Combination of resistance to Verticillium longisporum from zero erucic acid Brassica oleracea and oi...

Combination of resistance to Verticillium longisporum from zero erucic acid Brassica oleracea and oilseed Brassica rapa genotypes in resynthesized rapeseed (Brassica napus) lines.. High [r]

PLANT BREEDING REVIEWS

Volume 31

edited by Jules Janick

Purdue University

PLANT BREEDING REVIEWS

Volume 31

Plant Breeding Reviews is sponsored by:

American Society of Horticultural Science

International Society for Horticultural Science

Editorial Board, Volume 31

I L Goldman

C H Michler

Rodomiro Ortiz

PLANT BREEDING REVIEWS

Volume 31

edited by Jules Janick

Purdue University

Copyright # 2009 by John Wiley & Sons, Inc All rights reserved.

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Library of Congress Cataloging-in-Publication Data:

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Printed in the United States of America

10 9 8 7 6 5 4 3 2 1

2 Brassica and Its Close Allies: Cytogenetics

VI Genome Dissection and Development

IX Potential Role of Arabidopsis thaliana

X Chloroplast Genomes and their Phylogenetic

v

3 Genetic Enhancement for Drought

Belum V S Reddy, S Ramesh, P Sanjana Reddy,

and A Ashok Kumar

4 Breeding for Resistance to Stenocarpella

Johannes D Rossouw, Z A Pretorius, H D Silva,

5 Cassava Genetic Resources: Manipulation

Nagib M A Nassar and Rodomiro Ortiz

IV Cassava Diversity as Revealed by DNA Markers

6 Breeding Roses for Disease Resistance 277

Vance M Whitaker and Stan C Hokanson

7 Plant Breeding for Human Nutritional Quality 325

Philipp W Simon, Linda M Pollak, Beverly A Clevidence,

Joannne M Holden, and David B Haytowitz

III Progress in Breeding for Nutrient Content

Cumulative Subject Index 395

Cumulative Contributor Index 415

Jeff J Doyle L H Bailey Hortorium, Department of Plant Biology, CornellUniversity, Ithaca, New York 14853 USA

David B Haytowitz Nutrient Data Laboratory, United States Department ofAgriculture—Agricultural Research Service, Beltsville Agricultural ResearchCenter, Beltsville, Maryland 20705 USA

Stan C Hokanson University of Minnesota, Department of HorticulturalScience, 1970 Folwell Avenue, St Paul, MN 55108 USA

Joannne M Holden Nutrient Data Laboratory, United States Department ofAgriculture—Agricultural Research Service, Beltsville Agricultural ResearchCenter, Beltsville, Maryland 20705 USA

P B Kirti Plant Science Department, University of Hyderabad, Hyderabad,

Rodomiro Ortiz Centro Internacional de Mejoramiento de Maiz y Trigo MYT), El Batan, Texcoco, Apdo Postal 6-641, 06600 Mexico, D.F MexicoReid G Palmer United States Department of Agriculture, AgriculturalResearch Service, Corn Insects and Crop Genetics Research Unit, Department

(CIM-of Agronomy, Iowa State University, Ames, Iowa 50011 USA

Linda M Pollak Corn Insects and Crop Genetics Research Unit, United StatesDepartment of Agriculture, Agricultural Research Service, Department ofAgronomy, Iowa State University, Ames, Iowa 50011 USA

Z A Pretorius Department of Plant Sciences, University of the Free State,Bloemfontein, 9300 South Africa

ix

Shyam Prakash National Research Centre on Plant Biotechnology, IndianAgricultural Research Institute, New Delhi 110012 India

C F Quiros Department of Vegetable Crops, University of California, Davis,California 95616 USA

S Ramesh International Crop Research Institute for the Semi-Arid Tropics(ICRISAT), Patancheru 502 324, Andhra Pradesh, India

Belum V S Reddy International Crop Research Institute for the Semi-AridTropics (ICRISAT), Patancheru 502 324, Andhra Pradesh, India

P Sanjana Reddy International Crop Research Institute for the Semi-AridTropics (ICRISAT), Patancheru 502 324, Andhra Pradesh, India

Johannes D Rossouw Monsanto Singapore Co (PTE) Ltd., 151 Lorong Chuan06-08 New Tech Park, Singapore

H D Silva Monsanto Brazil, Rodovia Uberlaˆndia-Araxa´, Uberlandia, MG,Brazil

Philipp W Simon Vegetable Crops Research Unit, United States Department ofAgriculture, Agricultural Research Service, Department of Horticulture, Uni-versity of Wisconsin, Madison, Wisconsin 53706 USA

Vance M Whitaker University of Minnesota, Department of HorticulturalScience, 1970 Folwell Avenue, St Paul, MN 55108 USA

Anthony H D Brown

Dedication: Anthony H D Brown

Conservation Geneticist

Reid G Palmer

United States Department of Agriculture

Agricultural Research Service

Corn Insects and Crop Genetics Research Unit

Department of Agronomy

Iowa State University

Ames, Iowa 50011 USA

B Plant Mating Systems and Population Structure

III THE MAN

IV HONORS AND AWARDS

ACKNOWLEDGMENT

SELECTED PUBLICATIONS OF ANTHONY H D BROWN

This volume of Plant Breeding Reviews is dedicated to Anthony (Tony)

H D Brown, known internationally for his research in conservationand population genetics and plant breeding Dr Brown’s primarycontributions in the area of conservation genetics followed two majorthemes: optimum sampling strategies and core collections His life’s

Plant Breeding Reviews, Volume 31 Edited by Jules Janick

Copyright & 2009 John Wiley & Sons, Inc.

1

activities in this area were inspired primarily by his long friendshipand close working relationship with Sir Otto Frankel His research inpopulation genetics focused on the estimation of mating systems andtheir impact on plant population structure while his research inbreeding was on the use of wild relatives in crop improvement.

Dr Brown started with the Commonwealth Scientific and IndustrialResearch Organization (CSIRO) in 1972 as a research scientist andretired as a chief research scientist in 2006 He is now an HonoraryResearch Fellow in the Centre for Plant Biodiversity Research, CSIROPlant Industry, Canberra, Australia

I BIOGRAPHICAL SKETCH

Tony Brown was born on November 25, 1941, in Waverley, SydneyNSW, Australia It was wartime and his father, Arthur Brown, was inDarwin, Australia, serving as Squadron Leader in the Royal AustralianAir Force Arthur was from three or more generations of Australianstock Tony’s maternal grandfather, Hugh Milligan, son of Scottishimmigrants, was an eminent primary school headmaster Hugh’s taskwas to register Tony’s birth, the agreed name being Anthony Hugh DeanBrown Hugh urged that the last two names be hyphenated becauseplain ‘‘Brown’’ was insufficiently distinguished for a future MacquarieStreet specialist doctor However, Tony’s mother Joyce intervened andsaid, ‘‘Plain Brown is good enough for me, it should be for my son.’’This ensured that name hyphenation could await future needs YetHugh had other major influences on Tony, inspiring a love of plants, ofarithmetic shortcuts, and of parsing sentences That three initials were

an encumbrance emerged later in the United States, where names andofficial forms were triplet coded, allowing only one middle initial Andthe inevitable inversion happened after publications on alcohol dehy-drogenase, when the AHD became ADH, which spawned a growing list

of mutant miscitations in the Science Citation Index

In high school Tony was inspired to seek a research career as anagricultural chemist after watching a film made by CSIRO on how thediscovery of remedying trace element deficiencies in some depleted soils

of South Australia converted them to cropping Consequently, Tonyenrolled in the Faculty of Agriculture, University of Sydney, as a salariedemployee of the Colonial Sugar Refining Company This was an era whencompanies were competing to recruit future graduates At Sydney,inspirational lectures on Mendelian genetics by Professor ‘‘Spinny’’Smith-White led to a final fourth year in genetics Jim Peacock, later tobecome Chief CSIRO Plant Industry, was in his final year as Spinny’s PhD

student He shaped the honors course, and infused enthusiasm as theDNA era was unfolding Jim had spent some time in Adelaide University,and brought back insights from meeting Sir R A Fisher and the team ofpopulation genetics students there Jim devised an unforgettableexperiment for the honors genetics class with 16 blue and 16 yellowplastic beads in a jar to simulate genetic drift theory The jar was shakenand inverted 16 times and the color of the first two beads noted After 16repeats, the jar was opened and its contents adjusted to the new observedgene frequency Over 50 population replications were run over tens ofgenerations, or until fixation blissfully occurred Why was fixationhappening faster than predicted? Sampling with or without replacement?Late into the night the rattle of balls in the jar resounded down the collegecorridors, until crash .extinction: The neck of the jar wore through andbroke However, the experiment had sown the seed of a lifelong interest insampling issues.

