Biologia Del Maiz

8/13/2019 Biologia Del Maiz

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The Biology of Zea mays L. ssp mays (maize or corn) Office of the Gene Technology Regulator

The Biology of

Zea maysL. ssp mays

(maize or corn)

Version 1: September 2008

This document provides an overview of baseline biological information relevant to riskassessment of genetically modified forms of the species that may be released into theustralian environment!


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For information on the Australian Government Office of the Gene Technology egulator visit

!http"##$$$.ogtr.gov.au%


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TABLE OF CONTENTS

"#$%&'$ !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! !!!!!!!! !!

Section 1 Ta(onomy !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

Section 2 )rigin and *ultivation !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

2!1 *entre of diversity and domestication !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!2!2 *ommercial uses !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

2!2!1 %ai+e types and their uses !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!2!2!2 "rocessing of grain mai+e !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!2!2!, -orld mai+e production !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

2!, *ultivation in ustralia !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!2!,!1 *ommercial propagation !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!2!,!2 Scale of cultivation !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!2!,!, *ultivation practices !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

2!. *rop /mprovement !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!2!.!1 &reeding !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!2!.!2 enetic modification !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

Section , %orphology !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

,!1 "lant morphology !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!,!2 #eproductive morphology !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

Section . evelopment !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

.!1 #eproduction !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!.!1!1 se(ual reproduction !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!.!1!2 Se(ual reproduction !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

.!2 "ollen dispersal pollination and outcrossing rates !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!.!2!1 "ollen !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!.!2!2 "ollen dispersal and pollination !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!.!2!, )utcrossing rates and isolation distances !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

.!, $mbryogenesis fruit3seed development and seed dispersal !!!!!!!!!!!!!!!!!!!!.!,!1 $mbryogenesis !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!.!,!2 4ruit3seed development !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!.!,!, Seed dispersal !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

.!. Seed dormancy and germination !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

.!5 Vegetative growth !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Section 5 &iochemistry !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

5!1 6utrient components of the mai+e kernel !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!5!1!1 Starch !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!5!1!2 "rotein !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!5!1!, 'ipids !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

5!2 To(ins !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!5!2!1 6itrate poisoning !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

5!, llergens !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!5!. )ther undesirable phytochemicals !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

5!5 &eneficial phytochemicals !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!Section 7 biotic /nteractions !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

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7!1 6utrient reuirements !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!7!2 Temperature reuirements and tolerances !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!7!, -ater !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!7!. )ther abiotic stresses !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

Section 9 &iotic /nteractions !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

9!1 -eeds !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!9!2 "ests and diseases !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

9!2!1 /nsects and other invertebrate pests !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!9!2!2 Vertebrate pests !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!9!2!, iseases !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!9!2!. )ther adverse associations with mai+e !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

9!, )ther biotic interactions !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

Section 8 -eediness !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

8!1 -eediness status on a global scale !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

8!2 -eediness status in ustralia !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!8!5 *ontrol measures !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

Section "otential for Vertical ene Transfer !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

!1 6atural intraspecific and interspecific crossing !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!2 *rossing under e(perimental conditions !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

!2!1 /nterspecific crosses !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!2!2 /ntergeneric crosses !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

REFERENCES ...........................................................................................................................................


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PREABLE

This document describes the biology ofZea mays'! subspecies ;ssp!< mays withparticular reference to the ustralian environment cultivation and use! /nformationincluded relates to the ta(onomy and origins of cultivatedZ. maysssp! mays generaldescriptions of its morphology reproductive biology biochemistry and biotic andabiotic interactions! This document also addresses the potential for gene transfer tooccur to closely related species! The purpose of this document is to provide baselineinformation about the parent organism in risk assessments of genetically modified

Z. maysssp! maysthat may be released into the ustralian environment!

s mai+e is one of the best researched and characterised plants significant amounts ofinformation are available for many aspects of the biology of mai+e and the reader isreferred to the literature provided in this document as a starting point!

/n this document the terms mai+e and corn are used to refer toZ. maysssp! mays.)ther subspecies ofZea maysare referred to as teosintes!

%ai+e is an annual grass growing up to . m tall! The female inflorescences the earsdevelop in leaf a(ils on the stalk which terminates in the male inflorescence thetassel! The broad leaf sheaths are overlapping around the stalk and the leaves arearranged in two opposing rows along the stalk! %ai+e has a multitude of uses and isused in the preparation of food or drinks as animal feed or for industrial purposes!

