and determine rust resistant plants; (Bottom) Inspection of individual oat plants.

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Figure 3.4 Non-stress growing environment (Top) Fulghum x Garry (female parent of the High-vigour cross showing) showing its strong straw; (Bottom) Wide spacing of individual oat plants.

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Figure 3.5 Crossing of a rust resistant line, of oat, 0600 (Top) and VRAF (W4890) (Bottom). P4315, the highest yielder, was solely derived from an F2 plant that yielded 600 seeds, of which 540 seeds were spaced out in row 851 G59 in the F3 generation, as set out with other

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pedigrees in Chapter Two (Table 2.8). In the 1966 trial at Richmond NSW, it produced more tillers per plant (3.3) than Fulghum (2.6) and Cooba (2.5). The groat percentages of the grains were equal to those of Cooba, in a wide range of climates. Although the grains were smaller than those of Cooba, they were better for stock feed with their higher weight per bushel, due to its resistances to rust diseases and to lodging. Mr. Norm Markham, District Agronomist for West Wyalong, informed the Author on 21st April, 1972 that P4315 line was exceptional for grazing in the Grenfell area (Guerin 2003). Many other similar trials in eastern Australia have been conducted or recorded (Guerin 2003). Results at Cowra showed that 3 High-vigour lines headed by Blackbutt, produced from 7 to 8 tonnes of total yield (Table 3.7) (Guerin and Guerin 1992). This was better achieved by 4 grazings than by 2 grazings (i.e. the lenient series). The frost susceptible varieties like Cooba and Coolabah performed better with lenient than with heavy grazing. This significant finding, from Cowra, Richmond, Glen Innes, Orange and other trials throughout NSW (NSW Agriculture, 1960-1977) has not been taken into account by later research work. For example, Lovett and Matheson (1974) studied only lenient grazing and did not realise that oat varieties bred for heavy grazing were superior to wheat, barley and rye for total biomass energy and, because of higher grazing yields, one could confidently state much higher protein yields (Tables 3.9 and 3.10). Similarly Craig and Potter (1983) did not study beyond 2 grazings and their sowing in late May was too late for maximum oat grazing winter production which should significantly outyield pasture growth (Table 3.11 and 3.12). Algerian was badly frosted in the Nepean valley at the River Farm of Hawkesbury Agricultural College (Richmond, NSW) in 1966, as illustrated in Figure 3.5 and reflected in the yield data (Figure 3.6). The straw of Algerian, like that of Klein 69B, is tall and weak and unsuitable for grain only crops. In hectolitre or bushel weight (weight per unit volume) and groat percentage (weight of whole grains less the hulls), the grains of KIein 69B are like those of Algerian, which are inferior to those of the Glen Innes oats. In the 1966 trial referred to above, it was a dryland sowing which received only 50% of the 85 year mean annual rainfall and was a notably dry and frosty season. The five top yielding varieties in this trial were derived from the High-vigour cross, as shown in Figures 3.7-3.9. A total biomass yield of 5 grazings and a crop of hay amounted to over 10 tonnes per ha in the case of P4315, the highest yielding line in the trial. By contrast Algerian produced only 3.4 tonnes per ha, similar to Croft's yield from 5 grazing cuts at Orange in the Central West Tablelands of NSW (Crofts 1966). In addition, the Richmond 1966 trial received one-third of the nitrogen applied by Crofts at Orange. The fertilizer was applied only at sowing time and the lower seeding rate suited the season and the soil, a rich alluvial clay loam which encouraged the high tillering capacity of the High-vigour lines. The stocking capacity of P4315 was potentially equivalent to 42 adult sheep per ha for 255 days, on the basis of the results at Orange. Professor Lloyd-Davies, however, regards dry matter yields as more reliable than stocking rates4. Figure 3.6 shows the poor grazing capacity of the specialised grain oat, Swan, from Western Australia, in comparison with P4315 and Cooba at Temora, a site with less frost damage than Richmond or Glen Innes (both located in NSW).

4 Personal communication (2005).

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Figure 3.6 A typical Western Australian bred cultivar, Swan, showing poor dry matter recovery under a 5 grazing cut regime at Temora, New South Wales, 1969, in comparison with moderately frost-hardy Cooba and very frost-hardy P4315.

0

0.2

0.4

0.6

0.8

1

1.2

1 2 3 4 5

Num ber of cuts

Yie

ld (t

/ha)

Cooba Swan P4315

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Table 3.7 Heavy (4 P cuts) and lenient (2 P cuts) grazing and grain recovery (pG) at Cowra: F10 generation testing of the High-vigour lines (1966)a.

Heavy grazing (t/ha) Lenient grazing (t/ha) Cultivar (Inventory Number) 4P (DM)f pG Total Y 2P (DM) pG Total Y (R)Blackbutt (47) P4315 (38) P4314 (39) Coolabah (105) Fulghum (27) Cooba (22) Avon x Os (117) F x Avon (120) 871-1 G59 (43) 871 G59 (42) Bundy (15) F x Vic (122) Blythe (12) Avon x Fk (116) Acacia (1) Algerian (70)

2.49 2.95 2.72 2.47 2.83 2.56 2.11 1.78 2.49 2.25 2.30 2.32 2.50 1.90 2.26 2.11

5.26b 4.74b 4.81b 4.26b 3.65b 3.60b 3.30b 3.43c 2.70c 2.62c 2.41c 2.31c 1.68c 2.20c 1.80c 1.67c

7.75 7.69 7.53 6.73 6.48 6.16 5.41 5.21 5.19 4.87 4.71 4.63 4.18 4.10 4.06 3.78

0.86 1.11 0.80 1.36 1.33 0.99 1.17 1.18 0.97 0.92 1.18 1.02

- 1.26

- 1.08

6.21e 6.42e 6.12e 5.43e 4.38e 5.61e 3.83d 5.37e 3.37e 5.71e 6.06e 3.69e

- 5.20e

- 2.45d

7.07 7.53 6.92 6.79 5.71 6.60 5.00 6.55 4.34 6.63 7.24 4.71

- 6.46

- 3.53

SD CV (%)

n.a. -

n.a. -

- -

0.23 14.4

0.71 10.3

- -

a Pasture cuts were weighed green and converted to DM (dry matter) by using factor of 20% dry matter as found at Tamworth Agricultural Research Institute, which has a mild winter climate like that of Cowra but is more subject to frost damage than is Cowra; Date of sowing 25.3.66 (heavy grazing) and 25.3.66 (lenient grazing); b Date of harvesting was 12.12.66; c Date of harvesting was 22.12.66; d date of harvesting after heavy grazing was 22.12.66; e date of harvesting after lenient grazing was 30.11.63; f Dates of grazing were 17.5.66, 22.6.66, 25.7.66 and 29.8.66 (heavy grazing); g n.a. = Heavy grazing results not analysed.

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Table 3.8 Lenient grazing and grain trial: F17 generation testing of High-vigour bulk oats. Cultivar (Cultivar Number)

Pasture Yield (2 cuts) (t/ha)a

Grain Recovery Yield (t/ha)b

Total Yield (t/ha)c

P4315 (38) Sual (63) Cassia (103) Blackbutt (11) Cooba (22) Klein 69B (53) Coolabah (105) Bundy (15) Algerian (70) Mugga (58) Avon (101) Acacia (1) SD CV (%)

0.58 0.49 0.47 0.48 0.62 0.57 0.51 0.54 0.47 0.45 0.43 0.53 0.08 10

19.83 19.66 16.14 14.50 10.36 7.26 7.31 5.95 5.98 5.62 3.92 3.80 1.71 12.4

20.41 20.15 16.61 14.98 10.98 7.83 7.82 6.49 6.45 6.07 4.35 4.33

- -

a Total of 2 pasture cuts to measure 2 grazings; b Grain recovery yields were a NSW Department of Agriculture and world record for oat yields; c Total yield of grazing (as dry matter) and grain in tonnes/ha. Excellent season; crown rust and stem rust severe but not affecting P4315 or Sual. Date of sowing 4.5.73 at Tamworth Agricultural Research Institute; d = coefficient of variation.

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Table 3.9 Effect of multiple grazing cuts on grain and pasture yields on a range of oat cultivarsa.

Site 1 Yields (t/ha) Site 2 Yields (t/ha)c Variety

Cut 1b Cut 2 Cut 3 Cut 4 Cut Total Cut 1 Cut 2 Cut Total Grainc

Acacia (1) 1.07 1.66 0.97 0.31 3.96 1.12 0.99 2.11 1.38 Algerian (70) 1.02 0.94 0.66 0.21 2.81 1.27 0.79 2.06 1.26 P4315 (38) 1.28 2.19 1.34 0.26 5.12 1.17 1.71 2.85 2.10 Blackbutt (47) 0.90 1.76 1.43 0.46 4.56 1.07 1.47 2.52 2.07 Cooba (22) 1.10 1.52 1.08 0.30 3.96 1.17 1.59 2.75 1.56 Coolabah (105) 1.07 0.87 0.79 0.16 2.86 1.42 1.02 2.45 1.08 Fulghum (27) 0.93 1.13 0.80 0.24 3.11 1.71 1.55 3.25 1.32 Klein 69B (53) 0.99 1.52 1.27 0.36 4.14 1.30 1.33 2.66 1.83 Mugga (58) 1.20 1.40 1.30 0.38 4.31 1.12 0.97 2.08 1.65 Saia (60) - - - - - 1.55 0.97 2.52 1.40 Abyssiniand - - - - - 1.18 1.68 2.86 2.05 Windebrie 0.53 0.63 0.84 0.16 2.16 - - - - Cutting datef 65 148 247 276 - - - - - SD (t/ha)g 0.48 0.48 0.48 0.48 1.05 0.21 0.39 0.48 0.40 CV (%)h - - - - 39.3 9.9 21 11.5 17.3

a Trial conducted in New England, NSW in 1969; Site 1-sown 4 March 1969 with 4 replicates at Mr. Shireberg’s farm, Niangula, and Site 2 - sown 2 April 1969 with 3 replications at Mr. Lane’s farm, Reddestone, NSW; b Dry weight of pasture sampled at the designated cutting date; c Grain harvested after two pasture cuts; d Barley variety; e Winter wheat variety; f Days after sowing (not recorded for Site 2); g Standard deviation; h Coefficient of variation.

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Table 3.10 Grain and pasture yields from 1955 competing crop trialsa.

Pasture Yield (t/ha) Crop Variety

Cut 1b Cut 2c Totald Grain Yield (t/ha)e Frost Damagef Ear Emergence

(Days after sowing)g

Oats Fulghum (27) 2.36 2.51 4.87 0.717 3 203

Oats Acacia (1) 2.46 1.84 4.30 1.375 3 119

Oats W4477 1.57 2.22 3.79 1.008 2 206

Oats W4484 1.79 1.84 3.64 0.654 4 199

Barley Cape 1.79 1.59 3.39 0.790 4 204

Barley Pryor 1.62 1.59 3.21 0.840 4 204

Wheat Celebration 1.12 0.89 2.01 0.914 5 214

Winter Wheat Winglen 1.29 0.66 1.96 1.451 1+ 216

Winter Wheat

German P4 Cel. 1.18 0.63 1.81 1.116 3 217

a The trial was conducted in 1955 at Glen Innes in New England, NSW and data originally reported by Mr. Lancaster (1956, unpublished); b Cut was taken on 5 May 1955 (44 days after sowing); c Cut 2 was taken on 30 August 1955 (161 days after sowing); d Significant difference=1.10t/ha and standard deviation=23.4%; e Significant difference = 0.252t/ha and standard deviation = 17.0%; f On a scale of 0-6, Winglen was the least frost damaged and Celebration was the most damaged variety; g Sowing date was 22 March 1955.

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On the New England Tablelands, pastures which were ungrazed from mid February to June-July produced 515 kg per ha in 1958 and 280 kg per ha in 1959, an average of 397 kg per ha (Wheeler 1963), which is similar to that at Orange in 1966 (Crofts 1966). In the study by Wheeler (1963), the normal oat sowing without nitrogen, outyielded pasture by a factor of 3.9 in comparison with a factor of 5 to 10 in New England. These results indicate that these High-vigour oats have an even greater potential in the north of NSW where growers are obliged to use the frost resistant varieties. Sual, a rust resistant line of Sydney University derived from Algerian, also broke these world yield records by a similar margin to that of P4315, but was significantly inferior to P4315 in the total yield of 2 pasture cuts. Grain recovery yields of Cassia and Blackbutt also broke the world oat yield record and Cooba, the check variety, was equal to the oat record. Competing crop trials The NSW Department of Agriculture began competing crop trials of oats, wheat, rye and barley at Glen Innes in 1921. By 1935 it was well established that oats was the most productive crop under grazing during the severe winters of the New England Tablelands together with recovery of hay and grain. By that time, mostly oat varieties were being tested for production of grazing material, hay and grain from plantings in February. The reason for the early sowings was in order to obtain four grazings. As a continuation of the 1921-1939 trials, oats were compared at regular intervals with the latest varieties of wheat, barley and rye and more recently with the newer crop triticale. The objective trials of wheat, oats and barley recorded in this chapter are quite typical of many such comparisons made over the last 40 years in NSW. Therefore production statistics are too subjective. To make more sense of production statistics, details like those of the Research Stations, District Agronomists and Cooperating Farmers should be collected and given top priority by the Bureau of Statistics Forsberg and Reeves (1995). In The Oat Crop: Production and Utilization. Edited by R.W. Welch, p.223., 1985). This section of this chapter is an attempt to do this. Pasture research scientists in Australia have had great success with the long-season variety from the High-vigour cross, Blackbutt. Muldoon in 1986, found that Blackbutt had a higher dry matter percentage in primary growth than Cooba and Abysinnian barley. He also found that Blackbutt had the highest cumulative regrowth of any cereal, well ahead of the barleys, which declined considerably for the third regrowth. Muldoon was comparing oats, barley, wheat, cereal rye and triticale under irrigation at Trangie, NSW in 1978 and 1980 and has made a great contribution to pasture and grazing research (Muldoon 1986). On the Southern Tablelands of NSW, Dann et al. (1983) found that Blackbutt oats yielded considerably more than Isis wheat in herbage availability, grazing days, liveweight gain and grain yield. They also found that severe grazing (down to 2 cm) never reduced the number of spikes or panicles per unit area and that cattle were not significantly superior to sheep for liveweight gain on grazing oats but were greatly superior for financial return because of the 100% higher price/kg live-weight obtained, in that year, for the cattle (Dann et al. 1983). Table 3.13 shows the continuing supremacy of Blackbutt oats for grazing and grain in the relatively dry, frosty winters of NSW, even over the most recently bred variety, Eurrabie, selected at Temora under the conventional system of breeding. Only the yields of the highest yielding winter wheat, Tennant, are included in Table 3.13. In addition to the superior grain recovery yields of Blackbutt, its superior grain feed quality, high-test weight and low husk percentage should also be considered by Southern Tableland farmers in choosing a winter cereal grazing crop.

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Carbeen was included in grazing and grain recovery trials in South Australia (Craig and Potter, 1983). The trial was sown on 29 May, 1980 on a solodised solonetz soil with 100 kg/ha of superphosphate. The area was evenly grazed by sheep (an excellent source of fertiliser and even grazing). Comparing 0, 1 and 2 grazings, the erect varieties yielded more grain after one grazing than after 0 or 2 (Table 3.11 and 3.12). The most prostrate variety, Carbeen, was the only variety to yield more grain after 2 grazings than after 0 or 1 grazing in this trial. The second grazing was on 5 September 1980, after a pasture cut to 2.5cm above ground level. This section of the trial was grazed by 100 sheep for 3 days down to a uniform height of 2.5cm above ground level. The prostrate variety, Carbeen, from the High-vigour cross, significantly outyielded all other varieties in grain recovery. The effect of a single grazing event led to higher grain protein in A. strigose compared to common oats species (Table 3.11). Competing crop trials conducted in New England, NSW, demonstrate the superiority of the High-vigour lines. At Glen Innes, the winter of 1955 was cold and cloudy after the first grazing. Although Winglen winter wheat was the least visibly affected by frost damage (scoring 1+ out of 6), in Table 3.10, it was significantly inferior in total grazing to all the oat and barley varieties. The winter wheats were set back by the cold weather and produced very little feed for the late August grazing. The oat results showed that this cereal was the most effective for winter grazing and grain recovery, while both oats and winter wheats were the best grain yielding cereals. In this trial there was no significant difference between Acacia and Winglen winter wheat. This flexibility or versatility of oats compared with other cereals was further highlighted at the New England Experiment Farm at Glen Innes in 2 trials in 1953 (a dry season) and in 1954 (a wet season) (Table 3.14). Oats yielded higher than winter wheat, spring wheat and barley in both seasons, with Acacia oats outyielding all varieties of wheat and barley in both seasons, and significantly so in the dry season. Barley and spring wheat responded the most to wet season-oats the least, with the exception of Acacia oats which significantly outyielded all varieties of wheat and barley in both seasons. The relatively new crop Triticale (variety Empat) was inferior in pasture yield to the High- vigour lines (Table 3.15) as were the wheats. Barley is a “quick early” lenient grazing cereal which does not recover well and is susceptible to frost and waterlogging at Glen Innes (Tables 3.3, 3.4, 3.10 and 3.14). Freebairn (1986) described a very large area of the Coonabarabran and Coolah Shires, 900,000 ha of agriculture, with acid soils, low in natural N and P. Before clearing, this country was covered with ironbark, bloodwood, pine, gum, apple box and a variety of undergrowth. Freebairn highlighted the value of lupins, triticales and rye. He found Carbeen (selected from the High-vigour cross at Glen Innes) to be superior to other oat varieties for tolerance to acid soils and aluminium toxicity.