On graduating (in 1963), Tony was assigned by the Colonial SugarRefining Company to its sugarcane experiment station in Lautoka, Fiji.This was a major transition, from collegiate to colonial life, and he wasfully briefed at the head office in Sydney on how to behave toward thelocal population The company itself was in transition, hiring local staff

as officers, and the country was preparing for independence Tony’simmediate boss was Joe Daniels, a sugarcane breeder respected aroundthe world and a scholarly and imaginative leader All communicationswere directed through the mill manager, including scientific reports.Tony had an early lesson in communication when management enrolledhim in an in-house training course on report writing The fact thatmanagement chose Tony’s report on fiber content to be one anonymousexample of bad writing firmly made a point It was an object lesson in the

‘‘Gunning fog index,’’ which is a function of the average length ofsentences and the number of words with more than two syllables Theindex is intended to equate to the number of years of education that areader requires to understand the writing Clearly no one in the headoffice had sufficient schooling to read Tony’s report

Joe Daniels introduced Tony to the magnet of wild crop relatives Itmay seem hard to imagine in today’s bottom-line corporate world, butJoe was able to get the company’s approval for a visionary, ambitiousproject entitled ‘‘Re-creating the Pathway of Evolution of Sugarcane’’

by deliberately selecting Saccharum spontaneum for sucrose content

To achieve the screening of large populations for sucrose, Joe andTony installed an autoanalyzer to run around the clock For twoweeks, Tony was on an unforgettable night shift feeding thousands ofethanol extracts into the analyzer and reading the output That and theproject to tag leafhoppers, vectors of Fiji disease, with radiolabel P-32

1 DEDICATION: ANTHONY H D BROWN CONSERVATION GENETICIST 3

to track their movement in cane fields in retrospect seem incongruous

in a small island colony

After nearly four years in Fiji, Tony returned in 1966 to academia andgraduate school He chose to work with Dr Robert W Allard at theUniversity of California, Davis, primarily because of his classic plantbreeding book and his research on quantitative genetics On arrival,however, Tony found that Professor Allard was convinced that the newisozyme technique would open the door to empirical population genetics

in plants Professor Allard recommended that the PhD project should not

be on quantitative genetics but on isozyme variation in Zea mays Thiswould fit better with his assigning Tony as half-time research assistant toimplement an isozyme lab In so many ways, this was a opportunemoment to arrive in Davis and share the excitement and friendship of theAllard lab (particularly Drs M T Clegg, S K Jain, D R Marshall, and B

S Weir) The scientific collaborations begun at Davis continued inprojects for several decades and led later to sabbatical visits at StanfordUniversity and the University of California, Riverside UC Davis was thus

a watershed in Tony’s science and life including marriage

With the completion of his PhD in 1969, Tony was appointed as alecturer in Biology at the University of York, England A seminar byProfessor Warren Ewens in Leeds on the sampling theory of neutralgenes had a lasting influence on Tony’s research After three years, Tonyreturned to Australia to CSIRO Plant Industry as a research scientist inCanberra in 1972 There, two sons, Laurence and Christopher, wereborn At CSIRO, Tony collaborated with Dr Don Marshall, who hadpreceded him from UC Davis, and with Dr Bruce Weir then at Massey,New Zealand One early project was on the charge-state model ofelectrophoretic variation, from which a number of experiences flowed.One experience was to have their joint work scooped by Drs T Ohta and

M Kimura On another occasion, a manuscript by Tony and Don wasbeing subjected to the internal CSIRO editorial process and was sent bythe panel for review to Professor P A P Moran at Australian NationalUniversity Professor Moran was intrigued by the problem andconcerned about some aspects of the existence and convergenceproperties of the distribution He not only submitted his review, butmore important also phoned Tony after hours to discuss this paper Thisled Professor Moran to write a series of theoretical papers, and thisthinking was referred to by Dr J F C Kingman in the history ofcoalescent that he wrote for Genetics in 2000

More influential than the existence of wandering distributions andthe electrophoretic profile was Tony’s work on sampling strategies forplant genetic resources Sir Otto Frankel had challenged Tony and Don

to think critically on this subject Later he promoted their strategy ininternational meetings and used it to challenge conventional collectingpractice, particularly when he thought sampling was excessive In thefield, such theoretical strategies are but a guide, requiring adjustment

to reality This is particularly so for sampling the diversity of wildrelatives, where one is deliberately seeking populations in diversehabitats and of greatly varying size

Along with the excitement of discovering variation new to science inits native setting came the experience of diverse human situations.Tony’s first real germplasm collecting mission was an object lesson inadjusting theory to field reality This was a frenetic mission to Iran withIsraeli professors Dani Zohary and Eibi Nevo The trip went fromMehran near the border with Iraq, across the Zagros Mountains and thesouthern Caspian shores to Gonbad-e-Qabus, just two years before the

1979 revolution With portraits of the shah’s family in every hotel room,the future course of events was not evident At the hotel in Andimesk,the grim faces of the hotel staff were unforgettable as they examined thescientists’ passports Although the target of the trip was wild cereals,particularly wild barley, the diversity being grown by farmers in themany barley fields was inspiring This led to a sampling deliberatelyaimed at testing the allozyme diversity and genetic structure of theselandraces, particularly to see whether the richness of diversity soapparent to the eye was just a mixture of a few genotypes That researchultimately led to Tony’s principal commitment as Honorary ResearchFellow with the International Plant Genetic Resources Institute in theirproject on the significance of crop genetic diversity still present on farms

in traditional agroecosystems (with Drs Toby Hodgkin and DevraJarvis) The research focus was to develop a scientific basis of the useand conservation in situ of this diversity

When are germplasm collections large enough? This question came

to Sir Otto with force after a visit to the excellent large world ricecollection developed by Dr T T Chang at the International RiceResearch Institute Few collections would have the resources to attainthe size and organization of that for rice Furthermore the humanimpulse to collect beyond need would lessen the utility of manycollections Sir Otto therefore proposed that core collections should beset up and called on Tony to read the draft It was stunning to considerthat Otto Frankel, renowned internationally as a champion of crisisgermplasm collecting, should now advocate the slashing of collections

by an order of magnitude After much debate, a compromise concept ofthe ‘‘reserve’’ collection emerged But for years thereafter Sir Otto oftensaid, ‘‘Tony, I have you to blame for the reserve collection,’’ feeling

1 DEDICATION: ANTHONY H D BROWN CONSERVATION GENETICIST 5

that the reserve concept had dulled his original challenge to gene-bankmanagers to prune their holdings.