SECT!"N# TA$"N"%

The genusZea belongs to the tribe ndropogoneae in the subfamily "anicoideae inthe family "oaceae ;reviewed in )$* 200,= >S 2005S 2005

Table 1 Zeaspecies and subspecies*

Species Chromosomenumber

Subspecies Synonyms

1.

Zea diploperennisHH Iltis et al

2n= 20 - -

2.

Zea luxurians(Durieu & Asch.)RM Bird

2n= 20 - Euchlaena luxuriansDurieu & Asch.

Zea mayssspluxurians(Durieu &Asch.) HH Iltis

3.

Zea maysL. 2n= 20 Zea maysssphuehuetenangensis (HHIltis & De!le") De!le"

Zea maysssp mays Zea curaguaMli#aZea indentata$turte%.

5


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Species Chromosomenumber

Subspecies Synonyms

Zea indurata$turte%.

Zea japonicaa#Hutte

Zea maysc% albaAle'.

Zea maysc% leucodonAle'.

Zea mays varflavorubra

Zea mays%ari#de#tata ($turte%.) LHBaile"

Zea mays var indurata($turte%.) LH Baile"

Zea mays %arjaponica

(a# Hutte) Alph.d

Zea mays %arsaccharata ($turte%.)LH Baile"

Zea mays %artunicataLarraa*a e+ A.$t.-Hil.

Zea mays %arvulgate,er#. & H er#er

Zea saccharate$turte%.

Zea maysssp mexicana($chrad.) HH Iltis Euchlaena mexicana$chrad.

Zea mexicana($chrad.) ,u#te

Zea mayssspparviglumis HH Iltis &De!le"

Zea mays %arparviglumis

.

Zea nicaraguensisHH Iltis & B/ Be#

2n= - -

.

Zea perennis(Hitchc.) Ree%es& Ma#*elsd.

2n= 0 - Euchlaena perennisHitchc.

$urce 4$DA (200)

Z. maysassp! maysis the only cultivated species= the other species and subspecies arewild grasses referred to as teosintes!

/n addition to the basic chromosome complement mai+e plants may contain one ormore supernumerary chromosomes called & chromosomes which do not pair with chromosomes during meiosis ;as reviewed in Aones et al! 2008

a The nameZeais from the reekzeameaning cereal or grain! The specific epithet maysis thought to

derive from the native rawak word mai+ or mahi+ used in the mericas to describe the plant= theword was adopted by the Spanish crew of *olumbusB first voyage who first collected the grain from themericas and took it to $urope ;Cyam D "ankhurst 15= esEardins D %c*arthy 200.


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The mai+e genome is large being somewhere between 2!, F 2!9 bpb

;rumuganathan D $arle 11and retrotransposons ;Cake D -albot 180= 'iu et al! 2009

SECT!"N& "R!'!NANCLT!*AT!"N

&.# Cen+re of ,i-ersi+y an, ,omes+ica+ion

The centre of origin of mai+e is the %esoamerican region probably in the %e(icanhighlands from where it spread rapidly! rchaeological records and phylogeneticanalysis suggest that domestication began at least 7000 years ago ;"iperno D4lannery 2001= %atsuoka et al! 2002

%ai+e can be grown in a number of environments ;reviewed in "aliwal 2000b=4arnham et al! 200,< from 58G 6orth ;eg *anada and the #ussian 4ederation< to .0GSouth ;eg *hile

b The amount of 6 in the nucleus of a eukaryotic cell is e(pressed as the total number of base pairs;bp< in a haploid ;1*< chromosome complement!

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http://www.maizegenome.org/http://www.maizegenome.org/


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&.& Commercial ses

%ai+e is one of the oldest cultivated grains and one of the most productive cropspecies with a global average yield of more than . tonnes per hectare ;reviewed in"aliwal 2000b= 4arnham et al! 200,

/t can be directly consumed as food at various developmental stages from baby cornto mature grain!

high proportion of mai+e produced is used as stock feed eg .0H in tropical areasand up to 85H in developed countries ;reviewed in "aliwal 2000g= 4arnham et al!200,

%ai+e can be processed for a range of uses both as an ingredient in food or drinks egcorn syrup in soft drinks or mai+e meal or for industrial purposes! %ai+e is the maEorsource of starch world wide and is used as a food ingredient either in its native formor chemically modified ;-hite 1.

production and are used in food manufacturing ;&oyer D Cannah 1.= "aliwal2000h= Cobbs 200,= %c*utcheon 2009

&.&.# aize +ypes an, +heir ses

number of mai+e types can be discerned on the basis of endosperm and kernelcomposition ;"urseglove 192= "aliwal 2000c= arrah et al! 200,

4lint mai+e kernels are characterised by their high percentage of hard endosperm ;see4igure ,< around a small soft centre! 4lint mai+e is grown predominantly in 'atinmerica and $urope for food use!

ent mai+e is the most commonly grown for grain and silage and is the predominanttype grown in the >S! Card endosperm is present on the sides and base of thekernel! The remainder of the kernel is filled with soft starch= when the grain startsdrying the soft starch at the top of the kernel contracts producing the depression for

which it is named!