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Table 3.11 Effect of a single grazing, grain recovery, total yield and grain protein (%) on a range of oat cultivarsa.

Cultivar Species Cut 1 (t/ha)b Grain (t/ha)c Total Yield (t/ha) Protein (%)d

Moore A.sativa x A.byzantina 0.207 3.615 3.822 12.2

Avon (101) A.sativa 0.224 3.508 3.732 12.0

Swan A.sativa x A.byzantina 0.230 3.070 3.300 12.2

West (213) A.sativa x A.byzantina 0.273 3.010 3.283 12.6

Coolabah (105) A.sativa 0.198 2.826 3.024 13.3

Cooba (22) A.sativa 0.143 2.542 2.685 15.3

Cassia (103) A.sativa 0.230 2.281 2.511 14.3

Carbeen (19) A.byzantina 0.166 2.341 2.507 14.2

Stout (212) A.sativa 0.326 1.967 2.293 15.8

Saia (60) A.strigosa 0.104 1.861 1.965 19.2

LSDe - 0.112 0.538 - -

a Trial conducted in South Australia in 1980 and reported by Craig and Potter (1983). All yields expressed in dry matter; b One pasture cut only was taken; c Grain yield after one grazing; d Protein content of grain harvested; e Least significant difference (t/ha).

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Table 3.12 Effect of two grazing cuts on grain recovery and pasture yields on a range of oat cultivarsa.

Cultivar Cut 1 and 2 (t/ha)b

Grain Yield (t/ha)c

Total Yield (t/ha)

Volumetric Weight (kg/hl)

Grain weight (g/1000 seed)

-2mm fraction(%)d Growth Habite

Carbeen (19) 1.391 3.227 4.618 44.0 34.8 9.9 3

Swan 1.477 2.195 3.672 49.0 45.0 4.2 7

Moore 1.252 2.321 3.573 47.1 42.0 5.4 8

Coolabah (105) 1.203 2.133 3.336 44.1 32.8 21.5 6

Cooba (22) 1.252 1.802 3.054 45.5 31.8 18.5 4

West (213) 1.317 1.534 2.851 47.8 36.3 10.0 8

Avon (101) 1.476 1.346 2.822 43.9 37.2 12.6 7

Cassia (103) 1.071 1.529 2.600 46.9 33.2 24.3 7

Stout (212) 1.315 0.850 2.165 46.0 35.8 9.5 7

Saia (60) 0.518 1.426 1.944 52.1 19.5 86.8 7

LSDe 0.445 0.538 - - - - -

a Trial of the same 10 varieties subjected to 2 grazings and grain recovery (Craig and Potter, 1983); b Sum of dry matter yields from 2 grazing cuts; c Dry matter grain recovery yield after 2 grazing cuts; d w/w (%) of grains passing through a 2mm screen; e Growth habit score on 1 to 10 scale where 1 = prostrate and 10 = erect; f Least significant difference.

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Table 3.13 Dry matter of pasture and grain recovery trial, Gunning, NSW (1999)a.

Cultivar Origin (oats) P cut 1 (t/ha)

P cut 2 (t/ha)

Grain recovery (t/ha)

Blackbutt (11) Glen Innes 3.49 1.46 3.70

Nile (209) Tasmania 4.00 1.33 3.10 Maiden (Triticale) 4.20 1.48 2.62 Eurabbie (Oats) Temora 4.17 1.37 1.80 Tennant (Wheat) 2.87 1.39 2.55 SDb 0.65 0.26 0.50 CV (%)c 10.72 12.65 14.51 Grazing date 11th June 20th August

a From Powell (2000). The above trial was sown on 1st April, 2000 and was harvested on 22nd December; b = significant difference; c = coefficient of variation.

Table 3.14 Grain yields from competing a cereal crop trial conducted in New Englanda.

Crop Variety Yield Dry Season (t/ha)b

Yield Wet Season (t/ha)c

Oats Acacia (1) 1.743 3.289

Oats Algerian (3) 1.617 2.237

Oats Fulghum (27) 0.983 -

Oats Lampton (56) - 2.930

Winter Wheat Winglen 1.268 2.111

Winter Wheat Cel. x Ten. D 1.268 2.804

Wheat Celebration 0.951 1.544

Wheat Lawrence 0.666 -

Barley Abyssinian 1.141 2.268

Barley Trabut 0.824 2.066

Barley Pryor 0.666 2.804 a Trials under dry (1953) and wet (1954) conditions; b Trial conducted in 1953 at New England Research Station at Glen Innes; significant difference = 0.254 t/ha (standard deviation=13.7%); no grazing; sowing date was 10 April 1953; c Trial conducted in 1954 at New England Research Station at Glen Innes; significant difference = 0.781 t/ha (standard deviation = 21.8%); no grazing; sowing date was 11 May 1954.

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Table 3.15 A continuous grazing (P) and grain recovery (pG) trial in Central NSW; F34 generation testing of High-vigour varities (1990) at Blayneya.

Cultivar (no.) P t/hab pG t/hac t/had

Carbeen (19) Blackbutt (11) Birch II (W) Empat (T) Cooba (22) WB135 (B) Birch 41 (W) Rosella (W) Osprey (W) Owlet (W) M5291 (W) Concort (R) SD CV (%)

2.32 2.16 1.63 1.48 2.55 2.29 1.85 1.91 1.68 1.58 1.08 0.93 0.46

14.6%

2.62 2.76 3.03 2.94 1.71 1.49 1.87 0.80 1.02 0.79 0.67

- 0.70 18%

4.94 4.92 4.66 4.42 4.26 3.78 3.72 2.71 2.70 2.37 1.75 0.93

- -

a Sowing date: 5.3.90; harvested: 7.1.91; growing season: 308 days; rainfall 820 mm. W = wheat; T = triticale; B = barley; R = ryegrass (measure of pasture growth). NSW Department of Agriculture trial conducted at Blayney (cool climate) was sown into excellent moisture and tilth at seeding rates of 70 kg wheat, 100 kg oats, 80 kg barley, 90 kg triticale, 25 kg/ha Concord ryegrass. Fertilizer was 62 kg/ha urea pre-sowing + 110 kg/ha DAP at sowing = 48 kg N + 22 kg P/ha. b Pasture cut taken on 9.5.90, then continuous hard grazing to 19.6.90; c Grain recovery since 19.6.90. No pasture cut taken on 19.6.90; d Total of b + c, not total biomass (Gammie, 1990).

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In the 1969 New England trials (Table 3.9), only Algerian, Coolabah and Fulghum were not significantly superior to winter wheat for 4 successive pasture cuts. P4315 and Abyssinian barley were the highest yielders for 2 grazings and grain recovery. The barley would not have withstood 4 grazings, as proven by other trials. In both trials, P4315 was the top yielder, followed by Blackbutt oats, its sister line from the High-vigour cross of 1957. In the warmer climates of the Western Plains of Central NSW, all oats were significantly superior to winter wheats for first, second and total pasture cuts in a dry winter, 1964 at Condobolin (Table 3.16). For grain recovery, the Temora line, VRBurke (W4464) gave the highest yield, which was significantly superior to that of Winglen winter wheat but not Windebri. In the grain only section of this trial, W4464 and Avon oats were significantly superior to both winter wheats. Bundy was not inferior to the best winter wheat yields and was significantly superior in grain recovery, and in the grain only trial, to Belar, which it was intended to replace as an early mid-season oat with greater frost resistance and equal milling quality (Guerin, 1965). The yellowish brown colour of the grain, however, was not as acceptable for milling as the creamy brown grain of Belar. The better grazing yields of Cooba and its improved milling traits explain why Cooba replaced both Belar and Bundy. Summer rainfall climates adversely influence grain colour in most seasons. Millers therefore buy brighter coloured oat grain grown in the drier summer climates. Carbeen was tested by Craig and Potter (1983) in South Australia. There it was the only variety not to suffer reduction in grain yields when grazed twice. Under severe winter conditions, however (Tables 3.11 and 3.12), Blackbutt proved to be the most productive oat for total yields. This higher productivity of Blackbutt amounts to an extra tonne when sowing is in early March rather than in late March. Associated with the extra yield is an increase in the protein content. The dry matter pasture yields and the grain recovery yields may be taken to be equal in energy value but the herbage dry matter would have a protein equivalent of 15, or twice that of oat grain, according to Robinson (1949). This is important for lactating cows or ewes, and for growing high quality fat lambs. The management of oat pasture requires knowledge, skill and supplementation with native pasture or dry feed at night time, for sound animal husbandry. The new High-vigour oats are greatly superior to wheat, rye and barley for biomass and protein yields. Winter wheat is too slow growing, as found also by Dann et al. (1977) at Canberra. Barley and Winter Rye have been excluded from the analysis of means (Table 3.9) of the early March sowing. The former was too frost susceptible and the latter too low in productivity for this environment. They are not suited for the heavy grazing required for high protein yields.

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Table 3.16 Effect of two grazing cuts and grain recovery (Site 1) and grain only (Site 2) on a range of oat cultivarsa.

Site 1 (t/ha) Variety

Cut 1 Cut 2 Total Cut Grainb

Site 2 Grain Only (t/ha)

Avon (101) 0.73 0.88 1.59 1.38 3.32

Belar (9) 0.54 0.74 1.28 0.97 2.20

Bundy (15) 0.44 0.68 1.12 1.37 2.55

Cooba (22) 0.46 0.87 1.33 1.41 2.33

Kent (52) 0.84 0.85 1.69 1.10 2.35

VR Burkec 0.46 0.76 1.22 1.54 3.29

Windebrid 0.30 0.44 0.74 1.47 2.30

Winglend 0.39 0.48 0.87 1.23 1.95

SD 0.04 0.04 0.05 0.18 0.30

CV (%) 16.2 15.1 10.1 11.4 12.5

Cutting datee 105 160 - - -

a Site 1-sowing date: 24 March 1964; 4 replications; conducted at Condobolin, NSW; Site 2-sowing date: 5 May 1964; 4 replications; conducted on Mr.C.Grady’s farm at Trundle, NSW for grain only. All yields are for dry matter; b Grain yield after 2 grazings; c Early maturing line (W4464) bred at Temora, NSW; d Winter wheat varieties; e Days after sowing.

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Figure 3.7 (Top) The Author (on left) shows greater damage to Algerian from a combination of frost and grazing pressure than that to Klein 69B (right) the Argentine oat, which showed excellent frost resistance and grazing recovery almost equal to Blackbutt; (Bottom) The Author (on right) shows poorer performance of Algerian compared to High-vigour line P4314. Further images of the grazed plots at Hawkesbury Agricultural College trials in Richmond NSW in 1966, are presented in Appendix B.

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Figure 3.8 A comparison of the five selections from the High-vigour cross for total biomass yield (P + pH) with conventionally bred cultivars at Hawkesbury Agricultural College, Richmond, NSW (1966).

0

5

10

15

20

25

Bla

ckbu

tt

P431

5

856

871\

1

871

Coo

ba

Kle

in 6

9B

Fulg

hum

Coo

laba

h

Fulm

ark

Alg

eria

n

Yie

ld (t

/ha)

Total pasture yield (P) Hay recovery (pH) (P+pH)

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Figure 3.9 A comparison of a standard cultivar, Algerian, with five selections from the High-vigour cross, with 5 separate pasture cuts at Hawkesbury Agricultural College, Richmond, NSW (1966). The extent of the grazing is shown in individual plots within the trial presented in Appendix B.

0

0.5

1

1.5

2

2.5

1 2 3 4 5Number of cuts

Yie

ld (t

/ha)

P4315 856871 871/1Algerian Blackbutt

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- 99 -

Figure 3.10 The Author at Temora Agricultural Research Station taking notes near seed increase blocks. Selecting hardy, productive and rust resistant dual-purpose oats by wide spacing of plants in the Author’s Isolection breeding system, produced P4315 (Top), Blackbutt (Centre) both from the same High-vigour cross; (Bottom) Mugga, also bred by the Author, is the hardiest of the oats tested in Glen Innes NSW, equivalent hardiness to winter wheat. Mugga was selected from VRBop x Belar.

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Recommendations for extension With the much higher and more stable yields of the new bulk varieties (Tables 3.3, 3.4 and 3.5), four years of testing instead of the usual 7 years would therefore be sufficient for cultivar release. This would be the case especially if the number of testing sites could be increased. Instead of the usual 15 years under the pedigree system, which could be run concurrently, the time from cross to farmer can be halved to 6 years for F3 bulks and 8 years for F4 directed bulks. New possibilities are opened up by this rapid breeding technique for producing dual-purpose oat cultivars. For NSW oat research programs to be attractive to funding, it will probably need to replace the popular variety Cooba with a variety that yields at least 20% higher. The F3 bulk, P4315, tested from 1960 to 1976, was at times up to 100% higher in yield than Cooba but lacked milling suitability. It may soon be that this stipulation of millibility will be waived by the NSW Department of Agriculture, allowing such potential cultivars to be released to farmers. As well as P4315, there is a millable F4 directed bulk, P4318, which is also tolerant to barley yellow dwarf virus (BYDV), as is P4315. There could be little, if any, advantage in releasing a high yielding F3 bulk to a farmer who is interested in growing oats for grain only. On the other hand, farmers who are in need of supplementing their winter pastures (Crofts 1966), and who practise optimum soil and crop management (Guerin 1961) stand to gain significantly. Supporting the latter type of farmer would increase Australia's relatively low average oat yields (approximately 1.4 tonnes/ha), bringing them closer to the yields reported in this chapter. The model farmer will extend both seed and cultural knowledge to make long-term profits. A further recommendation based on the results of the NSW Department of Agriculture oat breeding and testing program, i.e. the results of which are presented in this chapter, is that F3 bulks and F4 directed bulks should be maintained indefinitely, not alone as gene banks similar to Suneson’s composites (Allard and Hansche 1964), but also to monitor yield changes, over time, relative to all check varieties, including Algerian, Fulghum and Cooba.

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CONCLUSIONS This study highlights several important considerations for plant breeding. Firstly, The important lesson from Isolection breeding is to aim for close (not distant) crossing within the same ecospecies or morphological type, like winter oat x winter oat or spring type x spring type, rather than winter x spring type crosses. This concentrates the multiple genes needed for each type. Wide crossing within the hexaploid oats (or wheat) requires backcrossing in order to regain the ecotype, while the full yield potential being sought is not regained. As well as concentrating together the multigenes required for a winter oat (or a spring oat), the Isolection system also enables the rapid breeding of new varieties. Thus, from the High-vigour cross of 1957, the Author was able to recommend the release of both P4319 and P4315 as early as 1966. However, only P4319 was released and it was subsequently named Blackbutt in 1974. Secondly, plant breeders should select as many plants as possible in the F2 to retain as much hybridity as possible, leading to homeostasis and versatility, as compared with F6 selected inbreeds, which show much more environmental variation. This is important in the case of oats, where cross-pollination is only 0.5% as compared with 5% in wheat and triticale. Nevertheless, James Carroll, the Author’s predecessor, maintained that fixity is never finalised and this is certainly the case with landraces like Algerian and Fulghum (as discussed in Chapter Two). Early, not later, generation selection is suggested by the Mendelian diagram for a trihybrid or three factors. Thirdly, the results of this study indicate that heavy grazing, that is 4 or more pasture cuts, are essential to assess true dual-purpose capacity of oat varieties. Finally, the results of NSW-wide biometrically designed and analysed yield trials have strongly supported the principle of the Isolection breeding system, which may be stated as follows: The continuing ability to observe and select new varieties of cultivated crops depends on isolating single (individual) plants with optimum space and nutrient status in the early generations after the cross and situated in the locality for which the crop is being bred. While the principle of Isolection is essential to pin-point and select new varieties, yield per unit area rather than yield per single plant is the ultimate goal and this requires statistically sound replicated yield trials in order to determine the highest yielding commercial variety of oats for the farmer. The best dual-purpose oats now derive from the High-vigour cross which has produced Blackbutt and Carbeen. The results of this oat breeding and testing has proved the advanced state of competence in oat breeding, testing and production that has been achieved in Australia.