II RESEARCH ACCOMPLISHMENTS

A Conservation Genetics

Sir Otto Frankel was a member of the FAO Expert Panel on PlantExploration and Introduction, and was preparing for the 1973 FAO/IBPTechnical Conference on Crop Plant Genetic Resources in Rome He feltthat previous papers on plant collecting were strongly biased towardthe practical details of collecting expeditions and that little emphasishad been given to the science of plant exploration The original paperpresented at the technical conference by Drs Don Marshall and TonyBrown entitled ‘‘Optimum Sampling Strategies in Genetic Conserva-tion’’ (subsequently published in the book Crop Genetic Resources forToday and Tomorrow edited by O H Frankel and J G Hawkes) wascontroversial but has since been widely accepted and expanded tocover other issues, such as sampling in biological control programs.Tony followed his early work on sampling strategies by extensivework on developing the concept of core collections This concept, firstintroduced in 1984, was to facilitate the use of genetic resources in themajor crops By the mid-1980s it was felt that many collections,especially in the major crops, had grown so large that their mere sizewas likely to deter their extensive use by individual scientists,breeders, or students, except for a few characters that could be readilyand rapidly discerned on single plants It was proposed that givingpriority in evaluation to a smaller number of accessions would faci-litate greater use of germplasm collections, particularly for a range ofcharacters In a series of papers over the last 20 years, Tony has pro-vided much of the underlying scientific rationale for the establishmentand use of core collections When first introduced, the core collectionconcept, because it challenged accepted dogma, was controversial, but

it now has become widely applied in practice

B Plant Mating Systems and Population Structure

A key theme spanning Tony’s research career has been the ment of plant mating systems and their impact on population variationand structure, and the implications of these differences in terms ofevolution and the collection and conservation of genetic diversity

One of his earliest papers with Professor Allard, which was based onhis PhD research, reported the use of isozyme polymorphisms to esti-mate mating system parameters in open-pollinated maize populations.Over his career Tony developed procedures not only for the estimation

of mating system parameters in both predominantly inbreeding andoutbreeding populations but for also for apomictic species Tony alsoworked with a wide range of colleagues in applying these techniques

in species as diverse as Eucalyptus, Lupinus, wild Hordeum, and anumber of colonizing weed species (with Drs Jeremy Burdon andSpencer Barrett)

Tony’s work on population structure was focused on geneticpolymorphism, heterozygosity, multilocus associations, and populationdifferentiation A theoretical project with Dr Marc Feldman, which isenjoying renewed attention with the burgeoning DNA sequence data,dealt with the measuring and testing of multilocus associations Anotherexample is the analysis of published isozyme data undertaken with Dr.Dan Schoen, which showed that not only do inbreeding and outbreedingspecies differ in overall levels of genetic diversity, but they also differ inthe amount of among population variance of gene diversity Inbreedingspecies exhibited much greater variation in how their populations arestructured than do the populations of individual outbreeding species

We have to be clear that the comparison is the variability between thepopulations of one species; that is, populations 1 and 2 of species A, notpopulation 1 of species A with population 1 of species B

Hordeum Research Another major research theme of Tony’s was thegenetic structure of natural populations of the wild progenitor(Hordeum spontaneum) of cultivated barley and the utilization of thevariation in the wild species in barley improvement His interest inwild barley was sparked in part by his experience at Davis and because

of an invitation by Professor Nevo in 1976 to spend a six-monthsabbatical in the Institute of Evolution, University of Haifa, in Israel.This opened an opportunity for Tony to collect wild barleys in the fieldand use enzyme markers to understand their genetic structure Thefinding of extensive isozyme variation in wild barley raised the issue as

to whether this indicated equally high levels of potentially usefulvariation for barley improvement Screening of accessions collected inIsrael and Iran revealed extensive resistance to barley scald and a range

of other diseases and quality traits Tony developed advancedbackcross lines carrying scald resistance from the wild species,which proved to be a useful tool in investigating the genetics ofseedling resistance and the tagging and pyramiding of resistance genes

1 DEDICATION: ANTHONY H D BROWN CONSERVATION GENETICIST 7

This work led to the release of a commercial cultivar (Tantangara)carrying a known scald resistance gene from wild barley An alliedproject, also conducted with Dr Dave Garvin, was the use of molecularmarkers in breeding adapted proanthocyanidin-free barley.

Glycine Research The legume genus Glycine includes G max(soybean) and its wild progenitor, G soja These annual species arenative to northern Asia and so would seem to have little or nothing to

do with Australia Yet, surprisingly, their closest relatives, and the onlyother members of the genus Glycine, are native to Australia This group

of wild perennial species, Glycine subgenus Glycine, represents thetertiary gene pool for the soybean and is thus of potential economicimportance Collecting and characterizing these perennials has been amajor focus of Tony’s work

The potential of this uniquely Australian resource was recognized atCSIRO by Dr Don Marshall, working initially with Paul Broue` and JimGrace Subsequent staff changes led to Tony taking over the program in

1982 At that time there were fewer than 10 species recognized insubgenus Glycine, but that has changed dramatically In 1982, theInternational Board of Plant Genetic Resources (IBPGR) held a workshop

on soybean genetic resources at Urbana, Illinois, where Tony met thosewho were already, or would become, among the key figures in soybeandiversity research, including Drs Theodore Hymowitz, Reid Palmer,Randy Nelson, Christine Newell, and Duncan Vaughn At the time ofthis workshop, papers on crosses between soybean and perennialGlycine species by the Hymowitz group and by the CSIRO group (Broue`and Marshall) were in draft, and there was tremendous excitement aboutthe potential of the perennials for plant breeding, particularly as sources

of drought- and disease-resistance genes

Tony came to the workshop with rough maps of the Australian Glycinedistributions, based on existing herbarium records This led to proposalsfor germplasm collecting trips to various poorly explored regions ofAustralia As a result, IBPGR funded field trips conducted by Tony tonorth Queensland (1983), the Kimberley district, Western Australia, andNorthern Territory (1984), south Queensland (1985), and easternVictoria (1985) The first two of these trips were co-led by anotherleading figure in perennial Glycine research, Dr Theodore Hymowitz(University of Illinois), with whom Tony led two later trips funded bythe U.S Department of Agriculture, to Western Australia (1993) and tothe central arid zone of the continent (1996) As a result of these trips, thenumber of accessions in the CSIRO perennial soybean germplasmcollection grew from a few hundred in 1982 to over 2,000 at present

It was through Tony’s role as curator of the perennial Glycine seedcollection that he began a longtime collaboration with Dr Jeff Doyle(Cornell University) and his wife, Jane Doyle, when Dr Doyle contactedCSIRO requesting seed for systematic studies in 1982 Tony’s detailedknowledge of Glycine has guided their collaboration, which hasproduced numerous papers on the molecular phylogenetics of thesubgenus The chloroplast phylogeny of Glycine corroborated theexistence of the genome groups that were based on cytological dataamassed by the CSIRO and the University of Illinois groups, and offeredthe first hypothesis of relationships among these groups of species Theavailability of a phylogeny based on defined molecular markers sharedamong all species also allowed the affinities of newly described species

to be determined without recourse to the painstaking studies of

con-ducted by Tony and colleagues in the 1980s and by the group at Illinois.Phylogenies based on nuclear markers subsequently showed someincongruence with chloroplast sequences, and some relationships inGlycine remain unresolved Despite these limitations, molecularsystematic approaches have replaced artificial hybridization as thestandard method for categorizing new species in the subgenus

The field collections made these studies possible and also enabledtraditional taxonomic studies of the genus by other workers, notably Drs.Mary Tindale, Lyn Craven, and Bernard Pfeil Thanks to all of their effortsand the ongoing work of Dr Hymowitz and colleagues, the number ofspecies in the subgenus has grown to 25, including 3 new speciesformally named in 2006 Of the 25, 11 were first found in the field trips led

by Tony Additional species remain to be described in this diverse group

almost exclusively to the Australian continent; one species makes itacross the Torresian Strait to the tip of Papua New Guinea But the

distributed throughout the Pacific, in Timor, the Philippines, NewCaledonia, Vanuatu, Taiwan, and the Ryukya Islands Tony’s work,along with parallel studies by Dr Hymowitz and his group, has beeninstrumental in unraveling the origins of these species, pinpointingtheir diploid progenitors, and elucidating processes such as multipleorigins and lineage recombination, by which these allopolyploidshave become genetically diverse and geographically widespread.With this wealth of data, and the resources developed by years ofcollecting, subgenus Glycine has become a significant model forstudying the process of allopolyploid evolution in plants Tony’sdynamic collaborations on Glycine are a stellar example of cooperation

1 DEDICATION: ANTHONY H D BROWN CONSERVATION GENETICIST 9

to solve biological problems and are a tribute to his congenial andcollegial personality.