4loury mai+e is being grown predominantly in the ndean region! /ts endosperm ismainly composed of soft starch making it easy to grind and process into foods!

-a(y mai+e kernels contain almost entirely amylopectin as their starch ;rather thanthe normal 90H amylopectin and ,0H amylose

"op mai+e kernels are characterised by a high proportion of hard endosperm which ismuch higher than in any other mai+e kernel! "op mai+e is grown on a small scalecompared to other types but popped kernels are consumed world@wide as a snackfood! The ta(onomic relationship of pop mai+e with other mai+e is still under


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discussion ;reviewed in Iiegler 200,

Sweet mai+e is grown for green ears ;sweet corn

more recessive mutations blocking conversion of sugar to starch!

&.&.& Processing of grain maize

/mportant ways of processing grain mai+e include

Traditional processing:rain mai+e is eaten by numerous people especially in'atin merica the >S frica and sia! vast number of recipes e(istinvolving whole grains mai+e meal or mai+e flour! %ai+e grains may or may not

be roasted before lye@cooking lime@cooking and3or fermenting to preparetraditional foods or drinks! n important way of preparing mai+e grains for

cooking involves lime@cooking steeping and removal of the pericarp resulting inJni(tamalB which can be used in the preparation of various soups or doughs;masa

Dry-milling:%ai+e grains are either subEected to the

o 4ull@4at %illing "rocess resulting in mai+e meal in which germ and

crude fibre content are highly similar to whole mai+e grains=

o &olted %illing "rocess by which the mai+e meal is sifted to

e(clude large particles germ tip cap and bran pieces! &y@productsof this process include mai+e flour and hominy grits= or

o Tempering@egerming %illing "rocess by which moisture is added

in the milling process ;tempering< to facilitate almost completeremoval of the germ ;degerming< and bran fractions! /n addition tothe prime grits meals and flours obtained from the endospermfraction of the grain hominy feed for use in the manufacture ofcattle swine poultry and auatic feed can be obtained!

These descriptions are based on uensing et al ;200,

Wet-milling:uring the highly comple( processes of wet@milling the constituentsof the mai+e grains F carbohydrates proteins oil and crude fibre F are separatedand prepared for the use in food food manufacture animal feed and industrialuses! series of steps involving steeping the grain coarse and fine grindingcentrifugation and evaporation of steep water are employed!

o The starch fraction of the mai+e grains has a variety of uses

including: 6ative starch used in baby foods snack foods saladdressings paper products insulating materials paints tablet

binders= modified starch is used in bakery products sauces andgravies icings and gla+es pastes and glues ceramics dyes andsandpaper= glucose and fructose ;in corn syrup< are used in

beverages cheese spreads desserts fruit Euices fro+en seafoods


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lia

.reen corn

Area

&ha'

rld 9615011 95213 890582 15035039

4$A 269570 23500 2975870 270510Austra

lia35100 5283 599 5000

Yie()&*%+ha'

rld 75397 75861 9520 95636

4$A 10511 125277 13583 15923

Australia

9586 11578 1358 105000

,ro)uc-ion &-'

rld 572580

58725207

950935192

85200592

4$A 2592500

0

35108570

0

500510

0

5117527

0

Australia

27531 85236 650 05000

$urce /i*ures 'r 186 : 188 (i#cl.) ;ere ta$?A? ;e!site (http@@'astat.'a.r*@) i# 2007. ?he 2007 'i*ures ;ere taS *orn &elt hybrid and not until the 1.8 F .season that the first ustralian@bred hybrids ;C7 and C112< were released;*olless 19

/n ustralia now mai+e may be grown in every State and in the 6orthern Territory;6T< as an irrigated or dryland crop depending on rainfall conditions ;eg "rice 19="/D4 2007= 4arrell D )KNeeffe 2009= "/- @ Tasmania 2008

southern 6ew South -ales ;%urrumbidgee %urray and 'achlan #iver Valleys11
http://faostat.fao.org/http://faostat.fao.org/