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REFERENCES Archer, K.A. and Swain, F.G. 1977. Evaluation of oat forage for finishing prime lambs on the Northern Tablelands, NSW. Australian Journal of Experimental Agriculture and Animal Husbandry 17: 385-392. Auckland, A.K. 1967. Soybeans in Tanzania Part 2. Seasonal variations and homeostasis in soybeans. Journal of Agricultural Science (Cambridge) 69: 455-464. Craig, A.D. and T.D. Potter. 1983. The effect of grazing on the grain yields of ten oat cultivars in the south east of South Australia. Agricultural Record 10, 15: 4-7. Crofts, F.C. 1966. More winter feed and drought reserves from high density, nitrogen fertilized oats. Agricultural Gazette NSW 77, 5: 258-262. Dann, P.R., A. Axelsen, and C.B.M. Edwards. 1977. The grain yield of winter-grazed crops. Australian Journal of Experimental Agriculture and Animal Husbandry 17: 452-461. Dann, P.R., A. Axelsen, B. S. Dear, E.R.Williams and C.B.H. Edwards. 1983. Australian Journal of Experimental Agriculture and Animal Husbandry 23: 154-161. Evans, L.T. 1996. Crop Evolution, Adaptation and Yield. Cambridge University Press, pp. 288-9. Fagan, T. W. and W. E. J. Milton (1931). The chemical composition of eleven species and strains of grasses at different stages of maturity. Welsh Journal of Agriculture 7: 229-246. Fageria, N.K. 1992. Maximizing Crop Yields, Marcel Dekker, Inc. Falconer, D. 1952. The problem of environment and selection. American Naturalist 86: 293-298. Falconer, D.S. 1975. Introduction to Quantitative Genetics. Longman. Falconer, D.S. and T.F.C. Mackay. 1996. Introduction to Quantitative Genetics. Longman, Melbourne. Fitzsimmons, R.W. 1978. Developments in the oat growing industry. Agricultural Gazette NSW 89: 35-37. Forsberg, R.A and D.L. Reeves. 1995. In The Oat Crop: Production and Utilization. Edited by R.W. Welch, p.223. Freebairn, R.D. 1986. Rotations and Soil Acidity, Agriculture NSW, The Regional Institute. Frey, K.J. 1964. Adaptation Reaction of Oat Strains Selected under Stress and Non-Stress Environmental Conditions, Crop Science 4: 55-58. Gotoh, K and S. Osanai. 1959. Efficiency of selection for yield under different fertilizer levels in a wheat cross. Japanese Journal of Breeding 9: 101-106. Guerin, P.M. 1965. Bundy - The breeder's report: A new midseason oat variety for Northern NSW. Agricultural Gazette NSW 76: 667-669.

Guerin, P.M. 1966. Mugga - The breeder's report: A new winter-hardy oat for the Northern Tablelands. Agricultural Gazette NSW 77: 675-678. Guerin, P.M. and T.F. Guerin. 1992. A rapid low-technology method of breeding high-yielding oats with dual-purpose characteristics. In Fourth International Oat Conference in Adelaide. Edited by A.R. Barr, R.G. McLean, J.D. Oates, G. Roberts, G. Rose, K. Saint, and S. Tasker, pp.191 – 195.

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Guerin, P.M. 2003. Scientific Laws of Creation: Breeding High-Yielding Crops. Self-published. Hodgson, J., R.D. Baker, A. Davies, A.S. Laidshaw, and J.D. Leaver. 1981. Sward Measurement Handbook. Maidenhead: The British Grasslands Society. Lovett, J.V. and E.M. Matheson. 1974. Cereals for winter grazing on the Northern Tablelands of NSW. Australian Journal of Experimental Agriculture and Animal Husbandry 14: 790-795. McRae, F.J., D. McCaffery and P. Matthews. 2005. Winter Crop Variety Sowing Guide 2005. Agdex 110/10, NSW Agriculture. Muldoon, D.K. 1986. Dry matter accumulation and changes in forage quality during primary growth and three regrowths of irrigated winter cereals. Australian Journal of Experimental Agriculture 26: 87-98. Pearce, G.R., R.G. Simpson, R. Ballard, and D. Stephenson. 1987. Feeding value of dead pature grasses. In Temperate Pastures, Edited by J.L. Wheeler, C.J. Pearson, and G.E. Robards, pp. 423-431. Sydney: Australian Wool Corporation and CSIRO. Powell, C. NSW Agriculture, 2000: Winter Crop Variety Experiments for 1999. Roberts, G. 1989a. Avena sativa cv.Yarran. Australian Journal of Experimental Agriculture 29, 1: 144-145. Roberts, G. 1989b. Avena sativa cv. Hakea. Australian Journal of Experimental Agriculture 29, 1: 146-147. Robinson, D.H. 1949. Fream's Elements of Agriculture. London: John Murray.

Spurway, R.A. 1975. Oats stand out as a winter forage crop for the Northern Tablelands. Agricultural Gazette NSW 86: 24-26. Wheeler, J.L. 1962. Experimentation in grazing management. Herbage Abstracts 32: 1-7. Wheeler, J.L. 1963. Winter forage crops in the New England Tablelands. Australian Journal of Experimental Agriculture 8: 62-68. Wilson, B. 1968. (Editor) Pasture Improvement in Australia. Murray, Sydney, p.176.

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CHAPTER FOUR

BREEDING OATS FOR IRRIGATION IN AUSTRALIA

Glen Innes, on the New England Tablelands in NSW, has proven to be the best centre for breeding oats for the heavy soils of the Riverina at Leeton, southwestern NSW. Plant selections made on the black self-mulching soils of the Glen Innes Research Station of northern NSW have resulted in the varieties Acacia, Bundy, and Mugga; all now replaced by Blackbutt. Both areas require resistance or tolerance to stem rust, water logging, red-legged earth mites, BYDV, lodging, shattering and second growth. Although frost damage is less of a problem in the irrigation areas than on the northern tablelands of NSW, the frost resistant bulks from the cross F.Ga x VRAF.VRSF demonstrated good tolerance to water logging on heavy soils. Blackbutt also excelled as both a dual-purpose and a grain only variety, therefore a triple-purpose variety, and has been recommended for both northern and southern irrigation areas. INTRODUCTION In order to succeed in plant breeding for irrigation, it is necessary to study yields both in dryland and in irrigation trials, so that we can learn more about the complexities of yield itself, the different environments and requirements of the farmer. Non-irrigated as well as irrigated trials are therefore both considered relevant in this chapter. Alternative crop studies are also necessary. Oats are regarded as needing more water to produce a unit of dry matter than any other small grain cereal except rice (Coffman 1961). Rice is the most important irrigated crop in the Riverine Plain of NSW. Wheat, however, is also extensively grown under irrigation but is very prone to crop failure if the varieity is susceptible to rust diseases (Leigh and Noble, 1972). Oats is less affected by rust diseases and soil-borne pathogens than wheat and therefore, often resulting in higher yields under irrigation. Oats can utilise more water than any other cereal except rice (Coffman, 1961) and is less prone to crop failure than wheat in the event of serious rust disease and the soil-borne pathogens which affect wheat and barley (Leigh and Noble, 1972). This is especially important in countries like Australia and China, where reliable sources of irrigation water are available and sometimes under-utilised, because of a need to apply a soil-conditioner like gypsum (Davidson and Quirk, 1961). Yields of oats in NSW are compared with other cereals in Table 4.1.These statistics suggest that oats respond well under irrigation, however, this has only been the case since the 1980s. The results of trials presented in this chapter show the response of a range of oat cultivars, inlcuing the High-vigour varieties described in Chapter Three, as well as other cereals in the Riverine Plains of NSW.

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Table 4.1 NSW cereal crop yields under dryland and irrigation (t/ha)a.

Total Irrigation Onlyb Years

Wheat Oats Barley Wheat Oats Barley

1930-1939 0.93 0.72 0.98 0.82 0.63 0.84

1970-1979 1.43 1.06 1.24 1.43 0.92 1.18

1982-1990 1.60 1.24 1.49 1.52 1.33 1.27

a Data from Fitzsimmons (1990). b This data was taken from Leeton, NSW.

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AUSTRALIAN RESEARCH ON IRRIGATED OATS AND FORAGE PRODUCTION Australian research over the past 20 years has demonstrated the significant role oats have to play in irrigated farming and forage production. Dann et al. (1983) have shown that Winglen, Windebri and Isis winter wheats produce significantly less early fodder than oats. Blackbutt oats out-yielded Isis wheat in grain yield by 125% in 1976 but the seed shattered badly in 1977. Severe grazing (down to 2 cm) never reduced the number of spikes or panicles per unit area of the Blackbutt variety. One disadvantage of oats in general is a tendency to shatter when ripe. To reduce shattering loss, oats can be windrowed up to 14 days before it can be direct headed. Duncan (1983) found that triticale and rye both run to head early in the spring, producing a spiky head with the same disadvantages as barley. He also found that Blackbutt and Acacia oats (both bred at Glen Innes) were more resistant to waterlogging than Cooba, which was also less tolerant to heavy frosts than Blackbutt and Carbeen. Duncan found that oats were the most popular fodder crop for filling the winter feed gap and also served as a useful “clean-up” crop in weedy areas, after pasture improvement, because pastures are not competitive enough to keep out thistles, barley grass and burrs. Muldoon (1986) has made a deep study of irrigated winter cereals and compared oats, barley, wheat, cereal rye and triticale under irrigation at Trangie in 1978 and 1980. Dry matter accumulation was described by mathematical equations which allowed cultivars to be compared under different cutting regimes. Muldoon (1986) also allowed dry matter and digestible dry matter yields to be predicted for irrigated cereals in western NSW. He showed that early maturing barley and triticale cultivars had lower digestibilities than oats. Oats and wheat had similar digestibilities and these began to decrease rapidly 40-50 days before head emergence (mid-August). Blackbutt oats had the highest cumulative regrowth of any winter cereal. Regular cutting (a total of 4 cuts) maintained the nitrogen content and digestibility of all cultivars above 2.7% and 72% respectively (Muldoon 1986). Simmons (1987) noted that varieties with more prostrate growth withstand close grazing, trampling and heavy frosts better than more upright growers. Blackbutt was outstanding in this respect, although not as frost hardy as some winter wheats. Mugga oats also had greater frost resistance than Blackbutt but was not as productive in grazing as Blackbutt. Powell (2000) found that Blackbutt oats sown as late as 1st April 1999 and harvested on 22nd December, yielded significantly more grain after 2 grazings than Nile and Eurabbie oats, Maiden Triticale, Tennant and all other wheat varieties. This followed grazing cuts on 11th June and 20th August, giving no significant differences, in the relatively dry, frosty winter at Gunning, on the Southern Tablelands of NSW. Eurabbie is a semi-dwarf oat, obviously not satisfactory for grain recovery yields after severe grazing. Hubbell et al. (2000) in America found similar weight gains in 225 kg steers grazing winter wheat (1.25 kg/day), oat forage (1.34 kg/day), rye (1.25 kg/day) and perennial ryegrass (1.18 kg /day). McRae (2003) found similar results for Australia: Quality tests on the forage value of oats, wheat, barley, cereal rye and triticale, when grown under similar conditions, show no significant differences in protein, energy and digestibility. For overall forage production, however, he found that oats produce more than wheat, barley, cereal rye or triticale.

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Dove (2004) found great variability in observed liveweight gains of animals grazing winter wheat, suggesting that more attention should be paid to cereal stage of growth and more accurate livestock comparisons and needs. TESTING OF HIGH-VIGOUR VARIETIES The NSW Department of Agriculture has conducted dual-purpose and grain only oat trials for many years in the Riverine Plain of NSW, a major irrigation area. The early varieties, P4315, Avon, Cooba and Coolabah were compared with the late varieties, Blackbutt, Acacia, Algerian, Klein 69B and Mugga over the period of 1963 – 1973 (Table 4.2). The varieties were kept in 2 separate groups, for ease of harvesting. The mid-season variety Bundy was common to both early and late trials. Due to the frequent tendency of Avon to make second growth and to delay harvest, early varieties sometimes were over-ripe and suffered some shattering losses as a result. Grazing yields are not given in these trials as they are usually too lenient in character to assess grazing potential under irrigation. The High-vigour bulk, P4315, topped the yields both in the grain only and in the grain recovery after grazing trials (Table 4.2). Similarly, Table 4.3 compares only grain recovery (grazing yields excluded) with grain yields on dryland in the Riverina. P4315 tops both sections without irrigation. This line therefore demonstrates a triple-purpose capacity, that is, high dual-purpose and high grain only yields. Blackbutt, under irrigation showed the same capacity but the earlier compounded bulk, P4315, was more adaptable, being triple-purpose under both irrigation and dryland conditions (Tables 4.2 and 4.3). Irrigation does not always give the highest yields unless the season is dry and drainage is good. The research plots at Cowra (results presented in Table 3.7, Chapter Three) were not irrigated and yet out-yielded the irrigated trial at Coleambally, NSW (Table 4.4). In this trial at Colleambally in 1985, Blackbutt demonstrated its high grain yield potential. The comparison between the irrigated site at Coleambally with the dryland trial at Adelong, showed the superior grain recovery yield of Blackbutt over those of wheat, barley and rye. The very cool climate of Adelong NSW, the Tasmanian oat variety, Nile, was significantly the highest yielder of all the crops and varieties tested. More recently, Sydney University's rust resistant Algerian-derived cultivar, Sual, has given high yields under irrigation although the straw is tall and weak, which requires this variety to be grazed (data not presented). This is impossible if water-logging occurs. The semi-dwarf variety, Eurabbie, released in 1998, is too short after heavy, late grazing. This results in harvesting difficulties. Eurabbie is also susceptible to rusts and BYDV, making it unsuitable for irrigation (McRae et al. 2004). Avon, a high yielding grain oat from West Australia, was found to be unsuitable for growing in irrigation areas because of its tendency to remain green for too long, holding back harvest operations. Most virgin soils of the Riverine Plain have extremely low infiltration rates (Leigh and Noble, 1972). Gypsum has made possible successful pasture establishment (Davidson and Quirk, 1961). On these soils, pasture establishment and rotation provide a sound basis for arable cropping. Due to the original nature of these soils, they still require oat varieties resistant to occasional waterlogging. For this reason, selected varieties at Glen Innes including Bundy,

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Acacia, Mugga (Guerin, 1966) and the High-vigour bulks have demonstrated their tolerance to these conditions, as have Klein 69B, Coolabah and Cassia (Fitzsimmons, 1986). The latter three should not be considered for grazing here in the Riverine Plain, because of poor quality grains per se. Resistance to summer storms at Glen Innes is also useful for selecting varieties for growing under irrigation. In the grain only trial, the dual-purpose cultivar bred in northern NSW, Blackbutt, yielded over 5 t/ha at Coleambally. Blackbutt outyielded all the specialised grain only oat varieties from West Australia, South Australia and Victoria (Figure 4.1). This triple purpose capacity was displayed by Blackbutt, due to its strong straw or general resistance to lodging, derived from the wide diameter culms of Garry (one of its parents). The trial at Colleambally included the dwarf oat Echidna, which appeared to lack drought resistance in the dryland trial site at Adelong, NSW (notwithstanding the cool climate). Coleambally and Adelong yields are compared in Figure 4.1. The Glen Innes bred cultivars Bundy, Mugga, Acacia, Carbeen and Blackbutt have been high-yielding under irrigation. This appears to be associated with their resistance to water-logging at Glen Innes, where drainage is sometimes impeded on the black basaltic soils, otherwise soils of high fertility, especially in dry seasons. The strong straw of Blackbutt as well as its moderate resistance to BYDV and some strains of stem rust made it suitable for irrigation (Verrell and Gammie, 1992). Oats in general do not have the same disease problems as other cereals. The most serious diseases of oats are leaf (crown) rust, bacterial leaf stripe, BYDV and the head smuts, to the last three of which Blackbutt is tolerant or resistant to these final three diseases. Reconmmendations for oat breeding programs Oat breeding for irrigation should consider the following. Firstly, resistance to waterlogging especially in heavy soils. Acacia x Lampton lines bred by Carroll at Glen Innes were outstanding in this respect in the 1960s. Secondly, the strong straw of the High-vigour lines from F.Ga x VRAF.VRSF, namely Blackbutt and P4315, seems to derive from the tall strong-strawed variety, Garry (Ga). The shorter culms of P4315, for instance, had thicker stronger walls than those of Cooba, a variety without Garry parentage. It is for this reason that Cooba is not suitable for grain only, especially under irrigation. Finally, a separate point is the harvest index, or grain-hay ratios. Yield improvements to Ballidu, Orient and Avon have been attributed solely to an increase in harvest index (Sims, 1963). Results reported at the Adelaide conference (Guerin and Guerin, 1992) suggest that 1309 has a higher harvest index than Cooba with a hay yield equal to that of Cooba. Both Cooba and 1309, sister lines of the same cross, VRAF.VRSF, have high quality grain and good milling characteristics, therefore 1309, the parent of the High-vigour, is a better choice for crossing, or black-crossing, than Cooba, which is moreover lower in frost resistance. The grain/hay ratios reported (Guerin and Guerin, 1992) demonstrates that the High-vigour line, 856 G59 (P4314) has the highest harvest index recorded and yet is significantly higher in hay yields to both Cooba and 1309. This high harvest index appears to be derived from Fulghum.