In the two and a half decades since Tony assumed responsibility forCSIRO’s perennial Glycine research, he published papers on a number

of other topics, including disease resistance, seed size, floral biology,several on population genetics, and the distribution of calcium oxalatecrystals in Glycine and allies And through all of this, Tony drew on hisGlycine work as a complement to his studies on Hordeum and otherplants, to refine and illustrate his views on germplasm contributions, thearea to which he has dedicated himself for many years

III THE MAN

Dr Brown has a passion for conservation genetics, from his formativeyears with sugarcane to the present with Glycine species Hisadmiration for Otto Frankel, his diligent research at CSIRO, and hisaffiliation with the International Plant Genetic Resources Institute (nowBioversity International), Rome, Italy (1982–present), are evident in hismany contributions CSIRO Plant Industry as his home base has been anexcellent and supportive research environment, where Tony workedjointly with many outstanding colleagues, including Drs JeremyBurdon (who is the current chief of Plant Industry), Curt Brubaker,Andrew Young, Jake Jacobsen, and several others Indeed, thesecharacteristics of Plant Industry owe much to Sir Otto who, as a formerchief of division, instilled a vision of excellence in plant research.The discovery of taxa new to science is the unique reward for thecollector of wild species related to important crops Each of his manytrips had memorable incidents for Tony, and three are mentioned here

If you happened to be one of the few vehicles driving the remote dirtPeninsula ‘‘highway’’ in Cape York, north Queensland, in July of 1983,you may have seen three collectors (Ted Hymowitz from Illinois and JimGrace and Tony from CSIRO) sprawled on the lawn outside theLakeland pub below the billboard saying ‘‘Ice Cold Beer.’’ This was noearly knock off; they actually were sampling rare, tiny Glycinetomentella plants The billboard had nothing to do with site selection;

a collector must check all habitats The roadside pub, a lone building inthe rural landscape, was a haven for the thirsty traveler, and it surrounds

a haven for wild plants that grazing animals would otherwise decimate.Thus, sampling strategies for germplasm collection adapt to reality.Day’s end of another trip found Tony with U.S soybean breeder

Dr Bill Kenworthy, graduate student Michael Doyle, and Jim Grace in

the Carnarvon Gorge of central Queensland, which proved a special site,rich in diverse new taxa Excitement died, however, when halfwayalong the 150-km return trip, it became clear there was insufficient fuel

to reach home at Injune There followed a long silent drive, meeting noother vehicles Despite eking out the last drop of fuel, the vehicle slowed

to a stop 15 km north of their destination Tony and Jim remained withthe vehicle; Bill and Michael chose to jog and walk to town for help,guided by moonlight and the smell of road-killed cattle and kangaroos.During the long wait at the vehicle, the silent darkness was broken

by another vehicle, the first sighted since Carnarvon and, luckily,approaching the road to town, from the property right where the vehiclehad stopped When they apprised the driver of the pickup of theirsituation, he pointed to the rifle hung above his rear window and replied

in a rural Texas accent, ‘‘Just as well you told me If I’d been forced tostop on the road in the dark by two desperados on foot, looking for a ride

to town, I’d answer with this.’’

Meeting the wildlife is a feature of any field trip in Australia A trip tocollect wild Australian Gossypium species, and to evaluate the risks tothem of GM cotton, with botanist Professor Herbert Hurka fromOsnabrueck, Germany, brought them to a remote Corona farm 70 kmnorth of Broken Hill, western NSW Herbert was intrigued by the cagedtalking sulphur-crested cockatoo The farmer’s wife had warned themthat the bird had lived in a hotel in the ‘‘silver city’’ but was banishedbecause of bad language Clearly the garrulous bird enjoyed the attention

of the team of rare visitors, and Herbert lingered to converse with itwhile the CSIRO team sampled When he turned to leave, the bird had afail-safe method to retrieve attention To the visiting professor, itscreeched ‘‘A***hole’’—a fully effective way to grab Herbert’s notice.Sir Otto Frankel was one of the major influences of Tony’s scienceand life His unyielding insistence on high standards and exactness led

to many legendary stories Memorable for Tony was a Christmas Evelunch at which Tony hosted Sir Otto and Lady Margaret, along withProfessor Herbert and Ute Hurka and family members At one point,Tony introduced a wine he was particularly enjoying, and asked whowould like some of this excellent Orlando chardonnay, Otto’s responsewas immediate and emphatic: ‘‘That wine is good, but is certainlyNOT excellent!!’’ Silence fell; then he asked, ‘‘Which vintage?’’Stunned, Tony checked the label and replied, ‘‘1988.’’ Back camethe riposte: ‘‘1987 is better!!’’ Those quips have often proved useful,not only when recalling Otto’s outspokenness

Elements of Tony’s migratory career and strategic sampling interestsare reflected in the lives of his two sons Currently, Laurence is a lecturer

1 DEDICATION: ANTHONY H D BROWN CONSERVATION GENETICIST 11

in migration history in the Department of History, University ofManchester, UK; and Chris is an investment banker with the Mergersand Acquisitions Section of UBS, New York, USA Tony has a newyounger family of three stepchildren who are themselves embarking ondiverse careers.

IV HONORS AND AWARDS

Dr Brown has been extensively recognized for his contributions andachievements to conservation genetics To further broaden his expertise,Tony has been a visiting professor at Stanford University and theUniversity of California, Riverside, visiting research fellow at HaifaUniversity, Israel, and a visiting scientist at the Universitaet Osnabrueck,Germany Tony has excelled in his editorial duties for the journalsGenetics, Molecular Biology and Evolution, and Conservation Genetics aswell as serving as editor or coeditor of 10 books, and conference andsymposia proceedings Of the eight plant collecting missions, Tony hasbeen leader or coleader of six in Australia, one in Israel, and one in Iran.Tony has been the International Plant Genetic Resources Institute (IPGRI)technical advisor and on the organizing committee of 18 internationalworkshops in 10 different countries Perhaps the most rewarding honorwas the award as Honorary Research Fellow by the IPGRI, Rome, Italy.The initial award was in 1997 and Tony has been reappointed three times,most recently with Bioversity International, IPGRI’s new name

ACKNOWLEDGMENT

The authors thank Dr Don Marshall of Plant Breeding Solutions Pty.Ltd., Hamilton, NSW, Australia, for his contributions to the text and forhis critical review of this chapter

SELECTED PUBLICATIONS OF ANTHONY H D BROWN

Brown, A.H.D., J Daniels, and B.D.H Latter 1968 Quantitative genetics of sugarcane I Analysis of variation in a commercial hybrid sugarcane population Theor Appl Genet 38:361–369.

Brown, A.H.D and R.W Allard 1969 Inheritance of isozyme differences among the inbred parents of a reciprocal recurrent selection population of maize Crop Sci 9:72–75.

Brown, A.H.D 1970 The estimation of Wright’s fixation index from genotypic cies Genetica 41:399–406.

frequen-Brown, A.H.D and R.W Allard 1970 Estimation of the mating system in open–pollinated maize populations using isozyme polymorphisms Genetics 66:133–145.

Brown, A.H.D 1971 Isozyme variation under selection in Zea mays Nature 232:570 Brown, A.H.D and R.W Allard 1971 Effect of reciprocal recurrent selection for yield on isozyme polymorphisms in maize (Zea mays L.) Crop Sci 11:888–893.

Marshall, D.R and A.H.D Brown 1973 Stability of performance of mixtures and lines Euphytica 22:405–412.

multi-Brown, A.H.D., D.R Marshall, and L Albrecht 1974 The maintenance of alcohol dehydrogenase polymorphism in Bromus mollis L Aust J Biol Sci 27:545–559 Marshall, D.R and A.H.D Brown 1974 Estimation of the level of apomixis in plant populations Hered 32:321–333.

Brown, A.H.D., A.C Matheson, and K.G Eldridge 1975 Estimation of the mating system

of Eucalyptus obliqua L Herit using allozyme polymorphisms Aust J Bot 23:931–949 Brown, A.H.D 1975 Efficient experimental designs for the estimation of genetic para- meters in plant populations Biometrics 31:145–160.

Brown, A.H.D 1975 Sample sizes required to detect linkage disequilibrium between two

or three loci Theor Pop Biol 8:184–210.

Brown, A.H.D., D.R Marshall, and L Albrecht 1975 Profiles of electrophoretic alleles in natural populations Genet Res Camb 25:137–143.

Brown, A.H.D., D.R Marshall, and B.S Weir 1975 Population differentiation under the charge state model Genetics 81:739–748.