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compared to appro(imately half of the crops in northern 6S- and Lueensland;#obinson D Nirkby 2002

Table 3: Characteristics of some of the current maize growing regions in NSW

/epresen-a-i0e si-e

&i-hin area'

2uirin)i 3Li0erpoo(,(ains

&Nor-hern4n(an)'

Lee-on

&Sou-hern4n(an)''

4n0ere((

&Nor-hernTab(e(an)s'

Casino

&Nor-hCoas-'

A0era%e )ai(yma+min-empera-ure6C &Oc- #ar'

28.7@13. 29.7@1. [email protected] [email protected]

A0era%e )ai(yma+min-empera-ure6C &Apr Sep-'

[email protected] [email protected] 18.9@2. [email protected]

A0era%emon-h(yraina(( mm&Oc- #ar'

76. 33.9 83.6 112.7

A0era%emon-h(yraina(( mm&Apr Sep-'

7.3 39.2 0 0.

.roin%season

Au*ust :>ct!er

$epte!er :%e!er

$epte!er :%e!er

Au*ust :a#uar"

Eamp(e oarab(e soi(-ype

Blacnited States are yield and yield stability and therelatively short time that hybrids remain on the commercial market ;less than 10 yearson average< is due to their replacement by more high@yielding hybrids rather than any

problems with disease or insect susceptibility ;uvick D *assman 1

reduction in barreness at high density and reduced rate of leaf senescence duringgrain filling ;uvick D *assman 1

!utation *reeding

%utation breeding in mai+e encompasses a number of approaches includingspontaneous mutation cell@ or tissue@culture induced mutation ;also referred to assomaclonal variation


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&oyer D Cannah 1.

been used successfully in the development of commercially available imida+olinone@tolerant mai+e varieties ;nderson D eorgeson 18= 6ewhouse et al! 11=reviewed in Tan et al! 2005< ;see also Section 9!1

&.1.& 'ene+ic mo,ifica+ion

The most important advances in mai+e improvement via genetic modification arediscussed below! Singletary ;200,< has provided a detailed review on this subEect!

%onocot plants such as mai+e are generally more difficult to transform than dicotspecies! Cowever both efficient transient and stable transformation protocols formai+e have been developed and improved over time includingAgro*acterium@mediated transformation transformation through protoplast fusion particle

bombardment and silicon carbide whiskers ;eg ordon@Namm et al! 10= ould et

al! 11= 4rame et al! 1.= /shida et al! 17= -right et al! 2001= 4rame et al! 2002

The latest advances of genetic modification in mai+e include the development ofcircular mai+e minichromosomes ;%%*sThe maEor focus in the production of % plants has been on resistance to insects andtolerance to herbicides! 4or e(ample resistance to corn rootworm has been achieved

by using cry@genes from%acillus thuringiensis and tolerance to the herbicidesglufosinate ammonium and glyphosate using thepat@gene from "treptomycesviridochromogenesand the cp,epsps-genefromAgro*acteriumspp! respectively!

nother focus in early mai+e research was to increase lysine levels in mai+e whichwas achieved by transformation with the cordapAgene from Coryne*acterium

glutamicumencoding lysine@insensitive dihydrodipicolinate synthase! These e(amplesare approved for commercial release overseas and for use in human food in ustraliaand overseas ;4S6I 2002= "C/S3>S 2002= "C/S3>S 2005="C/S3>S 2007a= "C/S3>S 2007b= "C/S3>S 2009

%ai+e grains like those of other cereals do not contain the amino acids lysinetryptophan and methionine at levels that are sufficient in the diet of humans and othermonogastric animals ;as reviewed in 'ai D %essing 2002< ;see also discussion inSection 5!1!2

SECT!"N/ "RP2"L"'%

The following descriptions are adapted from "aliwal ;2000f< and 4arnham et al;200,

/.# Plan+ morphology

The typical mai+e plant is a tall ;1 F . m< annual grass ;monocot< which forms a


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seasonal root system bearing a single erect stem ;culm< made up of nodes andinternodes although some cultivars may develop elongated lateral branches ;tillers

6odes gradually taper to the top of the plant! 'eaves are broad and a single leaf

develops at each node in two opposite ranks F the leaf arrangement mai+e isdistichous ;reviewed in $sau 199b

plants develop more leaves than temperate cultivars!