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Table 4.2 Comparisons of early and late maturing cultivars under Irrigation in the Riverina for a 10 year period for grain only (G), and grain recovery (pG) (1963-1973).

Cultivars (Inventory Number)

G (mean of 7 trials) (t/ha)

pG (mean of 4 trials) (t/ha)

P4315 (38)a

Coolabah (105)a

Cooba (22)a

Bundy (15)a Avon (101)a Blackbutt (11) Bundy (15) Mugga (58) Coolabah (105) Cooba (22) Klein 69B (53) Acacia (1) Algerian (70)

3.76 3.61 3.42 3.36 3.30

4.36b 4.00b 3.79b 3.57b 3.23b

- - -

2.88 2.40 2.06 2.54

-

3.26c 2.75c 2.22c

- -

2.21c 2.06c 2.03c

Trials were conducted in the Murrumbidgee Irrigation Area, NSW, under irrigation from 1963-73; a early maturing cultivars; b late maturing cultivars, 2 trials; c late maturing cultivars, 4 trials.

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Table 4.3 Comparisons of grain only (G) yields and grain recovery (pG) in the Dryland Riverina for a 10 year period (1963-1973).

Cultivars (Inventory Number)

G (mean of 9 trials) (t/ha)

pG (mean of 4 trials) (t/ha)

P4315 (38) Cooba (22) Avon (101) Coolabah (105) Bundy (15)a

2.03 1.90 1.88 1.80 1.71

1.59 1.47 1.55 1.32 1.38

a The low yields of Bundy (Guerin, 1965), a drought tolerant variety, suggests an abundance of soil moisture in these trials, with P4315 exhibiting a triple-purpose capacity.

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Table 4.4 Year 24 of testing Blackbutt oats under irrigation versus dryland (1985): F29 generation trial. Cultivar (Inventory Number)

Irrigation sitea Gc (t/ha)

Cool dryland siteb pGd (t/ha)

Blackbutt (47) Cassia (103) Hakea (109) Echidna 206) Dolphin (205) Bundalong (204) Barmah (202) Nile (209) Carbeen (41) Cooba (22) Yarran (114) Coolabah (105) Mortlock (208) Bulban (203) West (214) Rysun, rye Malebo, barley Quarrion, wheat Rosella, wheat Osprey, wheat Forrest, barley SD CV (%) Date of sowing Date of harvest

5.29 4.89 4.78 4.76 4.63 4.36 4.25 4.05 3.75 3.61 3.02 2.89 2.89 2.71 2.62

- - - - - -

n.a. -

11.4.85 18.12.85

2.40 - -

1.58 - - -

2.85 2.31 2.01 2.12 1.90 2.00

- -

1.94 1.94 1.77 1.66 1.35 1.22 0.44 13.6

26.3.85 21.12.85

a Coleambally NSW trial, lodging prevalent; b Adelong NSW; c Grain only trial; d Grain recovery after grazing, which was not measured by a pasture cut; n.a. = not analysed.

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Figure 4.1 A diagrammatic comparison of eight cultivars under irrigation (grain only) and cool dryland (grain recovery) in t/ha (Colleambally, NSW in 1985).

0

1

2

3

4

5

6

Blackbutt Echidna Nile Carbeen Cooba Yarran Coolabah Mortlock

Variety

Yie

ld

Irrigation Site - Grain Only (t/ha) Cool Dryland Site - Grain Recovery (t/ha)

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CONCLUSIONS Glen Innes, on the New England Tablelands in NSW, has proven to be the best centre for breeding oats for the heavy soils of the Riverina in southwestern NSW. Plant selections made on the black self-mulching soils of the Glen Innes Research Station of northern NSW have resulted in the varieties Acacia, Bundy, and Mugga; all now replaced by Blackbutt. Both the Riverina and northen NSW require resistance or tolerance to stem rust, water logging, red-legged earth mites, BYDV, lodging, shattering and second growth. Although frost damage is less of a problem in the irrigation areas than on the northern tablelands of NSW, the frost resistant bulks from the cross F.Ga x VRAF.VRSF demonstrated good tolerance to water logging on heavy soils. Blackbutt also excelled as both a dual-purpose and a grain only variety, therefore a triple-purpose variety, and has been recommended for both northern and southern irrigation areas. In summary, the key characteristics required for breeding a successful oat variety for irrigation, in addition to stem rust, BYDV, and insect resistance, are (A) resistance to water logging, (B) strong straw and (C) a high harvest index.

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REFERENCES Coffman, F.A. 1961. Oats and Oat Improvement. American Society of Agronomy.

Dann, P.R., A. Axelsen, B. S. Dear, E.R. Williams and C.B.H. Edwards. 1983. Australian Journal of Experimental Agriculture and Animal Husbandry 23: 154-161.

Davidson, J.L. and J.P. Quirk. 1961. The influence of dissolved gypsum on pasture establishment on irrigated sodic clays. Australian Journal of Agricultural Research 12: 100-110.

Dove, H. 2004. Grazing grains—Cereals in grazing systems. CSIRO, Canberra, ACT.

Duncan, M. 1983. Winter fodder crops: northern tablelands. Agdex 120/20, Agriculture NSW.

Fitzsimmons, R. 1986. Winter cereal variety performance 1985-6 Season. Agdex 110/34, Agriculture NSW.

Fitzsimmons, R.W. 1990. Winter cereal production statistics. Australian Institute of Agricultural Science. Occasional Publication No. 48.

Guerin, P.M. 1966. Mugga - The breeder’s report. Agricultural Gazette NSW 77: 675-678.

Guerin, P.M. and T.F. Guerin. 1992. Breeding oats for irrigation in Australia. In Fourth International Oat Conference, Adelaide, Edited by A.R. Barr, R.G. McLean, J.D. Oates, G. Roberts, G. Rose, K. Saint, and S. Tasker, pp. 187 – 190.

Hubbell, D.S., Daniels, L.B., Harrison, K.F., Johnson, Z.B., Brown, A.H., Kegley, E.B., Coblentz, W.K. and Coffey, K.P. 2000. Arkansas Animal Science Department Report, pp.70-71.

Leigh, J.H. and J.C. Noble. 1972. Riverine Plain of NSW, its Pastoral and Irrigation development. CSIRO Division of Plant Industry, pp 1-63.

McRae, F.J. 2003. Crop agronomy and grazing management of winter cereals. GRDC Update.

McRae, F.J., D.W. McCaffery and D.J. Carpenter. 2004. Winter Crop Variety Sowing Guide, NSW Agriculture.

Muldoon, D.K. 1986. Dry matter accumulation and changes in forage quality during primary growth and three regrowths of irrigated winter cereals. Australian Journal of Experimental Agriculture 26, 1: 87-98

Simmons, K. 1987. Oats. Agfact P3.2.2. Wagga Wagga, NSW Agriculture.

Sims, H.J. 1963. Changes in the hay production and the harvest index of Australian oat varieties. Australian Journal of Experimental Agriculture and Animal Husbandry 3, 10: 198-202.

Verrell, A.G. and R.L. Gammie. 1992. Winter Cereal Variety Sowing Guide. NSW Agriculture.

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CHAPTER FIVE

THE INFLUENCE OF ENVIRONMENT

ON OAT GRAIN QUALITY

Oat grain quality (grain weights per 1000 seeds and groat percentages) was found to be an effective measure of the environmental stress imposed on an oat variety at a particular geographical and climatic centre. The results of various oat trials conducted across NSW show that the environment has an effect on the maturation and filling of the oat grain. The results compiled by the Author suggest that northern NSW (i.e. the summer rainfall zone) could be further sub-divided into 5 climatic regions, from east to west, for the purpose of recommending oat varieties. Glen Innes, at an elevation of 1,128m and latitude 29° 42” S. on the New England Tablelands, proved to be the ideal climate for developing high groat percentage and large grain size. A sixth climatic zone, located in Leeton, NSW (uniform rainfall zone), at elevation at 152m and latitude 34° 33” S., was the second most favourable centre, but required irrigation for full grain development. INTRODUCTION The environment in which oat varieties are bred and tested can have a significant effect on the final grain quality, in terms of weight and groat percentage. Both of these oat grain parameters are critical in determining how the harvested grain crop will be utilised. High oat grain quality for the farmer means a crop variety that yields high volumetric weight and high groat percentage. The 1000-grain weight, or seed size of an oat variety, can be low if the yield per ha is high. In general, oat cultivars belonging to the Avena sativa group, with high protein percentage, have been lower in grain yield than those with medium to low protein (Frey, 1992). Although Frey found no correlation between grain yield and grain protein concentration in some lines from a cross with A. sterilis, which has high-protein genes. Milling suitability is a different requirement, often in conflict with the grower’s needs, although they may not be aware or informed of this. Larger grains, like those of Belar (Guerin, 1965), a low yielding oat, are favoured by millers. So is Cooba (Mengersen, 1963), which is a high yielder in southern areas, but a lower yielder in the northern, rust and frost liable areas of NSW. The fact that millers pay more for certain varieties may sway the farmer into growing low yielding or unadaptable varieties. The registered seed grower of such varieties, even if they are better informed, are unlikely to advise against the variety they are selling if it is unsuited to the farmers' needs. Farmers thus get locked into growing a variety often for a small temporary cash advantage, ignoring potential long-term profits for the farm, where value is added to the grain in terms of wool, lambs and beef. TESTING OF HIGH-VIGOUR VARIETIES Testing environments Trials were conducted at several locations in NSW representing 6 different climates and environments. Northern NSW (i.e. the summer rainfall zone) was further sub-divided into 5

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climatic zones, from Glen Innes in the east to Tamworth, Gunnedah, Curlewis, Narrabri in the west. A sixth climatic zone, located in Leeton, NSW (uniform rainfall zone), at elevation at 152m and latitude 34° 33” S., was also used as a trial site. Leeton is an irrigated region in the southern rainfall area of the Riverina of NSW. Methods used Grains harvested from the trials were assessed for hectolitre weight (i.e. weight per 1000 grains) and for groat percentage. About 40 distinct oat varieties and selections were assessed for grain quality over a 5 year period. Trial results Testing in 6 different climatic zones demonstrated that 1000-grain weight and groat percentage were varietal characteristics. There were, however, separate components of quality which develop independently, depending on the location the crop is grown, and the season. Glen Innes, at an elevation of 1,128m and latitude 29° 42” S. on the New England Tablelands, proved to be the ideal climate for developing high groat percentage and large grain size. Table 5.1 shows that Garry, Acacia and VRAF.VRSF No. 23 excel in hectolitre weight, but of these VRAF.VRSF No. 23, a parent of the High-vigour cross, 28 x 23, has the highest groat percentage. Table 5.2, in giving the mean 1000 grain weights of Bundy and Belar, shows an increasing environmental stress (reduction of hectolitre weight) from east to west across the state, that is, from the Tablelands to the Western Plains of NSW. An exception to this trend was where the High-vigour cross produced their highest groat percentage at Narrabri in 1961, the warmest site and season of all those contrasted. These values were 75.8% for P4315 (cultivar inventory number 38), 75.0% for P4314 (cultivar inventory number 39) and 74.2% for P4316 (cultivar inventory number 46). Under these conditions they also exceeded Cooba in hectolitre weight. Table 5.3 shows the results of grain quality tests on the High-vigour lines in F7 generation over 3 sites. In making the F4 directed bulk, Blackbutt, from an F3 bulk, there is a tendency to produce a higher weight grain within this High-vigour cross (Cross A). Within Cross C, however, the reverse trend is shown, which is not surprising as 0614 x B, G5, has an even larger grain than Orient (Palestine x Dawn), the largest grained Australian oat variety and very susceptible to grazing and frost damage. A range of seeds from both the High-vigour cross as well as conventionally bred varieties, are presented in Figure 5.1. Recommnedations for extension For greater efficiency in the breeding and recommendation of oat varieties to farmers, northern NSW (within the summer rainfall zone) can be divided into at least 5 distinct sub-zones between 29°S and 32°S latitude. The central division (32°S to 34°S latitude) of uniform rainfall has 5 sub-zones, as is the case with the southern division (34°S latitude to the Victorian border). In the case of wheat breeding, rainfall isohyets are used for varietal recommendations. In the case of oats, a winter crop in Australia with greater vernalization requirements than wheat, which is a purely grain crop, isohyets of the length of the frost-free period (Commonwealth Bureau of Meteorology, 1965) correlate more closely with oat yields and quality.

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Table 5.1 Grain quality, as groat (gt %), of Australian cultivars and accession lines.

Glen Innesa Leetonb Curlewisc Cultivar (Inventory Number) kg/hl wt gt% wt gt% Wt gt%

Acacia (1) Algerian (70) Belar (9) Bundy (15) Fulghum (27) F. Ga W4597 (31) F. Ga W4605 (32) F. Ga 1183 (28) F. Ga W4584 (30) F. Ga 2116 E57 (33) Garry (34) K 69B.G.R. (54) Mugga (58) Orient (210) VRAF.VRSF (23) 28 x 23, 843 (37) 28 x 23, 851 (38) 28 x 23, 856 (39) 28 x 23, 871 (42) 28 x 23, 886 (46)d 28 x 23, 898 (48)

56.12 47.70 52.37 48.63

- 51.13

- - -

53.00 58.39 44.58 53.62

- 54.24

- - - - - -

42.0 42.6 45.1 43.0 46.2 36.1 31.0 32.3 37.9 37.6 35.3 40.6 39.6 51.0 44.0 45.4 40.3 41.6 46.8 33.8 43.2

70.0 68.0 72.0 68.0 71.5 71.5 71.5 70.5 71.8 76.5 68.0 71.3 70.5 66.6 74.0 72.2 73.9 74.0 72.0 72.5 71.2

- -

42.0 41.3

- 32.1

- - - - - - - - - - - - - - -

- -

70.7 68.7

- 69.5

- - - - - - - - - - - - - - -

- -

38.6 37.8

- 36.4 28.0

- - - -

40.4 -

51.9 - - - - - - -

- -

72.8 69.5

- 71.2 71.0

- - - -

73.0 -

70.6 - - - - - - -

a New England Tablelands, NSW; b Murrumbidgee Irrigation Area, NSW; c Liverpool Plains, NSW. Trials were conducted during 1959; d later reselected in 1961 as P4319 for extensive yield testing and released to farmers as Blackbutt in 1974; kg/hl = kg/hectolitre (weight in kilograms per unit volume); Wt = weight of 1000 seeds (groat + hull) in grams.

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Table 5.2 Oat grain quality, as weight in grams of 1000 seeds (groat + hull) and as groat %, of cultivars at four sites in Northern NSWa.

Glen Innesb Tamworth Ib Gunnedahc Tamworth IIc Narrabri Ic Narrabri IIb Cultivar (Inventory Number) wt. gt.% wt. gt.% wt. gt.% wt. gt.% wt. gt.% Wt. gt.%

Acacia (1) Algerian (70) Avon (1010) Belar (9) Blackbutt (11) Bundy (15) Burke (17) Cooba (22) Coolabah (105) Fulghum (27) Fulgrain (104) Klein 69B (53) Mugga (58) Orient (210) Victorgrain (114) 843 G59 (37) 851 G59 (38) 856 G59 (39) 871 G59 (42) 871-1 G59 (43) 886 G59 (46) Belar-Bundy meansd Increasing stress (1 to 6)

32.9

- 38.5 37.2

- 35.6

- 34.7

- - - -

31.5 41.6

- 36.5 29.7 34.4 36.8

- 30.1

36.4

1

72.0

- 65.3 72.7

- 69.3

- 75.5

- - - -

72.0 71.7

- 73.0 74.3 73.0 73.6

- 71.5

32.8 32.8

- 36.2

- 35.5 25.0 29.9

- 30.1

- 28.0 30.9 45.2

- 36.0 26.8 31.1 34.5

- 30.1

35.9

2

67.8 70.5

- 71.1

- 70.9 70.0 70.8

- 67.9

- 66.7 71.5 71.0

- 69.9 69.2 69.6 69.5

- 71.5

- - -

38.1 32.7 31.5

- 29.8

- 28.1 27.6 31.4

- 35.0 27.6

- 25.4 25.7

- -

28.6

34.8 3

- - -

71.1 69.2 69.9

- 72.7

- 71.6 69.8 68.5

- 70.0 69.0

- 72.9 73.1

- -

73.3

- - -

30.3 32.7 33.3

- 30.9 39.9 28.9 25.6 32.4

- 40.9 31.4

- 27.1 28.7 33.8 34.5 27.5

31.8

4

- - -

70.5 69.2 70.4

- 74.3 62.8 71.7 63.0 64.8

- 67.7 66.7

- 72.0 70.8 71.0 71.6 71.0

- - -

29.0 25.9 28.8

- 29.3

- 26.9 22.2 32.1

- 38.0 25.6

- 23.7 28.0

- -

26.7

28.9 5

- - -

68.0 71.1 67.4

- 73.3

- 68.5 60.1 68.8

- 70.3 65.7

- 71.9 72.9

- -

73.2

28.5 28.9

- 25.9

- 25.1 19.0 24.6

- - -

30.2 31.7 32.3

- -

25.2 26.3

- -

28.9

25.5 6

72.0 75.4

- 71.4

- 73.5 71.1 74.5

- - -

71.3 74.0 69.0

- -

75.8 75.0

- -

74.2

a wt = weight of 1000 seeds (groat + hull) in grams. gt% = groat percentage, b 1961, c 1963, d averages of seed weight of 2 cultivars common to all sites and years. Grain finishing stress increases from east to west, from moist to dry climates.