Marshall, D.R and A.H.D Brown 1975 The charge state model of protein polymorphism

in natural populations J Molec Evol 6:149–163.

Marshall, D.R and A.H.D Brown 1975 Optimum sampling strategies in genetic conservation pp 53–80 In: O.H Frankel and J.G Hawkes (eds.), I.B P.2 Crop Genetic Resources for Today and Tomorrow Cambridge Univ Press, Cambridge.

Brown, A.H.D., D.R Marshall, and J Munday 1976 The adaptedness of variants at an alcohol dehydrogenase locus in Bromus mollis L (Soft Bromegrass) Aust J Biol Sci 29:389–396.

Weir, B.S., A.H.D Brown, and D.R Marshall 1976 Testing for selective neutrality of electrophoretically detectable protein polymorphisms Genetics 84:639–659.

Brown, A.H.D., E Nevo, and D Zohary 1977 Association of alleles at esterase loci in wild barley Hordeum spontaneum Nature 268:430–431.

Brown, A.H.D 1978 Isozymes, plant population genetic structure and genetic tion Theor Appl Genet 52:145–157.

conserva-Brown, A.H.D., E Nevo, D Zohary, and O Dagan 1978 Genetic variation in natural populations of wild barley (Hordeum spontaneum) Genetica 49:97–108.

Brown, A.H.D., D Zohary, and E Nevo 1978 Outcrossing rates and heterozygosity in natural populations of Hordeum spontaneum Koch in Israel Hered 41:49–62 Brown, A.H.D 1979 Enzyme polymorphism in plant population Theor Pop Biol 15:1–42.

Nevo, E., D Zohary, A.H.D Brown, and M Haber 1979 Genetic diversity and tal associations of wild barley, Hordeum spontaneum, in Israel Evolution 33:815–833 Doll, H and A.H.D Brown 1979 Hordein variation in wild (Hordeum spontaneum) and cultivated (H vulgare) barley Can J Genet Cytol 21:391–404.

environmen-Brown, A.H.D 1980 Genetic basis of alcohol dehydrogenase polymorphism in Hordeum spontaneum J Hered 70:127–128.

1 DEDICATION: ANTHONY H D BROWN CONSERVATION GENETICIST 13

Brown, A.H.D and L Albrecht 1980 Variable outcrossing and the genetic structure and predominantly self-pollinated species J Theor Biol 82:591–606.

Brown, A.H.D., M.W Feldman, and E Nevo 1980 Multilocus structure of natural lations of Hordeum spontaneum Genetics 96:523–536 Corrigendum May 1981, p 238A Green, A.G., A.H.D Brown, and R.N Oram 1980 Determination of outcrossing rate in a breeding population of Lupinus albus L (White Lupin) Z Pflanzenzuchtg 84:181–191 Brown, A.H.D and M.W Feldman 1981 Population structure of multilocus associations Proc Natl Acad Sci U.S 78:5913–5916.

popu-Brown, A.H.D and D.R Marshall 1981 Evolutionary changes accompanying tion in plants pp 351–363 In: G.G.E Scudder and J.L Reveal (eds.), Evolution Today, Proc Second Int Congr Syst and Evol Biol Univ British Columbia, Vancouver Hunt Institute for Botanical Documentation, Pittsburgh.

coloniza-Marshall, D.R and A.H.D Brown 1981 The evolution of apomixis Hered 47:1–15 Marshall, D.R and A.H.D Brown 1981 Wheat genetic resources pp 21–40 In:

W J Peacock and L.T Evans (eds.), Wheat Science, Today and Tomorrow Cambridge Univ Press, Cambridge.

Brown, A.H.D and J.V Jacobsen 1982 Genetic basis and natural variation of alpha– amylase isozymes in barley Genet Res Camb 40:315–324.

Brown, A.H.D and J Munday 1982 Population genetic structure and optimal sampling of land races of barley from Iran Genetica 58:85–96 Erratum 60:237.

Nevo, E., E Golenberg, A Beiles, A.H.D Brown, and D Zohary 1982 Genetic diversity of environmental associations of wild wheat, Triticum dicoccoides in Israel Theor Appl Genet 62:241–254.

Brown, A.H.D 1983 Barley, pp 57–77 In: S.D Tanksley, and T.J Orton (eds.), Isozymes

in plant genetics and breeding, Part B Elsevier, Amsterdam.

Brown, A.H.D and J.J Burdon 1983 Multilocus diversity in an outbreeding weed, Echium plantagineum L Aust J Biol Sci 36:503–509.

Brown, A.H.D and M.T Clegg 1983 Analysis of variation in related DNA sequences pp 107–

132 In: B.S Weir (ed.), Statistical analysis of DNA sequence data Marcel Dekker, New York Brown, A.H.D and B.S Weir 1983 Measuring genetic variability in plant populations.

pp 219–239 In: S.D Tanksley and T.J Orton (eds.), Isozymes in plant genetics and breeding, Part A Elsevier, Amsterdam.

Burdon, J.J., D.R Marshall, and A.H.D Brown 1983 Demographic and genetic changes in populations of Echium plantagineum L J Ecology 71:667–679.

Brown, A.H.D 1984 Multilocus organization of plant populations pp 159–169 In:

K Wohrmann and V Loeschcke (eds.), Population biology and evolution Springer Verlag, Berlin.

Clegg, M.T., A.H.D Brown, and P.R Whitfeld 1984 Chloroplast DNA diversity in wild and cultivated barley: Implications for genetic conservation Genet Res Camb 43: 339–343 Hanson, A.D and A.H.D Brown 1984 Three alcohol dehydrogenase genes in wild and cultivated barley: characterization of the products of variant alleles Biochem Genet 22:495–515.

Grant, J.E., A.H.D Brown, and J.P Grace 1984 Cytological and isozyme diversity in Glycine tomentella Hayata (Leguminosae) Aust J Bot 32:665–677.

Grant, J.E., J.P Grace, A.H.D Brown, and E Putievsky 1984 Interspecific hybridization in Glycine subgenus Glycine Willd (Leguminosae) Aust J Bot 32:655–663.

Schroeder, H.E and A.H.D Brown 1984 Inheritance of legumin and albumin contents in

a cross between round and wrinkled peas Theor Appl Genet 68:101–107.

Zurawski, G., M.T Clegg, and A.H.D Brown 1984 The nature of nucleotide sequence divergence between barley and maize chloroplast DNA Genetics 106:735–749.

Brown, A.H.D., J.E Grant, J.J Burdon, J.P Grace, and R Pullen 1985 Collection and utilization of wild perennial Glycine pp 345–352 In: R Shibles (ed.), Proc World Soybean Research Conference III Westview Press, Boulder, Colorado.

Doyle, M.J and A.H.D Brown 1985 Numerical analysis of isozyme variation in Glycine tomentella Biochem Syst Ecol 13:413–419.

Brown, A.H.D., J.E Grant, and R Pullen 1986 Outcrossing and paternity in Glycine argyrea by paired fruit analysis Biol J Linn Soc 29:283–294.

Burdon, J.J and A.H.D Brown 1986 Population genetics of Echium plantagineum L.—a target weed for biological control Aust J Biol Sci 39:369–378.

Doyle, M.J., J.E Grant, and A.H.D Brown 1986 Reproductive isolation between isozyme groups of Glycine tomentella (Leguminosae), and spontaneous doubling in their hybrids Aust J Bot 34:523–535.

Grant, J.E., R Pullen, A.H.D Brown, J.P Grace, and P.M Gresshof 1986 Cytogenetic affinity between the new species Glycine argyrea and its congeners J Hered 77: 423–426.

Brown, A.H.D and J.J Burdon 1987 Mating systems and colonizing success in plants pp 115–131 In: A.J Gray, M.J Crawley, and P.J Edwards (eds.), Colonization, succession and stability 26th Symposium of British Ecol Soc Blackwell Scientific, Oxford Henry, R.J and A.H.D Brown 1987 Variation in the carbohydrate composition of wild barley (Hordeum spontaneum) grain Z Pflanzenzu¨chtung 98:97–103.