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Table 5.3 Grain quality of mainly High-vigour oats (Cross C and Cross A, 28 x 23), F7 generation testing.

Tamworth Gunnedah Narrabri Cultivar (no.)

wt.a gt.%b wt. gt.% wt. gt.%

Cooba (22) Coolabah (105) Fulghum (27) Orient (210) 0614 x B, G5 (Cross C), F2 0614 x B, G11-1 0614 x B G32-7 Belar (9) Bundy (15) 850-28-1 (start of Cross A) 851-0, P4315 (38) 851-7 851-7-1 856-0, P4314 (39) 856-1, P4317 (40) 856-201-1 869-67-1 871-0, P4322 (42) 871-1, P4318 (43) 871-41-1 871-48-3 871-53-1 871-56-2 871-80-1 871-80-2, P6093 (44) 871-80-3 871-80-4 871-80-6, P6094 (45) 871-165-1 871-165-9 886-0, P4316 (46) 886-n, Blackbutt (47)

30.9 39.9 28.9 40.9 42.4 33.2 35.9 30.3 33.3 36.2 27.1 28.7 33.2 28.7 31.6 36.9 32.6 33.8 34.5

- 39.4 39.2 37.7 38.4 37.7 38.2 39.9 36.6 37.6 36.8 27.5 32.7

74.3 62.8 71.7 67.7 72.9 73.0 71.1 70.5 70.4 69.6 72.0 70.4 71.5 70.8 72.6 73.6 70.6 71.0 71.6

- 69.0 69.0 72.0 71.1 70.6 70.5 71.0 71.3 70.9 67.8 71.0 69.2

29.8 -

28.1 35.0 36.1 35.6 34.5 38.1 31.5 34.0 25.4 27.4 29.9 25.7 30.3 31.7

- 27.9 28.0 35.6 32.0 36.2 31.2 33.4 35.7 32.8 33.2 33.9

- 42.0 28.6 31.4

72.6 -

71.6 70.0 74.4 73.8 72.8 71.1 69.9 76.3 72.9 71.9 73.4 73.1 75.3 76.4

- 73.2 75.4 73.1 70.4 74.8 73.7 72.3 70.5 72.7 71.7 72.9

- 72.7 73.3 73.3

29.3 -

26.9 38.0 30.7

- 30.2 29.0 28.8 27.5 23.7

- 27.0

- 26.0 24.4

- 33.5 31.5 36.2

- 35.1 28.9 32.8 30.8 28.7 33.5 28.5 28.9 37.9

- 25.9

73.3 -

68.5 70.3 71.6

- 71.1 68.0 67.4 68.3 71.9

- 72.9

- 72.0 72.1

- 73.3 69.9 73.4

- 70.1 68.5 70.3 69.5 68.4 71.4 67.7 70.0 71.3

- 71.1

a grams/1000 seeds; b groat percentage. Tests made at the NSW Department of Agriculture, Agricultural Research Station, Glen Innes. Note large grain size of 871 family and of 0614 x B, F2. Grain samples were from yield trials conducted at Tamworth, Gunnedah and Narrabri in 1963.

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Figure 5.1 Grain shape and sizes of the parents of the High-vigour cross.

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Figure 5.2 Grain shape and sizes of the High-vigour varieties and lines (Blackbutt, Carbeen and P4315) alongside conventionally bred cultivars. The NSW Department of Agriculture High-vigour selections have been proven in some cases to combine grain quality per se with milling and dual-purpose characteristics. These findings indicate the potential of these lines for current oat breeding programs in the winter rainfall areas and in summer rainfall countries on the eastern sides of continents, like China and the eastern sides of North and South America.

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CONCLUSIONS The results presented in this chapter suggest that northern NSW could be divided into 5 climatic regions, from east to west, for the purpose of recommending oat varieties. Glen Innes, at an elevation of 1,128m and latitude 29° 42" S on the New England Tablelands, proved to be the ideal climate for developing high groat percentage and large grain size. A sixth climatic zone, located in Leeton, NSW, at elevation at 152m and latitude 34° 33" S., was the second most favourable centre, but required irrigation for full grain development. At least three oat breeding and testing centres are needed for NSW: Glen Innes, Tamworth and Temora. Frost and grazing resistant oat varieties are greatly needed to implement sound rotation of crops, early fallowing to kill wild oats (Avena fatua), rather than heavy reliance on herbicides, which are bringing out resistances in this weedy species, and early sowings of oats in February and March to prevent erosion and allow flexibility in the management of livestock. The grain quality tests reported in this chapter may become increasingly significant particularly in the light of the relatively recent discoveries of the effects of oats on human and animal health and the indispensable role of oat bran, formerly not so highly regarded.

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REFERENCES Commonwealth Bureau of Meteorology. 1965. The Climate and Meteorology of Australia, Bulletin No. 1.

Frey, K.J. 1992. Setting and achieving oat breeding goals to meet specialised and new end uses. 4th International Oat Conference, Adelaide.

Guerin, P.M. 1965. Bundy – The breeder’s report: A new mid-season oat variety for northern NSW. Agricultural Gazette NSW 76: 667-669.

Mengersen, F. 1963. Choosing oats for grazing and grain in Southern NSW. Agricultural Gazette NSW 12, 678-683.

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CHAPTER SIX

PLANT BREEDING METHODS AND TECHNOLOGIES

FOR INCREASING OAT CROP YIELDS

This chapter discusses plant breeding methods and technologies and their potential for increasing oat crop yields and oat crop improvement. It specifically introduces the importance of hybrid vigour and a non-stress environment for higher percentage heritability selection and therefore providing a more productive conventional plant breeding method for the improvement of crops. This chapter draws together the results from trials presented in Chapters Three and Four to show the superiority of the Isolection method over the conventional oat breeding method for development of high yielding, multi-purpose oat varieties. GM technology and crops derived from cloning, a process devoid of hybrid vigour, are compared with proven plant breeding methods. INTRODUCTION In this chapter, GM technology and conventionally bred crops are briefly compared with proven plant breeding methods, with respect to hybrid vigour and the economic viability of both systems. These proven methods of plant breeding are also considered historically. These methods are (A) traditional landrace cropping, (B) conventional Mendelian breeding and (C) Isolection Mendelian breeding. There are important reasons for considering the limitations of comparing the yields of GM and conventionally bred crops and these are discussed in this chapter.

GM PLANT BREEDING TECHNOLOGY Overview GM crops have one parent only, to which is transferred only one, or a limited number, of genes from an organism in another genus (hence the term “transgenic”). Currently this gene (or genes) gives the plant resistance to chemical spraying for control of either weeds or pests. Conventional crops, on the other hand, derive from crossing intraspecific varieties to unite a multitude of “matching genes” from two parents, conferring hybrid vigour. This hybrid vigour, in varying degrees, applies to all conventionally bred crops but not to GM crops. For example, close crossing within the ecospecies, like winter oat x winter oat, a feature of the Isolection system, gives more hybrid vigour to the progeny than wider crossing, like winter oats x spring oats. This fact was established at Glen Innes in NSW (see previous chapters). Furthermore, yield comparisons are invalid without specifying the environment and its interaction with the varieties being compared. There are proven agro-ecological factors of weed and pest control: crop rotation and length of fallow, specially suited for high-yielding conventional varieties, depending on regional soils and climates (Fettell, 1980).

Information on comparative crop trials of GM versus non-GM crops in Australia has been limited (Guerin and Guerin 2003). Studies are, however, emerging outside Australia

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comparing GM crops with conventionally bred crops, though few are scientifically designed for meaningful comparison of GM crops with those that are conventionally bred. For instance, a report by the British Soil Association, on GM crops in North America, found that with the exception of crops possessing Bacillus thuringiensis (Bt) for pest resistance, the GM crops yielded lower than conventional crops (Anonymous, 2002a). Despite higher yields with Bt corn, US farmers lost $US 1.31 per acre ($A6 per ha). The Soil Association reported widespread contamination of seed sources, crops and the human food chain, with GM crops costing the US economy $US12 billion over the previous two years. Another report from the Canola Council of Canada seemed to favour GM crops (Anonymous, 2002b). The Council reports an average increase of $C14.33/ha in net returns to Canadian farmers growing transgenic canola, but 37% of Canadian farmers were staying with conventional lines because the cost, $C37/ha, of the Technology Use Agreement, was prohibitive. In addition, if a hybrid GM canola was being reported, it should have been compared with a hybrid non-GE canola, which would normally yield higher than its transgenic counterpart. In Arkansas, researchers found that transgenic soybeans yielded almost 10% lower than conventional soybeans (Lappé and Bailey, 1999).

Description of GM technology Gene technology enables plants to be cloned from a single cell of the parent plant. Gene transfer technology then enables cloned genes for a desired trait to be blasted into cultured plant cells with a “gene gun” that forces the genes into the cell. The cells are cultured to form a tissue mass that will grow into a plant carrying the gene or genes for herbicide or pest resistance. This results in a source of inefficiency for breeding programs and high cost, in transferring the new gene into a commercially desirable conventional variety of the crop species being modified. Out of millions of plant cells that are bombarded with metal particles coated with DNA, only very few cells actually take up the DNA. If the tissue piece were then cultured, the untransformed or native cells of the invaded plant (selected to take the gene) would rapidly grow and swamp the few cells that had the added gene. Therefore, selectable marker genes are used to favour the growth of the cells that carry the new gene. A selectable marker gene is a gene that confers resistance to a substance that is toxic to normal plant cells. This marker gene is delivered to plant cells with the introduced gene and the cells are cultured in the presence of the toxic compound, as well as plant hormones to induce the cells to divide and grow. Only cells that contain the marker genes as well as the new gene (for pest or herbicide resistance) are able to inactivate the toxic compound, in order to survive and grow into complete plants. The selectable marker genes may be antibiotic resistance genes conferring resistance to antibiotics, or herbicide resistance genes that confer resistance to herbicide.

The importance of limiting gene transfers into wild populations Outcrossing of GM with non-GM plants complicates the study of taxonomy and should be rigorously excluded from the Vavilovian5 centres of origin of specific crops and wild relatives. For example, GM mustard plants were found to be 20 times more likely to interbreed with related species than non-GM mustard plants (conventionally bred for the same herbicide resistance) (Burgelson et al. 1998). It has been reported that GM tomatoes have been grown without the consent or knowledge of Authorities in Guatemala, a Vavilovian centre, where hundreds if not thousands of indigenous tomato varieties are grown. It is also claimed that cross-pollination distances needed for strict isolation have been ignored, even for 5 Geographical centres of origin that possess plant varieties with unique genotypes and naturally occurring biodiversity, and are usually isolated by geographical barriers.

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pharmaceutical crops, so long as potential dangers, in the 1995 joint consultation between WHO and FAO, were “judged to be unrelated to food safety” (Anderson, 2000). CONVENTIONAL CROP PLANT BREEDING This is independent of genetic modification and may be divided into three productive methods or systems developed over 8-10,000 years and according to the results of particular plant breeders: (A) Traditional, (B) Conventional Mendelian and (C) the Isolection Mendelian breeding systems. These mechanisms are natural, like the agents of wind, pollinating insects and honeybees, all of which are prevented from causing evolution by means of the genetic barriers between species and even ecospecies. Here, however, the breeder controls the hybridizations and selections. All three methods can benefit from the heritability of selections made in non-stress conditions (hand spacing of plants, not drill sowings) as has been achieved with oats in Australia (refer to Chapter Three).

Traditional landrace cropping This is and has been a very successful period of maintaining peasant landraces of different species and ecospecies in the various so-called Vavilovian centres of origin of our cultivated crop plants. These are mixtures of homozygous plants most suitable for their particular soil and climatic conditions, e.g., small-seeded, rust-resistant varieties or eco-species in continental climates and large-seeded, early maturing types, in Mediterranean climates. These centres are also reservoirs of genes for high yield. Maize trials show that the degree of heterosis, when open-pollinated varieties are used in hybrid combinations, is considerably higher with varieties from Latin America (rich in Vavilovian centres) than with US Corn Belt varieties (Mangelsdorf, 1952).

There is ample evidence that our various crop species have had single and sudden origins. The great genetic variability present in isolated peasant farmers’ landraces suggests that they were created, not from single plants, but from a multitude of “first parents” to produce their multicultural (due to companion cropping) varieties with resistance to a broad spectrum of rusts, blight and climatic variability. The companion cropping of peasants often reduces disease and increases total yields.

Vavilov recorded the various large-seeded varieties of the Mediterranean centre of origin, relative to the continental centres. His critics put this down to the greater antiquity of Mediterranean agriculture but Vavilov found this to be no greater than that of Asia Minor, Afghanistan or China. Oat grazing trials at Glen Innes after 1957 vindicated Vavilov (see earlier chapters of this book). Farmers in the centres of origin should be encouraged to separately maintain their landrace varieties, free from introduced high yielding varieties, which soon succumb to rusts and blight. These unique centres are, or should be, universal reservoirs of germplasm in situ for all plant breeders, in preference to under-utilised gene banks (Harlan, 1992).

Inbreeders and outbreeders. Here we must distinguish out-breeders like maize from self-pollinated crops like wheat, oats and barley, peas and beans. The latter are designed to be resistant to inbreeding and respond well to pure-line breeding. There is enough natural crossing (4% in wheat, 0.5% in oats) to maintain their yields in the centres of origin.

Darwin was probably right in stating that selection, over thousands of years, had not made our crop plants higher yielding (Darwin, 1868). Not until the 20th century did hybridisations and

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introductions from the centres of origin combine to give significant increases in crop yields, and this is shown in the following sections.

Conventional Mendelian plant breeding During Gregor Mendel’s life (1822-84), hybridisations between different varieties, or ecotypes within the same species, formed the basis of the Mendelian laws of inheritance. G.H. Shull later showed that the depression in yield, following inbreeding of maize, was due to homozygosity. He hypothesised that hybrid vigour must be associated with the heterozygosity arising from crossing. In 1914, he proposed the term “heterosis” for this effect. His single-cross interline hybrids, however, yielded much lower than a standard maize variety on the same area. In 1917, D.F. Jones used double-cross interline hybrids to reduce the cost of seed sufficiently to justify hybrid seed production. This could increase maize crop yields by 25 to 35% and sometimes by 50%, as compared with the best selected open-pollinated varieties (Guzhov, 1989).

Natural selection. Regarding self-pollinated crops, it was assumed for half a century after Darwin that by selecting a certain type of plant for propagation, the species or variety would be continually transformed in the same direction. This was a result of acceptance of Darwin’s evolution theory and later of Galton’s “law” of inheritance, as applied to selection. Selection work commenced by W. Johannsen in 1901 on common garden bean, Phaseolus vulgaris nana var. Princess, refuted this theory in papers he wrote from 1903 to 1913 (Babcock and Clausen, 1918). Princess was actually a blend of highly homozygous pure lines. Johannsen found that selection within a pure line was without effect. Louis de Vilmorin’s wheat plants also remained identical in all respects after 50 years during which annual selection had been continued.

T.H. Morgan (1866-1945) also rejected the possibility of natural selection bringing about evolution and found that pleiotropy, the state in which one gene has effects on a number of different traits, could control several factors in Drosophila and even cause reduced fertility. This led to the hypothesis that genes occurred in linear order along the length of the chromosome. This concept could explain linkage, which enables a group of genes to be inherited together. This was a great help to conventional breeders. Conventional Mendelian breeding reached a high point with the Green Revolution, from 1950 to 1990, when world population doubled while food production quadrupled (referred to in Chapter One). Isolection Mendelian plant breeding (Isolection Method) The Isolection system of breeding (described in Chapter Three) was conceived and executed for the first time in Australia at the New England Agricultural Research Station, Glen Innes, in the drought year of 1957 (Guerin and Guerin, 1992). All the early generation oat plants were widely spaced, at 3.66-5.38 plants/m2, in contrast to 13.99-21.53 plants/m2 in the Temora Research Station drill-sown breeding plots. The object of this wide spacing was to eliminate environmental variance (due to competition and stress between plants) and to make more effective prostrate genotype selections. This method of breeding of oats (described in Chapter Three) has lead to the development of varieties of oats that are significantly higher yielding than traditionally or conventionally bred varieties. As an example, one trial at Richmond demonstrated the differences in yield in oat varieties bred by conventional and the new Isolection method clearly as all High-vigour (isolection-

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bred) varieties and lines (5 in total) reported higher pasture cut, hay and total yields than any of the conventionally bred varieties and lines. Another example of this higher yielding attribute of the Isolection-bred variety, specifically Blackbutt, was also demonstrated in a trial at Gunning in 1999 where it was tested with the conventionally-bred variety, Nile (Table 6.2). Nile is a cross between Blythe (winter type) x Avon (frost susceptible type) and was significantly inferior to Blackbutt in grain yield after 2 grazings. Numerous trials were conducted in Australia, particualry NSW, comparing the yield attributes of Isolection-bred and conventionally-bred oat cultivars and these have been presented in Chapters Three, Four of this book, over the 29 year period from 1961 to 1990. Table 6.3 draws together 12 of these trials, all of which provided statistically analysed results and that were conducted independently by researchers and agronomists other than the Author. This table presents the data previously reported in earlier chapters as yield ratios of Isolection-bred to the conventionally-bred oat variety (Cooba). Cooba was selected as a check variety for grazing and grain because it has been available to Australian farmers prior to the breeding of the Isolection-bred lines and varieties. In 83% of these trials, isolection-bred varieties reported higher yields than Cooba for pasture cut yields. In all (100%) of these trials, the grain recovery after grazing, grain only and total biomass yields were higher for Isolection-bred varieties than Cooba. In one trial, the Isolection-bred variety Blackbutt, reported three times the pasture cut yield of Cooba. In another trial, the Isolection-bred variety, Carbeen, yielded more than twice that Cooba for grain recovery after grazing. In summary, the ratios of Isolection bred varieties to conventionally-bred varieties for: pasture cut yield was 1.1, for grain recovery after grazing was 1.8; for grain only yield (i.e no grazing) was 1.5; and for total biomass yield (pasture cuts and grain after grazing) was 1.4. These ratios clearly demonstrate the superiority of the Isolection varieties and lines for total yield over the standard or check variety, Cooba, which is a conventionally-bred cultivar. In particular, the data in Table 6.3 highlights the stand out performance of Isolection-bred varieties and lines for producing grain yields after grazing. Features of the Isolection breeding method. The features of this breeding method are: (A) A high rate of success in crossing oats before starting, in order to produce a large number of homozygous F2 plants; (B) The two parents to be phenotypically similar (as in a narrow cross) but genotypically different; (C) The F2 generation plants to be widely spaced by hand, hence the name of Isolection system, to “isolate” pure breeding lines; (D) Linkage assists the rapid breeding method, by telling us that a winter cereal has morphological features like prostrate habit of growth and deep root system, correlated with resistance to frost, drought and grazing damage. Wild gene transfers are not necessary. Regarding the close crossing aspect of the Isolection breeding system, there is no lack of genetic engineering techniques to transfer genes from wild species to new cultivars but even within the limits of complete fertility and genetic exchange, there are severe restrictions on quantitative gene transfer (Röbbelen 1978). Röbbelen points out that even European potato breeding, based on only 5 introductions and with only 10 alleles for a given trait, can produce millions of genotypes. Wild oat species found in Spain and Asia Minor are very rare on deep fertile soils and seem to have few economic virtues (Rajhathy and Thomas 1974). With sound crop rotations and good farm management, there is no need for wild gene transfers. Even diploids (2n = 14) like Pilosa, Ventricosa, Prostrata, Damascena and Longiglumis cannot be crossed together.

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COMPARING GM WITH CONVENTIONAL CROPS This section highlights the differences between the 2 main systems of breeding, with respect to breeding mechanism, benefits, costs, risks and agro-ecological factors, which are summarised in Table 6.2. Yield comparisons between GM and conventional crops, which were recently investigated in a survey of available data, showed that there have been no readily available yield per se comparisons (Guerin and Guerin, 2003). There was no clear evidence that biometrically designed or analysed trials had been carried out or published in the United States. Similarly no such trials of GM cotton versus conventional cotton had been carried out openly at the Narrabri cotton station in NSW, Australia, before GM cotton (the only GM crop grown in Australia) was released to cotton growers. This was the first time that NSW Agriculture had failed to provide Australian growers with an independent yield analysis before allowing a completely alien type of variety to be grown6

There are some key comparisons between GM and conventional breeding. Firstly, conventional breeding is a natural technology and is more rapid than GM crop development. A greater length of time is required to backcross to elite conventional lines, make selections and build up seed supplies of new GM varieties for yield testing in comparison with conventional varieties. There are no yield comparisons in Australia of crops bred by conventional versus GM technology with the consequence that GM cotton has been released to farmers without any yield information. Breeders of conventional crops, on the other hand, can release a new variety every two or three years but are obliged to furnish State Departments of Agriculture with several years of biometrically analysed yield data.7 Secondly, only a limited number of genes and no hybrid vigour are added by the GM process. This makes GM technology unsuitable for the multigenic requirements of winter cereal breeding for high grazing and grain yields. In conventional or Isolection (Mendelian) plant breeding, one looks for traits, not genes: a big advantage over GM crop production, which adds only one or a few genes. Thirdly, GM crops have the advantage that they can be sprayed to kill weeds that emerge with the crop but the early competition involved will reduce crop yield. The no-till fallow of GM crops does, however, have other disadvantages (A) rodent, insect and disease incidence increase due to surface residues and (B) soil temperature may decrease by as much as 6°C at a depth of 2.5 cm in spring, giving poor germination (Anonymous, 1982). Fourthly, to gain full benefits from conventional cropping, farmers must plan for weed-free sowing conditions. Fallowing cultivations are essential for Central and Northern NSW and for Queensland, although no-till fallowing by herbicide spraying can replace some fallow cultivation (Percival, 1979). Finally, the cost of GM seed is high relative to conventionally bred varieties because of the seed patenting process.

6 The Author requested this information from the Director-General and officers concerned on a number of occasions, before the release of GM cotton, but never received any replies. 7 The Author released 3 new oat varieties: Bundy in 1965, Mugga in 1966 and Blackbutt in 1974, as a result of 7 years of oat plant breeding from 1957 to 1964.

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Table 6.1 Isolection-bred versus conventionally-bred oat varieties (Richmond, NSW)a.

Yield (T/ha) Breeding Method

Cultivar Cultivar Origin

5Pb Hayc July Pd Total

Frost Scoree

Isolection P4315 High-vigour cross, Glen Innes 6.55 3.62 1.45 10.17 1 P4314 High-vigour cross, Glenn Innes 6.21 3.70 1.23 9.91 1- Blackbutt High vigour cross, Glen Innes 6.67 2.86 1.35 9.53 1 871-1G59 High-vigour cross, Glen Innes 5.66 2.97 0.83 8.64 2 871G59 High-vigour cross, Glen Innes 5.60 2.99 0.74 8.59 2 Conventional Klein69B Argentina 5.01 3.37 0.72 8.38 2+ Cooba Temora 5.18 2.21 0.95 7.39 3+ Fulghum USA 4.87 2.20 0.64 7.07 3 F x Vic Temora 4.21 2.47 0.52 6.68 4+ Coolabah Temora 4.09 2.08 0.45 6.17 6+ FxAvon 21 Temora 3.89 2.23 0.36 6.12 4+ Avon x Fk Temora 3.96 1.93 0.28 5.90 7+ Avon x O Temora 4.04 1.81 0.33 5.85 8 FxAvon 20 Temora 3.45 2.11 0.23 5.57 7 Fulmark Temora 3.78 1.70 0.20 5.48 9 M1305 Temora 3.36 1.48 0.25 4.85 7 Algerian Algeria 3.38 0.60 0.19 3.98 8 SDf - 0.90 0.99 0.34 1.54 -

a This trial was conducted at Hawkesbury Agricultural College, Richmond, NSW in 1966; b5P = 5 Pasture cuts in dry matter yield per hectare; c Hay = hay recovered after 5P; d July P = Pasture yield during coldest month; e Frost scored 0 for no damage and 10 for extreme damage, during a cold, dry winter (rainfall only 50% of the 86-year mean). Date of Sowing: 25th March; f SD = significant difference, obtained by biometrical analysis performed by NSW Agriculture Biometricians at Rydalmere, NSW, Australia, during 1966-1967.

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Table 6.2 Isolection-bred versus conventionally-bred oat varieties (Southern Highlands NSW)a. Breeding method

Cultivar Origin (oats) P cut 1 (t/ha) P cut 2 (t/ha) Grain recovery (t/ha)

Isolection Blackbutt Glen Innes 3.49 1.46 3.70

Conventional Nile Tasmania 4.00 1.33 3.10 Eurabbie Temora 4.17 1.37 1.80 SDb 0.65 0.26 0.50 CV (%)c 10.72 12.6 14.5

a From Powell (2000). The above trial was sown on 1st April, 2000 and was harvested on 22nd December; b = significant difference; c = coefficient of variation. Grazing date for P cut 1, was 11 June 1999 and for P cut 2, 20 August 1999.

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Table 6.3 Yield ratios of Isolection-bred to conventionally-bred oat variety (Cooba) across climatic zones from statistically analysed trials over a 29 year perioda.

High-vigour Line Yield /Cooba Ratios Trial location Year

Pasture Cuts

Grain Recovery

Grain Only

Total Biomass

Climatic Zone

Tamworth 1961 1.2 1.9 - 1.3 Summer rainfall

Glen Innes 1962 1.1 2.9 - 1.3 Summer rainfall Glenn Innes 1962 1.2 1.6 - 1.2 Summer rainfall Richmond 1966 1.3 - - 1.4 Uniform/Summer rainfall Cowra 1966 1.0 1.5 - 1.3 Uniform rainfall Niangula 1969 1.3 - - - Summer rainfall Redderstone 1969 1.0 1.3 - - Summer rainfall Tamworth 1973 0.9 1.9 - 1.9 Summer rainfall South Australia 1980 1.1 2.2 - 1.5 Winter rainfall Colleambally 1985 - - 1.5 - Winter rainfall Adelong 1985 - 1.2 - - Winter rainfall Blayney 1990 0.9 1.5 - 1.2 Uniform rainfall Average 1961-1990 1.1 1.8 1.5 1.4 All zones

a The ratios in this table are derived from the data tables presented in Chapters Three and Four.

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Conventional plant breeding in Australia has been conducted hand-in-hand with crop rotations, judicious fallowing (cultivation of moist soil, or sheep grazing if the soil is dry). Contour tillage and contour banks can prevent erosion and store extra moisture. Sheep grazing prevents weed seeds from setting and increases soil organic matter. Both in Australia and America, judicious fallowing, has been recommended for the past 50 years (Guerin, 1961). Thus, a 9-month fallow can give a 100% yield increase over a 3-month fallow (Fettell, 1980). Fisher (2006) considers that GM crops are unlikely to deliver practical results in terms of yield potential increase for long time, and any increases from the technology are entirely conditional on much more investment in plant research, plant and crop physiology, conventional breeding and genetics. IMPORTANT CHALLENGES FROM GM CROP TECHNOLOGY

There are potential unintended consequences from GM technology. For instance, gene technologists claim that they are only controlling evolution. In fact they merely show that genetically modified organisms have a very low survival rate and that evolution, if it ever happened, was not by this process. This, however, should not be used as an argument for releasing genetically modified organisms. Cross-pollination can take place, giving rise to undesirable or weedy plants, animals or fishes, lacking in health and true hybrid vigour, or euheterosis.8 Genetic modification reduces euheterosis and depends upon backcrossing to elite, high yielding conventional varieties, before release. Growing GM crops also presents a risk of contaminating conventional crops. This has resulted in litigation and the loss of premium markets in the UK, Europe, Japan, China and other countries. GM crops have to contend with consumer resistance. This is based on evidence that long-term nutritional concerns are not being monitored. There is also a strong ethical component, upholding the genetic integrity of the species. This point need not, however, lower the value of gene technology, excluded from the natural environment, for fundamental research, as in glasshouses. In relation to oats, the health benefits from this grain are due to its unique properties. The β-gluten in rolled oats and especially in oat bran is unique in the treatment of cholesterol, blood pressure, heart disease and diabetes. The oat lipids are rich in antioxidants considered to be important in the treatment of cancer. The superior biological value of oats relative to other grains makes it closer in make-up to a legume. These attributes have been discussed in detail in Chapter One. It is imperative, therefore, that oats be preserved from transgenic transfers. Engineered traits in plants to date are limited to a few characteristics. The engineered plants must then be back cross into a conventional variety. This chapter stresses the importance of multigenic traits for high yields by close crossing within the species.

8 Euheterosis is hybrid vigour for sexual reproduction and seed yield. It is intra-specific.

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Table 6.4 Comparing features of GM crops with conventional cropsa.

Feature GM Crops Conventional Crops

Type of breeding Cloning and backcrossing to an elite conventionally bred variety

Independent of GM: a male x female cross.

Years to breed a variety 8-10 years Every 2-3 years

Number of genes added Usually one or two genes Possibly 50,000 allelic pairs of genes involved

Source of yield benefits Controlling weeds/pests Hybrid vigour

Land preparation All tillage is replaced by herbicide spraying Some tillage is needed to kill all weeds and residues

Weed infestation risk Weeds compete early with crop and reduce yield More emphasis on fallow tillage increases yield

Cost to farmer High cost of patented seed Relatively low cost seed

Consumer acceptance High resistance Universally accepted

a From Guerin and Guerin (2003).

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CONCLUSIONS The non-stress environment of the Isolection Mendelian system resulted in the breeding of superior dual-purpose oats, relative to the conventional Mendelian system, as well as in a more effective detection of heritability. This was shown up by a more rigorous assessment of resistance to grazing, frost and drought. Grain quality was also improved. A comparison of GM crops and conventionally bred crops show that GM crops lack versatility and economic advantage. This is because GM crops are, at present, designed for weed and pest control, not for agro-ecological factors, like crop rotation and contour tillage. The necessity for breeders to have to back-cross their GM varieties with elite conventional varieties effectively slows the process of releasing new varieties. Finally, the unintended consequences of releasing GM crops, particularly in the Vavilovian centres of landrace varieties, for maintenance of valuable germplasm such as oats, should not be underestimated.

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REFERENCES Anderson, L. 2000. Genetic Engineering, Food and our Environment. Scribe Publications, Melbourne.

Anonymous, 1982. Second Australian Agronomy Conference, Wagga Wagga, NSW.

Anonymous, 2002a. British Soil Association report on genetically engineered crops.

Anonymous, 2002b. Canola Council of Canada on genetically engineered crops.

Babcock, E.B. and Clausen R.E. 1918. Genetics in Relation to Agriculture. McGraw-Hill Book Co., Inc., New York, p. 250.

Burgelson, J. Purrington, C.B. and Wichmann, G. 1998. Promiscuity in transgenic plants. Nature, 395(September 3): 25.

Darwin, C. 1868. The variation of plants and animals under domestication. John Murray, London.

Fettell, N. 1980. Higher yields from long fallow in the Central West. Agricultural Gazette NSW 91, 1: 22-24.

Fischer, T. 2006. The Donald Oration 2004 – Part 1. Agricultural Science 18, 3: 12-16.

Guerin, P.M. 1961. Breeding new oat varieties for Northern NSW. Agricultural Gazette NSW 72: 1-7.

Guerin, P.M. and Guerin, T.F. 1992. A rapid, low-technology method of breeding high-yielding oats with dual-purpose characteristics. In: A. Barr (Editor), 4th International Oat Conference, pp. 191-5.

Guerin, P.M. and T.F. Guerin. 2003. A survey of yield differences between transgenic and non-transgenic crops. Archives of Agronomy and Soil Science 49, 3: 333-345.

Guzhov, Y. 1989. Genetics and Plant Breeding for Agriculture. Mir Publishers, Moscow, p239.

Harlan, J.R., 1992. Crops and Man. American Society of Agronomy, Madison, Wisconsin.

Lappé, M. and Bailey, B. 1999. Against the Grain. Earthscan, London.

Mangelsdorf, P. 1952. Hybridization in the Evolution of Maize. Heterosis. (Editor). Iowa State College Press, Ames, Iowa.