Hoffman, N.E., D Hondred, A.D Hanson, and A.H.D Brown 1988 Lactate ase isozymes in barley: Polymorphism and genetic basis J Hered 79:110–114 Brown, A.H.D., J Munday, and R.N Oram 1988 Use of isozyme-marked segments from wild barley (Hordeum spontaneum) in barley breeding Plant Breed 100:280–288 Brown, A.H.D 1989 The case for core collections pp 136–156 In: A.H.D Brown, O.H Frankel, D.R Marshall, and T Williams (eds.), The use of plant genetic resources Cambridge Univ Press, Cambridge.

dehydrogen-Brown, A.H.D 1989 Core collections: A practical approach to genetic resources ment Genome 31:818–824.

manage-Brown, A.H.D 1989 Genetic characterization of plant mating systems pp 145–162 In: A.H.D Brown, M.T Clegg, A.L Kahler, and B.S Weir (eds.), Plant population genetics, breeding and genetic resources Sinaeuer Associates, Sunderland, Massachusetts Brown, A.H.D., J.J Burdon, and A.M Jarosz 1989 Isozyme analysis of plant mating systems pp 73–86 In: D Soltis and P Soltis (eds.), Isozymes in plant biology Dioscorides Press, Portland, Oregon.

Brown, A.H.D., G.J Lawrence, M Jenkin, J Douglass, and E Gregory 1989 Linkage drag

in backcross breeding J Hered 80:234–239.

Doyle, J.J and A.H.D Brown 1989 5S nuclear ribosomal gene variation in the Glycine tomentella polyploid complex Syst Bot 14:398–407.

Hurka, H., S Freunder, A.H.D Brown, and U Plantholt 1989 Aspartate amino transferase isozymes in the genus Capsella (Brassicaceae): Subcellular location, gene duplication and polymorphism Biochem Genetics 27:77–90.

Kenworthy, W.J., A.H.D Brown, and G.A Thibou 1989 Variation in flowering response

to photoperiod in perennial Glycine species Crop Sci 29:678–682.

Brown, A.H.D 1990 The role of isozyme studies in molecular systematics Aust Syst Bot 3:39–46.

Brown, A.H.D., J.J Burdon and J.P Grace 1990 Genetic structure of Glycine canescens, a perennial relative of soybean Theor Appl Genet 79:729–736.

Doyle, J.J., J.L Doyle, and A.H.D Brown 1990 Analysis of a polyploid complex in Glycine with chloroplast and nuclear DNA Aust Syst Bot 3:125–136.

1 DEDICATION: ANTHONY H D BROWN CONSERVATION GENETICIST 15

Doyle, J.J., J.L Doyle, and A.H.D Brown 1990 A chloroplast DNA phylogeny of the wild perennial relatives of soybean (Glycine subgenus Glycine): Congruence with morpho- logical and crossing groups Evolution 44:371–389.

Doyle, J.J., J.L Doyle, and A.H.D Brown 1990 Chloroplast DNA phylogenetic affinities of newly described species in Glycine (Leguminosae: Phaseoleae) Syst Bot 15:466–471 Doyle, J.J., J.L Doyle, and A.H.D Brown 1990 Chloroplast DNA polymorphism and phylogeny in the B genome of Glycine subgenus Glycine (Leguminosae) Amer J Botany 77:772–782.

Doyle, J.J., J.L Doyle, A.H.D Brown, and J.P Grace 1990 Multiple origins of polyploids in the Glycine tabacina complex inferred from chloroplast DNA polymorphism Proc Natl Acad Sci USA 87:714–717.

Doyle, J.J., J.L Doyle, J.P Grace, and A.H.D Brown 1990 Reproductively isolated polyploid races of Glycine tabacina (Leguminosae) had different chloroplast genome donors Syst Bot 15:173–181.

Feuerstein, U., A.H.D Brown, and J.J Burdon 1990 Linkage of rust resistance genes from wild barley (Hordeum spontaneum) with isozyme markers Plant Breed 104:318–324 Schoen, D.J and A.H.D Brown 1991 Intraspecific variation in population gene diversity and effective population size correlates with the mating system in plants Proc Natl Acad Sci USA 88:4494–4497.

Abbott, D.C., J.J Burdon, A.M Jarosz, A.H.D Brown, W.J Muller, and B.J Read 1991 The relationship between seedling infection types and field reactions to leaf scald in Clipper barley backcross lines Aust J Agric Res 42:801–809.

Brown, A.H.D and J.D Briggs 1991 Sampling strategies for genetic variation in ex situ collections of endangered plant species pp 99–119 In: D.A Falk and K.E Holsinger (eds.), Genetics and Conservation of Rare Plants Oxford Univ Press, Oxford Lagudah, E.S., R Appels, A.H.D Brown, and D McNeil 1991 The molecular-genetic analy- sis of Triticum tauschii, the D-genome donor to hexaploid wheat Genome 34: 375–386 MacLeod, L.C., R.C.M Lance, and A.H.D Brown 1991 Chromosomal mapping of the Glb

1 locus encoding (1 !3), (1!4)–ß–D–glucan 4–glucanohydrolase EI in barley J Cereal Sci 13:291–298.

Schoen, D.J and A.H.D Brown 1991 Whole and part-flower self-pollination in Glycine clandestina and G argyrea and the evolution of autogamy Evolution 45:1651–1664.

Abbott, D.C., A.H.D Brown, and J.J Burdon 1992 Genes for scald resistance from wild barley (Hordeum vulgare ssp spontaneum) and their linkage to isozyme markers Euphytica 61:225–231.

Brown, A.H.D 1992 Genetic variation and resources in cultivated barley and wild Hordeum Barley Genetics 6:669–682.

Brown, A.H.D 1992 Human impact on plant gene pools and sampling for their servation Oikos 63:109–118.

con-Schoen, D.J., J.J Burdon, and A.H.D Brown 1992 Resistance of Glycine tomentella to soybean leaf rust Phakopsora pachyrhizi in relation to ploidy level and geographic distribution Theor Appl Genet 83:827–832.

Schoen, D.J and A.H.D Brown 1993 Conservation of allelic richness in wild crop relatives is aided by assessment of genetic markers Proc Natl Acad Sci USA 90:10623–10627.

Brown, A.H.D and D.J Schoen 1994 A revised measure of association of gene diversity values Hereditas 120:77–79.

Burdon, J.J., D.C Abbott, A.H.D Brown, and J.S Brown 1994 Genetic structure of the scald pathogen (Rhynchosporium secalis) in South East Australia: Implications for control strategies Aust J Agric Res 45:1445–1454.

Guerin, J.R., R.C.M Lance, A.H.D Brown, and D.C Abbott 1994 Mapping of malt endopeptidase, diaphorase and esterase loci on barley chromosome 3L Plant Breed 112:279–284.

Prober, S.M and A.H.D Brown 1994 Conservation of the grassy white box woodlands I Population genetics and fragmentation of Eucalyptus albens Benth Conservation Biol 8:1003–1013.

Abbott, D.C., E.S Lagudah, and A.H.D Brown 1995 Identification of RFLPs flanking a scald resistance gene on barley chromosome 6 J Hered 86:152–154.

Brown, A.H.D 1995 The core collection at the crossroads pp 3–19 In: T Hodgkin, A.H.D Brown, T.J.L van Hintum, and E.A.V Morales (eds.), Core collections of plant genetic resources John Wiley, Chichester.

Brown, A.H.D and D.R Marshall 1995 A basic sampling strategy: Theory and practice.

pp 75–91 In: L Guarino, V Ramanatha Rao, and R Reid (eds.), Collecting plant genetic diversity technical guidelines CAB International, Wallingford.

Frankel, O.H., A.H.D Brown, and J.J Burdon 1995 The conservation of plant diversity Cambridge Univ Press, Cambridge.

bio-Brown, A.H.D., D.F Garvin, J.J Burdon, D.C Abbott, and B.J Read 1996 The effect of combining scald resistance genes on disease levels, yield and quality traits in barley Theor Appl Genet 93:361–366.

Young, A., G.T Boyle, and A.H.D Brown 1996 The population genetic consequences of habitat fragmentation for plants Trends in Ecology and Evolution 11:413–418 Young, A.G and A.H.D Brown 1996 Comparative population genetic structure on the rare woodland shrub Daviesia suaveolens and its common congener D mimosoides Conservation Biol 10:1220–1228.