Percival, R.H. 1979. No-till fallowing in northern NSW. Agricultural Gazette NSW 90, 3: 42-43.

Powell, C. NSW Agriculture, 2000: Winter Crop Variety Experiments for 1999.

Rajhathy T. and H. Thomas. 1974. Cytogenetics of oats (Avena L). The Genetics Society of Canada. Miscellaneous Publications No. 2.

Röbbelen, G. 1978. Transfer of quantitative characters from wild and primitive forms. Proceedings of the Conference on Broadening Genetics. Base Crops, Wageningen.

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GLOSSARY

Agrobacterium tumefaciens. A soil-borne bacterium, which causes Crown Gall disease of beetroot and fruit trees, and can be used to transfer DNA into genetically unrelated species. Additive genes. Those in which more than one gene controls a character, with each allele of a gene making a definite contribution towards the character. This is a feature of many pigment systems, as in human skin colour. This is a feature of breeding for high yields by the Mendelian, or multigenic breeding system. Agronomy. The study of crop cultivation and soil management. Its aim is to increase yields and nutritional quality, for the particular soil and climate. Allele. A particular form of the gene. Alleles are usually in pairs. When these are similar, the individual is a homozygote, designated AA. If each allele is different, the individual is a heterozygote, designated Aa. Amino acid. A subunit of a protein molecule, which contains an amino group, NH2, and a carboxyl group, -COOH. An example is lysine, an important growth factor in oat grain, relative to other grains. Anaerobic. Conditions of soil waterlogging inimical to the aerobic bacteria, without which man would starve unless the land is drained. An anaerobic organism can obtain energy from the breakdown of glucose in anaerobic respiration. Obligate anaerobes cannot survive in oxygen and include food poisoning bacteria, which cause botulism. Most anaerobes are facultative, meaning that they can live in the presence or absence of oxygen. Anatomy. The study of the structure of living organisms. Angiosperm. Plants with their seeds enclosed in an ovary. They can be monocotyledons (grasses and tulips) or dicotyledons (apple or oak). Antibiotic. Substance obtained from a micro-organism that destroys or inhibits the growth of other micro-organisms, particularly disease-causing bacteria and fungi. Antibody. A protein manufactured by the immune system in response to the presence of an invading foreign body or antigen. Each antibody fits exactly to an antigen and destroys it. Antigen. A substance that causes the immune system to produce antibodies. Archetype. The hypothetical ancestral type postulated by evolutionists for a particular species. Asexual reproduction. The process by which organisms multiply without the formation of specialised sex cells or gametes. This results in genetically identical progeny called clones. Potatoes can be grown from seeds but are mostly cloned.

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Autopolyploid. A polyploid with multiple and identical sets of chromosomes (genomes), like lucerne (alfalfa), all the genomes being derived from the same species. Autosomal inheritance. The alleles are located on autosomes, i.e. not on sex chromosomes, so that mating is not affected by the sex of the parent and reciprocal crosses will give identical results. Autosome. A type of chromosome found in all cells not involved in sex determination and carrying the major part of genetic information in cells, including information on sexual characteristics. From Gr. soma,body. Backcross. A genetical term indicating a cross between a parental strain and its progeny from a previous cross. Mendel’s backcross to the recessive parent is called a testcross, a diagnostic test to reveal the genotype of the hybrid. Bacillus thuringiensis (Bt). A bacterium with the ability to produce a crystal protein toxic to certain insects, mainly Lepidoptera (caterpillars and butterflies) insects. Since the 1950s farmers have used the Bt toxin to control crop insect pests. GM crops transformed with the relevant gene from the Bt bacterium produce the same toxin. Bacterium. A one-celled prokaryote or organism. It is without a membrane-bound nucleus and, therefore, with no need for mitosis or meiosis during reproduction. They are present in soil, water, air and as free-living symbionts, parasites and pathogens. They are essential for feeding mankind through nitrogen and sulphur cycles as well as preventing cancer of the intestine, where they feed on healthy foods like oatmeal, potatoes, buttermilk and yoghurt. They show man their power to kill animals (anthrax) and mankind (tetanus). Bacteriophage. A virus that is parasitic within a bacterium. The viruses insert genes into a bacterium’s genome. These are used in GE (genetic engineering) as cloning vectors. Baculovirus. A group of viruses specific to insects, used as bio-insecticides. Base. The nitrogenous part of a nucleotide. DNA has four bases: adenine, thymine, cytosine and guanine. In RNA, thymine is replaced by uracil. The sequence of bases determines the genetic code. Base pairing. The chemical linking of two complementary bases. In DNA, adenine pairs with thymine and cytosine with guanine. In RNA, thymine is replaced by uracil. Base pairing holds the two strands of DNA together to form a double helix. Balanced diet. A diet including adequate natural foods to provide plenty of fibre, calcium, lysine, minerals, vitamins, protein, fats, oil, complex carbohydrates and water. Biogeographical region. One of 7 or 8 regions of the world which contain distinctive groups of plants and animals which are normally prevented from leaving those regions by various natural, geographical barriers. See Collins’s Dictionary of Biology by Hale and Margham (1988). See also Vavilovian centres of origin and Origins of cultivated plants. Biological control. Pest control by using natural predators (harmless to other organisms) to reduce the pest population.

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Biometry. Application of statistical methods to the study of biological measurements, as in crop variety yield trials. Biotechnology. Any natural biological procedure to produce a farm product, including the use of rennet (from calf’s stomach) in cheese making and the use of yeast in making bread, beer and wine. This includes Mendelian plant breeding but should exclude the unnatural procedures of gene technology like GE or GM, genetic modification. Biotype. Distinct physiological strain or race within an apparently morphologically uniform species. The biotype population consists of individuals with identical genetic constitution. Cell. The structural and functional unit of most organisms. The smallest unit of living matter capable of reproduction. Bacteria consist of a single cell but plants and animals are made up of billions of cells. A cell contains DNA and the organelles necessary for energy conversion and protein synthesis. Virtually every cell in an organism contains a complete set of that organism’s genes. This complete set of genes is the genome. There are 20 different amino acids that are the building blocks of the thousands of different proteins found in a cell. Chloroplast. A type of plastid containing chlorophyll, found within the cells of plant leaves and stems. The site of photosynthesis: the synthesis of organic compounds from carbon dioxide and water using the energy of sunlight. Chlorosis. A yellowing of plant leaves caused by lack of chlorophyll pigment due to mineral deficiency (magnesium or iron), or disease like virus yellows, both of which result in a decrease in photosynthetic rate. Chromosomal aberration. Abnormal structure or number of chromosomes, including deficiency, duplication, inversion, translocation, aneuploid, polyploid, or any change from the normal pattern. Chromosomes. Nucleoprotein bodies that are microscopically observable in the cell during cell division. They carry the genes, which are arranged in linear order. Every species is created with a characteristic chromosome number. They are thread-like bodies (from Gr.soma, body) comprising DNA and protein. Clone. All the individuals derived by vegetative propagation from a single original individual. Genetic engineering starts with clones. Clones lack hybrid vigour. Colchicine. A poisonous alkaloid (an organic compound containing nitrogen like nicotine, quinine, morphine and cocaine) extracted from the corms of Colchicum autumnale, autumn crocus. It is used to produce cells with double sets of chromosomes by arresting spindle formation and interrupting mitosis. Continuous variation. Multiple genes are responsible for this type of variation which is important in increasing crop yielding ability. This is one proof of the futility of gene technology, where only one or two genes are involved, for yield increase. Cross-pollination. This results in cross-fertilization or allogamy, the reverse of self-fertilization or autogamy. The female gamete fuses with a male gamete (carried in the pollen) from a different plant to produce offspring.

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Cultivar. A cultivated variety. Cytogenetics. The combined study of the inheritance of organisms and their chromosomal makeup. Cytology. The study of the structure and function of the cells. Cytoplasm. From Greek plasma, shape or body. The living contents of a cell, except for the nucleus. Jelly-like material in which the cell organelles are suspended. Cytoplasmic inheritance. The theory that inheritance, not explained by Mendel’s laws, is determined by the cytoplasm, not the cell-nucleus. The inheritance of some fertiliser treatments on flax, Linum usitatissimum, was recorded by A. Durrant (1962) in Heredity 17: 27-61. This was non-Mendelian inheritance but is very rare. Dicotyledonae. Plants with two seed leaves: potatoes, tomatoes, beans, sugar beet. DNA. Deoxyribose nucleic acid. This chemical code of information is specified by a sequence of four different nucleotide building blocks, based on consecutive sets of three of these nucleotides. A complex nucleic acid molecule in the chromosomes of most organisms which controls the structure of proteins, the double helix shape of which was discovered by John Watson and Francis Crick in 1953. Diploid. From Gr. diplos, double. Having 2 sets of homologous chromosomes in the nucleus, that is, double the number of chromosomes present in the sperm or egg-cells of the particular species. Discontinuous variation. Mendel’s discovery of distinct classes, like red versus white and tall versus dwarf, which refuted the “blending inheritance” concept of evolution, as distinct from the polygenic nature of quantitative traits like crop yields. Dominance. A genetic interaction where one allele of a gene masks the expression of the alternative allele in the heterozygote, the phenotype of which will be that of the dominant allele. The alternative allele is said to be recessive but is not lost to future generations. The term is unfortunate as the uninitiated may be led to believe that certain dominant characters are going to take over and multiply, proving that evolution is going on. Recycling law and the Hardy-Weinberg law explain what is really a stable situation. Ecology. Interactions between plants, animals and their environment suggest that organisms have been specially created, according to a design to ensure cooperation in enriching and sustaining the environment. From Gr.oikos, house; logos, reason. Environment. The surroundings of organisms. This must be taken into account when selecting plants for crop improvement. See Isolection breeding method. Enzyme. A class of proteins produced by organisms to speed up chemical reactions, from Gr.zyme, yeast. Escherichia coli. A bacterium found in the human gut which is being used extensively in biochemical and genetical studies.

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Eukaryote. A fungus, plant or animal with both nuclei and chromosomes in its cells. Excludes bacteria and blue-green algae, both of which are prokaryotes, which are without a nucleus. F1. The first filial generation after a cross.or mating. The hybrid generation in corn. F2. The second filial generation produced by crossing inter se or by self-pollinating the F1. Usually for wheat or oats, self-fertilization of the F1 is implied. F3 bulk. P4315 oats was derived from an F2 plant (851 G59), which yielded 600 seeds. These were bulked together in the F3 because of their great uniformity. This bulk of seed was given the Accession no. P4315 and proven to be the highest yielding oat that the Author has bred or is aware of. F4 directed bulks. A similar procedure was followed to produce Blackbutt because of its great uniformity in the F2 but in the F3 there was segregation for habit of growth. Only the prostrate plants were harvested and planted together in the F4, which was bulked together and given the number P4319. This proved to be the highest yielding late maturing variety and was named Blackbutt in 1974. Family. The taxon between order and genus and usually embracing more than one genus. Animal families end in -idae like Ursidae, the bears. Plant families have endings in -ceae like Rosaceae, the roses. Fitness. This is a nonsensical term for biology or genetics. The Author found oats to be higher yielding (fitter) than winter wheat but more susceptible to frost damage (interpreted by some as natural selection) than winter wheat, both sown on the same soil together in the same experiment for a number of years involving sheep grazing and exact dry matter weights. Neither fitness nor natural selection can be measured in a state of nature, where they have even less relationship to one another than in a statistically designed experiment. Fixity. A state of constancy such as is observed in the 6th generation after a cross is made. Free Trade. Trade arrangements where tariffs or other barriers to the free flow of goods and services are eliminated. Also known as laissez-faire (French for 'leave well alone'), free trade means no taxes on manufacturing goods and no tariffs paid when goods cross a border. It was an idea dear to the hearts of Victorian manufacturers and industrialists, who believed that anything that impeded free trade would reduce their profits. Gene. From Gr. genos, race. The unit of heredity: a small piece of the chromosome which influences the development of the organism carrying it. A gene is said to be a “paragraph” of DNA message. DNA carries the instructions of heredity in the form of bases or letters strung along the middle of a double spiral molecule. This message is divided into sections or paragraphs called genes. A single gene usually codes for a single protein chain. There is no satisfactory hypothesis to support evolution of the protein-synthesising mechanism: “Organisation, not substance, seems to make the difference between life and non-life.” See Gary Parker (1970), Origin of Life on Earth, Bible-Science Newsletter, VIII, No. 12, p. 4. Gene pool. The total collection of varieties maintained by a plant breeder for purposes of interbreeding to provide more and more valuable combinations of genes and their alleles

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present in the separately created species of plant to which the breeder is committed. The production of genetic diversity and its maintenance, as well as peaceful contact with the Vavilovian centres, must be the first goal of plant and animal breeders, not the production of clones to be injected with foreign genes, as in genetic engineering. This is related to Chapter 5 of this book, which sets out an oat Germplasm Inventory. Gene technology. This usually involves only one or two genes and is a flawed system that has grown out of stopgap methods of perpetuating unsustainable, agricultural monoculture, which naturally increases crop weeds, pests and diseases. Genetic engineering is the forcible injection into an organism of an alien gene (a gene from a separately created species), which has a very low success rate and is resented by the host plant. Genetic load. A measurement of the amount of deleterious genes in a population, calculated as the mean number of lethal genes per individual. This has to do with inbreeding and the incidence of mutations. Genetics. From the Gr. genesis, descent. The science of heredity and variation, which seeks to account for differences and resemblances exhibited among organisms related by descent. Mendelism deals with the mechanism by which these differences and resemblances between related organisms are inherited. Mendelism is a real predictable science of transmission genetics, in contrast with evolution, which is an ideology, but an unproven theory. Genome. The complete set of chromosomes in an individual. Genotype. The genetic constitution of an individual, all of whose genes are available for transmission to the offspring of that individual, but not all of which will be manifested in its phenotype or outward appearance. This is due to the masking of recessive traits (blue eyes, for instance) by dominant traits (brown eyes) but does not mean that the blue eyes will not be passed on in succeeding generations. Only genetics can account for hereditary transmission. Genus. A category bearing a name like Avena, for oats, represents a number of species, many of which are genetically isolated from one another and may be regarded as separate creations. Hardy-Weinberg law (1908). Proposed by G.H. Hardy in England and independently by W.Weinberg in Germany, this law explains how the various alleles (forms) of a gene could remain constant in a population and yet be inherited by the rules of Mendelian genetics. For example, dominant phenotypes are not necessarily in excess of recessive ones in a population. If a gene has two alleles, A and a, with a frequency of p and q respectively, the genotypic frequencies will be: AA Aa aa p2 + 2pq + q2 = 1.0 Such genetic equilibrium is said to occur only under conditions of zero selection, zero mutation, zero migration, random mating and a large population size. In practice, this gives rise to stability not evolution, under natural conditions, as stated under recycling law. Under controlled breeding, either allele can be selected and isolated. Harvest Index. The proportion of grain to total biomass produced by a crop variety.

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Heredity. The transmission of characteristics from generation to generation, involving genes and chromosomes. Heritability. The proportion of phenotypic variance in a population due to genetic differences. Heritability is a measure of genetic variability, necessary for the breeder’s selection to be effective. If heritability values for a particular character is low, this indicates high environmental variability and poor response to selection. Heterosis or hybrid vigour. The superiority of a hybrid resulting from a cross between two inbred lines, especially in maize, a cross-pollinated species, where the hybrids yield much higher than either parent. In the case of self-pollinated species, the mid-point of heterosis, advanced by the Author to explain the law of heterosis, is reached by crossing together individuals within an ecospecies (winter oat x winter oat), instead of crossing individuals from different ecospecies (winter oat x spring oat), even though both ecospecies, (subspecies is an older term), may have been formerly classed as belonging to one species, like Avena sativa, oats. The high yields and vigour obtained at this mid-point is advanced as a proof of the separate creation of the particular ecospecies. Thus there is no common progenitor for the ecospecies classed together under Avena sativa. The various ecospecies were created in and for the widely separated Vavilovian centres of creation, where the purity of the natural in-breeders has been maintained by geographical isolation.

Heterozygote. An individual possessing two different allelic forms of the same gene in all diploid cells, like Aa (expressed as the dominant A but carrying the recessive a for future generations). Homozygote. An individual possessing two identical forms of the same gene in all diploid cells, like AA, dominant, or aa, recessive. Hybrid sterility. This is a breeding barrier when two different species are crossed together, like the horse and the ass, each with different chromosome numbers, a different physique and preferred natural habitat. The chromosomes do not match properly during meiosis, resulting in the sterile mule. Inbreeding. Mating between human relatives leads to a loss of physical and mental health due to the accumulation of deleterious genes from a common ancestor, because Homo sapiens is an obligatory outcrossing species. Isolation. Geographical separation from other populations of the same species. The claim that isolation causes speciation (formation of new species) has never been proven. Isolation has long preserved the identity of separate creations, like kangaroos, eucalypts and other natives of Australia. Isolection breeding method. The Author developed a method of space-planting individual oat plants all from the same F2 plants and in comparison with similarly spaced plants from other F2 plants resulting from the same cross and other crosses. The purpose of this was to eliminate environmental variability and low heritability and to pinpoint superior F2 progenies. This gave rise to new high-yielding oat varieties and the concept of Isolection theory or law in opposition to the concept of selecting under conditions of high population. See also Malthusian Point.

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Johannsen, Wilhelm (1857-1927). A Danish breeder of beans, Phaseolus vulgaris, who coined the terms gene, genotype and phenotype. He called for a total rejection of evolution theory. Once a pure line, now called a variety, has been selected and isolated, it cannot evolve. Kölreuter, Joseph (1733-1806). The first of the great hybridisers and discoverer of the commercial value of hybrid seed production. In 1760 he published his Preliminary Reports of Experiments and Observations Concerning Some Aspects of the Sexuality of Plants, describing over 500 plant hybridisation experiments. He crossed two different species of tobacco, Nicotiana rustica and N. paniculate, securing a healthy, vigorous hybrid, which proved sterile when self-fertilized, the first mule of the plant kingdom. He then proposed that growers should use the hybrid seed once only to produce high-yielding tobacco hybrids. He proposed other schemes for increasing vegetative yields in crops and in forestry, which were not adopted until another 200 years had passed. Laws of Creation. Three laws may be identified as proof of the separate creation of every species and ecospecies. (1) The crossing together of different species and sometimes ecospecies results in sterility or greatly reduced yields, due to various genetic barriers. (2) The law of Isolection operates in the Vavilovian centres of origin, where self-pollinating ecospecies will occasionally cross within their own ecospecies to attain maximum heterosis and biodiversity, free of pollen from other or different ecospecies. (3) The law of Heterosis applies to both types of plants but is absolutely essential to cross-pollinating plants and to animals. The Author’s discovery of the mid-point of heterosis in self-pollinating plants is advanced as a proof that the various ecospecies have been separately created. Linkage. A number of genes or characters which are usually inherited together, on the same chromosome. This great proof of a natural tendency towards the cohesion of separate species, due to T.H. Morgan, is another argument against evolution. Linnaeus, Carl (1707-1778). The founder of the binomial system of classifying organisms, which is still used, proving that plants and animals are still exactly the same and therefore not evolving. He wrote the Systema Naturae in 1753. He was a genius for classification, a process of identifying unchangeable forms, still indispensable for all scientists. His belief in the fixity of species was confirmed by the experiments of Gärtner and Kölreuter. Malthusian point. Coined by the Author during plant breeding experiments, while watching over-dense plant populations die from drought. When Darwin claimed that natural selection would favour a small minority of such plants, he had obviously never tried a controlled experiment of this nature, in which every plant died, while in the same field there were individual spaced plants of gigantic size and large yields of grain. These latter were thriving well above the Malthusian point of death. The opposite point to this is the low plant density of spaced plants in the Isolection method of plant breeding. See Isolection breeding method. Mendel, Gregor (1822-1884). Published his masterpiece in 1866. His “Experiments in Plant-hybridisation” were not noticed or acted upon until 1900, when it was obvious to all unbiased scientists that evolution was no longer tenable. The absence of transitional forms in the second generation was a very good argument against the possibility of evolution, which was supposed to be a straight line process. Gärtner, Kölreuter and others had already observed that hybrids are inclined to revert back to the parental forms. We now know that the number of hybrids and their progeny from generation to generation is continually

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diminishing, according to formula 2n-1, where n is the generation number. For simplicity, take “A” to be a tall pea (one of Mendel’s) and “a” to be a short pea, which are both crossed together. In the first generation after a cross, we get the ratio:

1A : 2Aa : 1a and the 10th generation gives: 1,023 A : 2Aa : 1,023 a This is obviously a reversal back to the parental forms and is referred to as “the case of the disappearing hybrids.” Mendel’s laws. Mendel’s great mathematical ability and painstaking experiments showed the amazing order of nature and this order postulated two of the greatest laws of biology:

(1) The Principle of Segregation states: (a) In individuals differing in a single character and controlled by a pair of Mendelian factors, each trait appears as a unit, passes intact through the individuals of the first filial generation where it may or may not be visibly expressed and emerges unchanged in the second generation.

Or, restating it, (b) The units contributed by each parent separate in the germ-cells of the offspring without having had any influence on one another. Or, restating it, (c ) The members of a pair of factors separate into sister-gametes in germ-cell formation.

(2) The Principle of Independent Assortment was defined by Mendel after he crossed a variety of pea, Pisum sativum, with round seed and yellow albumen with another pea variety having wrinkled seed and green albumen, as follows: (a) The genes are assorted independently of each other in germ-cell formation. Or, (b) The relation of each pair of different characters in hybrid union is independent of the other differences in the 2 original parental stocks.

The above two laws show the essence of Mendelian inheritance to be particulate. The genetic constitution is composed of different and separate units. Each kind of unit can exist in a number of discrete forms. The hereditary transmission of any kind of unit is more or less independent of that of the other units, the restriction of independence being a partial one, concerned with the phenomenon of linkage (a post-Mendelian development). Linkage is an exception to Mendel’s Second Law: Genes in the same chromosome tend to remain linked together to the extent of 50 to 99.9 %. Disjunction of genes in the same chromosome, however, may take place due to crossing-over. Some other post-Mendelian developments should be noted:-

(a) A particular gene may affect the development of several distinct characters.

(b) A particular gene may be and probably is influenced by other genes. Thus all the genes of an organism may act together to form a single interacting system. A difference between two genetical systems may be in one factor only, e.g., tall or short peas.

(c) A gene may be influenced profoundly by external and internal environment.

(d) A gene may have several allelomorphs, like 13 eye-colours in Drosophila.

(e) The linear arrangement of the genes can be shown on chromosome maps, as has been done for maize and barley.

(f) Genes are affected in their expression by their actual position in the chromosomes.

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(g) Where some other crosses were found to follow other rules like “blending” inheritance, these exceptions could soon be explained by the fact that certain characters depend for their expression on the interaction of 2 or more pairs of genes. Thus the expected ratios could be modified in various ways, while the fundamental Mendelian laws of transmission remained the same. Indeed, with the exception of some traits inherited through the cytoplasm, the Mendelian transmission of genes accounts for all biological heredity. Morgan, Thomas Hunt (1866-1945). American geneticist who discovered sex linkage in 1910, using the fruit fly, Drosophila, together with a method of mapping genes on the same chromosome, the units of distance between them being called centimorgans. He is one of the founders of modern genetics. Morgan ably refuted the theory of evolution, both that of Lamarckism (by his fruit-fly experiments) and that of neo-Darwinism (by his downgrading of natural selection in The Scientific Basis of Evolution). Morphology. The study of the shape, general appearance or form of an organism, as distinct from anatomy, which requires dissection to discover the structure of an organism. Morphology is the plant breeder’s bible.

Mutation breeding. Mutagens are used to produce new genetic forms in agriculture. This is rarely successful because the vast majority of mutations are harmful. They are unnatural and likely to upset the delicate balance of genes of any species. Moreover, most mutagenic agents, whether radiations or chemicals, are also carcinogenic, that is, they induce cancers. Mutual aid. Prince Propotkin wrote Mutual Aid in 1902 to counter the over-stated ideology of “survival of the fittest.” He studied animals from Siberia to the Yorkshire Moors as well as human societies and found a law of love, sympathy and self-sacrifice. In humans, he based this ethical concept on conscience and a desire for human solidarity. In plants, which he did not study, there is the corresponding harmony of mutualism and symbiosis. Natural selection. An unscientific term coined by Charles Darwin. Oat varieties vary in their resistance to frost damage but this resistance may not be detected in a state of nature, where plants seed thickly and protect one another. Variety trials must be specially designed and planted and later subjected to sheep grazing so that the crowns of the plants are exposed to a uniform type of frost damage. The grain crop yield which recovers from this treatment may not be correlated with frost resistance as a measure of natural selection. For example, rye was found to be extremely hardy but was the lowest yielding, or least fit, of all the cereals tested. The correlation presumed by Darwin, therefore, does not hold under controlled conditions. Under natural conditions, non-human selection cannot be studied because organisms protect one another, just as tomato plants are protected from frost by planting them under spinach plants already two feet high and well established. Nitrogen fixation. Atmospheric nitrogen (80% of the atmosphere) can be made into amino-acids by free-living prokaryotes (the aerobic Azotobacter and the anaerobic Clostridium) or by Rhizobium bacteria occupying swellings in the roots of leguminous plants called nodules. The latter case is called symbiosis and is vital for nitrogen-poor soils, the bacteria getting carbohydrates from the plant host. The nitrogen is reduced by the enzyme nitrogenase to ammonia thus:- N2 + 3H2 = 2NH3. The ammonia reacts with keto-acids to form amino-acids.

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Origin of cultivated plants, centres of. (1) Chinese: millet, buckwheat, soy beans, apple, plum and citrus fruit; (2) Indian: rice, sugar cane and many tropical crops; (3) Indo-Malayan: bananas, rubber; (4) Central Asian: garlic, onion; (5) Asia Minor: grape, pear, soft wheat, rye and several leguminosae; (6) Mediterranean: hard wheat, sugar-beet and forage crops; (7) West Asian: almond, apricot; (8) Abyssinian: wheat, barley, flax and oats; (9) Central American: upland cotton, runner beans; (10) South American: potato, tobacco and sea island cotton; (11) Tropical African: cowpea and rice of O. glaberrima species; (12) European: blackcurrants and oats of A. sativa species; (13) Australian: macadamia nut, eucalyptus tree. These updating Vavilov’s original centres of origin. Phenotype. The observed traits of an organism. These may not correspond to its genotype, as when a recessive trait (e.g. frost resistance) is masked by another trait (e.g. earliness), in which case selection for frost resistance would be futile, especially if a natural selection method were relied upon. This is another reason, like pangenesis, why evolution does not happen through natural selection. Photosynthesis. The process whereby green plants absorb the radiant energy of sunlight by their chlorophyll, using carbon dioxide and water to manufacture carbohydrates, by the equation:- 6CO2 + 6H2O = C6H12O6 + 6O2 Physiology. The branch of biology that relates to nutrition, respiration, reproduction and excretion in plants and animals. Plasmid. A structure in cells consisting of DNA that can exist and replicate independently of the chromosomes. Bacterial plasmids are used to produce recombinant DNA for gene cloning. Plasmids are used in genetic engineering of micro-organisms. Plasmids are important in public health, as some types possess genes for antibiotic resistance. Polygene. One of a group of genes influencing a quantitative characteristic like height in man or yield in crops. Multifactorial inheritance is involved. Polypeptide chain. A sequence of amino acids joined together by peptide bonds, forming a protein. Polyploidy. Three or more sets of chromosomes in an organism. Most crops are diploid, e.g., maize, rice, barley, sugar beet, soy bean and other grain legumes, rubber and cacao. Others are allopolyploids: wheat, oats, tobacco, sugarcane, cottons and arabica coffee. A few are autopolyploids: potatoes and lucerne. Each was created according to its kind. Chromosome doubling can occur, rarely, in nature. Man can do it, using a drug extracted from the autumn crocus, but usually it is not worthwhile. The diploid number in barley is shown as 2n. This is 14 = 2 x 7, that is, twice the basic chromosome number in the temperate climate cereals. The diploid number of wheat is 6n = 42 = 6 x 7, that is, 6 times the basic number, hence it is called a hexaploid like oats. Any crop with a number of 3n or more is a polyploid: this excludes barley. Allopolyploids are polyploids whose somatic nuclei are said to contain 3 or more complete genomes derived from different species. There is no proof for this, as Kihara pointed out. They were probably created thus with non-homologous or only partly homologous genomes. And why not if it improved the overall design of which man is ignorant? Autopolyploids are polyploids whose nuclei contain 3 or more complete homologous (alike) genomes, obviously from the one species.

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Prokaryotes. Bacteria or blue-green algae that lack a membrane-bound nucleus and have no mitosis or meiosis. All are haploid, that is, contain one complete set of chromosomes, in contrast to eukaryotes, which can be haploid, diploid or polyploid. Quantitative inheritance or Polygenic inheritance. Inheritance of quantitative traits, like crop yield, stature, weight or colour intensity, depend on the cumulative action of many genes, each producing a small but significant effect on the same trait. The expression of a polygenic character is also affected by the environment, as human height is affected by diet. A generous environment is needed for high heritability and success in the Isolection system of plant breeding. Recombinant DNA technology. DNA that contains genes from different species, alien to one another, not by Mendelian breeding, but by an artificial, undesirable and unnecessary technique known as genetic engineering. Recycling law. Advanced here by the Author as a result of his plant breeding experiments. It represents a force within self-pollinating plants and landraces which maintains genetic stability and prevents yield variability. The best varieties bred were judged to be those which gave the highest yields on average over a number of years in one site and, still better, those which gave high yields over a number of years in more than one site, each with a different local climate. This suggested that genetic homeostasis (a term coined by Lerner) was operating to maintain the genetic stability necessary to yield well in the variable seasons which succeed one another in Northern NSW. Separate creation, not natural selection, is responsible for the genetic make-up of the landraces and crop ecospecies found in the various Vavilovian centres of crop origin, as judged by yield-testing them in different climates (those of Western Australian and Northern NSW). This seems to constitute a good argument against evolution, not unlike Kimura’s theory of neutral alleles, which are said to be independent of population size and therefore contrary to one of the requirements of random genetic drift. The Author hopes that recycling law is a clearer way of explaining genetic stability than is the Hardy-Weinberg Law. Species. The natural, or genetic, unit of classification for animals. Each has been created separately. For plants, geographical species, called either subspecies or ecospecies, from the Vavilovian centres of origin, separated by vast seas or mountain barriers, have been separately created. This is due to creative design: plants are more sensitive to the factors of environment, while most animals are mobile, warm-blooded and able to migrate away from excessive heat or cold. Man can live anywhere from the tundra to the tropics. Plants, on the other hand, are sedentary and have been created with different types of physiology to fit them for particular habitats and climates. Hence the validity of the Vavilovian centres of creation for plants. Symbiosis or Mutualism. A relationship between dissimilar organisms which benefit the partners or members of the association. Transgressive segregation. When 2 individuals, which possess some quantitative character or characters in common, are crossed together, it is possible to obtain in the progeny individuals, which possess that character or those characters in a higher degree than that of either parent. This is the key to increasing crop yields by natural, multigenic, Mendelian plant breeding. This is in contrast to genetic engineering or gene technology, which involves only one or two genes and causes yield reduction in such transgenic plants genetically modified by insertion of an alien gene.

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Vavilov, Nikolai Ivanovich (1887-1943). One of the greatest plant geneticist and appointed by Lenin in 1930. Established 400 research institutes and collected from overseas 26,000 species and varieties of wheat. His principle of maximum diversity pinpointed the world’s centres of crop origins, usually elevated areas with a wide range of micro-climates called Vavilovian centres. His work refuted the theory of the inheritance of acquired characteristics, basic to Darwinian and Lamarckian evolution. Vavilovian Centre. See Vavilov, Recycling law and Origin of cultivated plants.

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APPENDIX A

Australian Oat Statistics

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Statistical Oat Yields for NSW (1990-2001)

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1.5

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1988 1990 1992 1994 1996 1998 2000 2002

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(t/ha

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Statistical Oat Yields for NSW (1950-2001)

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1945 1955 1965 1975 1985 1995 2005

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Statistical Oat Yields for Australia (1990-2001)

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2.5

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1988 1990 1992 1994 1996 1998 2000 2002

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Statistical Oat Yields for Australia (1950-2001)

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1945 1955 1965 1975 1985 1995 2005

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Statistical Oat Yields for New South Wales and Queensland (1990-2001)

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0.5

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1.5

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3

1940 1950 1960 1970 1980 1990 2000 2010

Yie

ld (G

rain

onl

y) (t

/ha)

NSW Queensland

Statistical Oat Yields for Victoria, South Australia and Western Australia (1950-2001)

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0.5

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2.5

3

1940 1950 1960 1970 1980 1990 2000 2010

Yie

ld (G

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y) (t

/ha)

Victoria South Australia Western Australia

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APPENDIX B

Plots From A Heavy Grazing Trial (Richmond, NSW 1966)

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Individual plots at Richmond after heavy grazing (Top Left) High-vigour line, P4314; (Top Right) High-vigour line, P4315; (Bottom Left) Cooba, (Bottom Right) Avon x VR Bke on left and Algerian on right.

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Heavy grazing trial Richmond 1966. (Top) Cooba on left, Klein 69B in centre and Algerian on right; (Bottom) Plots after heavy grazing.

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