Brown, A.H.D., C.L Brubaker, and J.P Grace 1997 The regeneration of germplasm samples: Wild versus cultivated species Crop Sci 37:7–13.

Brown, A.H.D., C.L Brubaker, and M.J Kilby 1997 Assessing the risk of cotton transgene escape into wild Australian Gossypium species pp 83–94 In: G.D McLean, P.M Waterhouse, G Evans, and M.I Gibbs (eds.), The commercialisation of transgenic crops: Risk, benefit and trade considerations Bureau of Resource Sciences, Kingston, ACT, Australia.

Garvin, D.F., A.H.D Brown, and J.J Burdon 1997 Inheritance and chromosome locations

of novel scald resistance genes derived from Iranian and Turkish wild barleys Theor Appl Genet 94:1087–1091.

Roulin, S., P Xu, A.H.D Brown, and G.B Fincher 1997 Expression of specific (1 Glucanase genes in leaves of near-isogenic resistant and susceptible barley lines infected with the leaf scald fungus (Rhynchosporium secalis) Phys Mol Plant Path 50:245–261 Garvin, D.F., J.E Miller-Garvin, E.A Viccars, J.V Jacobsen, and A.H.D Brown 1998 Identification of molecular markers linked to ant28, a mutation that eliminates proanthocyanidin in barley seeds Crop Sci 38:1250–1255.

!3)-b-Prober, S.M., L.H Spindler, and A.H.D Brown 1998 Conservation of the grassy white box woodlands: Effects of remnant population size on genetic diversity of the outcrossing, allotetraploid herb, Microseris lanceolata Conservation Biol 12:1279–1290.

Young, A.G and A.H.D Brown 1998 Comparative analysis of mating systems in the rare woodland shrub Daviesia suaveolens and its congener D mimosoides Hered 80: 374–381 Brown, A.H.D 1999 The genetic structure of crop landraces and the challenge to conserve them in situ on farms pp 29–48 In: S.B Brush (ed.), Genes in the field: Conserving plant diversity on farms Lewis Publishers, Boca Raton, FL.

Brubaker, C.L., A.H.D Brown, J McD McStewart, M.J Kilby, and J.P Grace 1999 Production of fertile hybrid germplasm with diploid Australian Gossypium species for cotton improvement Euphytica 108:199–213.

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Burdon, J.J.P.H Thrall, and A.H.D Brown 1999 Resistance and virulence structure in two Linum marginale—Melampsora lini host-pathogen metapopulations with different mating systems Evolution 53:704–716.

Doyle, J.J., J.L Doyle, and A.H.D Brown 1999 Incongruence in the diploid B-genome species complex of Glycine (Leguminosae) revisited: Histone H3-D alleles vs chlor- oplast haplotypes Molec Biol Evol 16:354–362.

Doyle, J.J., J.L Doyle, and A.H.D Brown 1999 Origins, colonization, and lineage recombination in a widespread perennial soybean polyploid complex Proc Nat Acad Sci USA 96:10741–10745.

Marshall, D.R and A.H.D Brown 1999 Sampling wild legume populations pp 78–89 In: S.J Bennett and P.S Cocks (eds), Genetic resources of Mediterranean pasture and forage legumes Kluwer Acad Press, Dordrecht.

Young, A.G., A.H.D Brown, and F.C Zich 1999 Genetic structure of fragmented populations of the endangered grassland daisy Rutidosis leptorrhynchoides Conserva- tion Biol 13:256–265.

Young, A.G and A.H.D Brown 1999 Paternal bottlenecks in fragmented populations of the grassland daisy Rutidosis leptorrhynchoides Genet Res 73:111–117.

Abbott, D.C., J.J Burdon, A.H.D Brown, B.J Read, and D Bittisnich 2000 The incidence

of barley scald in cultivar mixtures Aust J Agric Res 51:355–360.

Brown, A.H.D and C.L Brubaker 2000 Genetics and the conservation and use of Australian wild relatives of crops Aust J Bot 48:297–303.

Brown, A.H.D and C.M Hardner 2000 Sampling the gene pools of forest trees for

ex situ conservation pp 185–196 In: A Young, T Boyle, and D Boshier (eds.), Forest conservation genetics: Principles and practice CSIRO, Melbourne, Australia.

Brown, A.H.D and A.G Young 2000 Genetic diversity in tetraploid populations of the endangered daisy Rutidosis leptorrhynchoides and implications for its conservation Hered 85:122–129.

Doyle, J.J., J.L Doyle, A.H.D Brown, and B.L Pfeil 2000 Confirmation of shared and divergent genomes in the Glycine tabacina polyploid complex (Leguminosae) using histone H3-D sequences Syst Bot 25:437–448.

Garvin, D.F., A.H.D Brown, H Raman, and B.J Read 2000 Genetic mapping of the barley Rrs14 scald resistance gene with RFLP, isozyme and seed storage protein markers Plant Breed 119:193–196.

Brown, A.H.D and C.L Brubaker 2001 Indicators for sustainable management of plant genetic resources—how well are we doing? pp 249–262 In: J.M.M Engels,

V Ramanatha Rao, A.H.D Brown, and M T Jackson (eds.), Managing plant genetic diversity CAB International, Wallingford, Oxon, UK.

Lin, J.-Z., A.H.D Brown, and M.T Clegg 2001 Heterogeneous geographic patterns of nucleotide sequence diversity between two alcohol dehydrogenase genes in wild barley (Hordeum vulgare ssp spontaneum) Proc Nat Acad Sci USA 98:531–536.

Teshome, A., A.H.D Brown, and T Hodgkin 2001 Diversity in landraces of cereal and legume crops Plant Breed Rev 21:221–261.

Brown, A.H.D., J.L Doyle, J.P Grace, and J.J Doyle 2002 Molecular phylogenetic relationships within and among diploid races of Glycine tomentella (Leguminosae) Aust Syst Bot 15:37–47.

Bleeker, W., A Franzke, K Pollman, A.H.D Brown, and H Hurka 2002 Phylogeny and biogeography of southern hemisphere high mountain Cardamine species (Brassica- ceae) Aust Syst Bot 15:575–581.

Doyle, J.J., J.L Doyle, A.H.D Brown, and R.G Palmer 2002 Genomes, multiple origins, and lineage recombination in the Glycine tomentella (Leguminosae) polyploid complex: histone H3-D gene sequences Evolution 56:1388–1402.

Rauscher, J.T., J.J Doyle, and A.H.D Brown 2002 Internal transcribed spacer repeat– specific primers and the analysis of hybridization in the Glycine tomentella (Legumi- nosae) polyploid complex Molec Ecol 11:2691–2702.

Brubaker, C.L and A.H.D Brown 2003 The use of multiple alien chromosome addition aneuploids facilitates genetic linkage mapping of the Gossypium G genome Genome 46:774–791.

Genger, R.K., A.H.D Brown, W Knogge, K Nesbitt, and J.J Burdon 2003 Development of SCAR markers linked to a scald resistance gene derived from wild barley Euphytica 134:149–159.

Genger, R.K., K.J Williams, H Raman, B.J Read, H Wallwork, J.J Burdon, and A.H.D Brown.

2003 Leaf scald resistance genes in Hordeum vulgare and Hordeum vulgare ssp neum: parallels between cultivated and wild barley Aust J Agric Res 54:1335–1342 Doyle, J.J., J.L Doyle, J.T Rauscher, and A.H.D Brown 2003 Diploid and polyploidy reticulate evolution throughout the history of the perennial soybeans (Glycine subg Glycine) New Phytologist 161:121–132.

sponta-Murray, B.R., A.H.D Brown, and J.P Grace 2003 Geographic gradients in seed size among and within perennial Australian Glycine species Aust J Bot 51:47–56 Rau, D., A.H.D Brown, C.L Brubaker, G Attene, V Balmas, E Saba, and R Papa 2003 Population genetic structure of Pyrenophora teres Drechs., the causal agent of net blotch

in Sardinian landraces of barley complex (Hordeum vulgare L.) Theor Appl Genet 106:947–959.

Doyle, J.J., J.L Doyle, J.T Rauscher, and A.H.D Brown 2004 Evolution of the perennial soybean polyploid (Glycine subgenus Glycine): A study of contrasts Biol J Linnean Soc 82:583–597.

Joly, S., J.T Rauscher, S.L Sherman-Broyles, A.H.D Brown, and J.J Doyle 2004 tion of the 18S-5.8S-26S nuclear ribosomal gene family and its expression in natural and artificial Glycine allopolyploids Molec Biol Evol 21:1409–1421.

Evolu-Murray, B.R., A.H.D Brown, C.R Dickman, and M.S Crowther 2004 Geographical gradients in seed mass in relation to climate.J Biogeography 31:379–388.

Rauscher, J.T., J.J Doyle, and A.H.D Brown 2004 Multiple origins and nrDNA internal transcribed spacer homoeologue evolution in the Glycine tomentella (Leguminosae) allopolyploid complex Genetics 166:987–998.

Cervantes-Martinez, T., H.T Horner, R.G Palmer, T Hymowitz, and A.H.D Brown.

2005 Calcium oxalate crystal macropatterns in leaves of species from groups Glycine and Shuteria (Glycininae; Phaseoleae; Papilionoideae; Fabaceae) Can J Bot 83:1410–1421

Genger, R.K., K Nesbitt, A.H.D Brown, D.C Abbott, and J.J Burdon 2005 A novel barley scald resistance locus: Genetic mapping of the Rrs15 scald resistance gene derived from wild barley, Hordeum vulgare ssp spontaneum Plant Breed 124: 137–141.

Rau, D., F.J Maier, R Papa, A.H.D Brown, V Balmas, E Saba, W Shafer, and G Attene,

2005 Isolation and characterization of the mating-type locus of the barley pathogen Pyrenophora teres frequencies of mating-type idiomorphs within and among fungal populations collected from barley landraces Genome 48:855–869.

Pfeil, B.E., L.A Craven, A.H.D Brown, B.G Murray, and J.J Doyle 2006 Three new species of northern Australian Glycine (Fabaceae, Phaseolae) G-gracei, G montis-dougle and G syndetika Aust Syst Bot 19:245–258.

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Brown, A.H.D and T Hodgkin 2007 Measuring, managing and maintaining crop genetic diversity on-farm pp 13–33 In: D Jarvis, C Paddoch, and D Williams (eds.), Managing biodiversity in agricultural ecosystems Columbia University Press, New York Jarvis, D.I., A.H.D Brown, V.I Imbruce, J Ochoa, M Sadiki, E Karamura, P Trutmann, and M.R Finckh 2007 Managing crop disease in traditional agroecosystems: The benefits and hazards of genetic diversity pp 292–319 In: D Jarvis, C Paddoch, and D Williams (eds.), Managing biodiversity in agricultural ecosystems Columbia University Press, New York.

Triono, T., M.D Crisp, A.H.D, Brown, and J.G West 2007 A phylogency of Pouteria (Sapotaceae) from Malesia and Australasia Aust Syst Bot 20:107–118.

Rau, D., G Attene, A.H.D Brown, L Nanni, F.J Maier, V Balmas, E Saba, W Schaefer, and R Papa 2007 Phylogeny and evolution of mating-type genes from Pyrenophora teres, the causal agent of barley ‘‘net blotch’’ disease Current Genetics 51:377–392 Jarvis, D.I., A.H.D Brown, et al 2008 A global perspective of the richness and evenness of traditional crop-variety diversity maintained by farming communities Proc Nat Acad Sci USA 105:5326–5331.

Brassica and Its Close Allies:

Cytogenetics and Evolution

Shyam Prakash

National Research Centre on Plant Biotechnology

Indian Agricultural Research Institute

New Delhi 110012 India

S R Bhat

National Research Centre on Plant Biotechnology

Indian Agricultural Research Institute

New Delhi 110012 India

National Research Centre on Plant Biotechnology

Indian Agricultural Research Institute

New Delhi 110012 India

I INTRODUCTION

II CYTOGENETICS

A Cytogenetic Architecture of Brassica Coenospecies

B Crop Species

Plant Breeding Reviews, Volume 31 Edited by Jules Janick

Copyright & 2009 John Wiley & Sons, Inc.

21

1 Nature of Diploid Species

2 Nature of Alloploid Species

3 Nuclear DNA

4 Karyotypes

5 Pachytene Chromosomes

6 Satellite Chromosomes and rDNA Loci

7 Archetype and Evolution of Genomes

III GENOME MANIPULATION

A Resyntheses of Natural Allopolyploid Brassica spp.

B Agronomic Potential of Synthetics

C Diploidization of Allopolyploid Species

D Raphanobrassica

E Higher Allopolyploids in U Triangle Species through Protoplast Fusion

IV WIDE HYBRIDIZATION

A Sexual Hybrids

B Somatic Hybrids

C Introgression of Genes

V CYTOPLASMIC SUBSTITUTION AND MALE STERILITY

VI GENOME DISSECTION AND DEVELOPMENT OF CHROMOSOME

VIII PLASTID GENOME

IX POTENTIAL ROLE OF ARABIDOPSIS THALIANA IN BRASSICA IMPROVEMENT

A A thaliana as a Model Crucifer

B Cytology and Possible Origin of the A thaliana Genome

C Synteny Conservation

D Synteny-Based Gene Discovery and Cloning

E Arabidopsis Knowledge–Based Gene Discovery and Brassica Improvement

1 Understanding Domestication

2 Understanding Metabolism

3 Testing for Gene Function by Complementary Transformation

X CHLOROPLAST GENOMES AND THEIR PHYLOGENETIC IMPLICATIONS

XI EVOLUTION OF MORPHOLOGICAL CHARACTERS

2 BRASSICA AND ITS CLOSE ALLIES: CYTOGENETICS AND EVOLUTION 23

RAPD Randomly amplified polymorphic DNA

com-of crop species In fact, genetic enrichment com-of crop species with genesfrom wild allies is a major approach for many crop improvementprograms Such gene transfer can be achieved both by conventionalplant breeding methods and through biotechnology

Cytogenetic investigations on Brassica, initiated in the second decade

of the 20th century, were confined to determining the chromosomenumbers and studying chromosome pairing in interspecific hybrids,which finally led to unraveling the genetic architecture of crop species.Small size of chromosomes, scarcity of distinctive cytogenetic landmarks,

24 S PRAKASH, S R BHAT, C F QUIROS, P B KIRTI, AND V L CHOPRA

and not being amenable to pachytene investigations were majordeterrants to cytogenetical analyses Advances in tissue culture techni-ques, including ovary and embryo rescue and protoplast fusion, since the1950s made varied cytogenetic material available to investigate genomehomologies and facilitated introgression of useful nuclear genes evenacross conventional generic boundaries Such investigations requirereliable markers for chromosome identification A significant step towardthis development has been the extensive use of molecular markers.Molecular biology in Brassica started with the determination of femaleparents of allopolyploid species using chloroplast DNA RFLPs by Palmer

et al (1983a) Use of genomic and fluorescence in situ hybridization(GISH and FISH respectively) methodology, in combination withribosomal DNA markers have given new directions in genome analysis

Table 2.1 Taxonomic components of Brassica and related genera and their usage.

B nigra black mustard condiment (seed)

var alboglabra Chinese kale vegetable (stem, leaves)

B rapa

spp oleifera turnip rape oilseed

var brown sarson brown sarson oilseed

var yellow sarson yellow sarson oilseed

ssp rapifera turnip fodder, vegetable (root) ssp chinensis bok choi vegetable (leaves)

ssp pekinensis Chinese cabbage vegetable, fodder (head) ssp nipposinica — vegetable (leaves)

ssp parachinensis — vegetable (leaves)

B carinata Ethiopian mustard vegetable, oilseed

B juncea mustard oilseed, vegetable

B napus

spp oleifera rapeseed oilseed

spp rapifera rutabaga, swede fodder

Eruca sativa rocket, taramira vegetable, nonedible oilseed Raphanus sativus radish vegetable, fodder

Sinapis alba white mustard oilseed

2 BRASSICA AND ITS CLOSE ALLIES: CYTOGENETICS AND EVOLUTION 25