The epidermis is the outermost layer that can be discerned in cross@sections of thestalk! Several layers of sclerenchyma tissue are underneath it and increase the strengthof the stalk! The arrangement of vascular bundles is comple( and they appearscattered throughout the parenchyma tissue that constitutes the remainder of the cross@

section although they are arranged more loosely towards the middle ;reviewed in$sau 199b

%ai+e F like many plants that evolved under tropical conditions F is a *.plant and

therefore is more efficient at utilising carbon dio(ide than *,plants! These

physiological characteristics are reflected in leaf morphology down to the microscopiclevel! 4or e(ample bundle sheath cells are richer in chloroplasts than mesophyll cells!The chloroplasts are also larger than those of mesophyll cells ;reviewed in $sau199a

/.& Repro,c+i-e morphology

%ai+e is a monoecious plant: one or more lateral branches the shanks develop in theleaf a(is of the plant! They terminate in a female inflorescence an ear ;see 4igure 1

>sually one or two lateral shoots in the upper part of the plant develop into femaleinflorescences! The shank consists of nodes and short internodes the lengths of whichvary between mai+e races! The ears arise from a(illary bud apices! The ear is coveredin a number of leaves called husks! Those leaves differ in appearance when comparedto those on the stalk: they surround and protect the developing ear! -here mai+e isleft to dry in the fields more husks are generally desired to protect the grains from

birds and insects! -here mai+e is harvested earlier it is often desirable for a cultivarto have a lower number of thin husks!

The ear does not usually show any lateral branching! The thick a(is of the ear thecob bears an even number of rows ;between . and ,0< of ovaries each containing asingle ovule! The number of ovules that will develop into kernels ranges from ,00 F1000 and is dependant on the cultivar3variety as well as factors occurring later indevelopment ;"urseglove 192

2,


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Silks ;long green strandsnless stated otherwise the following description is adapted from "aliwal ;2000f

/nitially male and female inflorescences have primordia of bise(ual flowers!Cowever during their development primordia of stamens abort in the a(illaryinflorescences and primordia of gynoecia abort in the apical inflorescence!

The apical meristem elongates once the leaf primordia are initiated! /t is transformedinto a reproductive meristem that develops into the tassel! "ollen is shed from thetassel continuously for a week or more as upper and lower florets in the male spikeletsshow developmental differences and the spikes mature asynchronously!

The female inflorescences ;ears< arise from a(illary buds and bear flowers in rowsalong the cob! evelopment of the flowers and the ovules on the ear proceeds from


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the base upwards ;acropetal

have short hairs trichomes which form an angle to the stylar canals and helpharbouring pollen grains! #eceptive silks are moist and sticky!

%ai+e is often considered as protandrous ;anthers reaching maturity before thegynoeciumnderany stress ;especially water stress F see Section 7!,


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donor and receptor ;eg are pollen donor and receptor arrangedin a concentric fashion neighbouring or distant from eachother/n ustralia the following isolation distances for the production of speciality mai+ecurrently are recommended:

/n L' the production of sweet corn reuires either a .00 m isolation distancefrom all other mai+e or staggering mai+e plantings so that there are at least 1. F 21days between pollination times to all other mai+e ;&eckingham 2009

/n 6S- the production of speciality mai+e eg wa(y white and popcorn varietiesreuires either 800 m isolation distance or staggering mai+e plantings at least si(week so as to avoid cross pollination ;4arrell D )KNeeffe 2009

s reviewed in 4eil and Schmid ;2002

1./.# Em5ryogenesis

The following information is adapted from Sass ;199< and Sheridan and *lark;1.

/n the first phase of embryogenesis irregular cell divisions result in a proembryoconsisting of appro(imately 12 F 2. cells within 100 hrs after fertilisation! The initial

basal cell divides into a number of large vacuolated suspensor cells! &y the sametime the initial apical cell has given rise to F 18 small cells that are densely filledwith cytoplasm!

Starting appro(imately 8 F days after fertilisation the second phase of

embryogenesis commences which leads to the establishment of meristems and theembryonic a(is by appro(imately 1, days after fertilisation ;transition embryo

29


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ppro(imately 1. F 15 days after fertilisation the coleoptilar embryo is establishedcharacterised by the differentiation and growth of scutellumf coleoptileg coleorhi+ah

and root and shoot apical meristem ;see 4igure ,

1 mg!

uring the last stage of embryogenesis ;during ,0 F .0 days after the first leafprimordium appears1./.& Fri+6see, ,e-elopmen+

The fruit of mai+e is a caryopsis a dry indehiscent single@seeded fruit ;4igure 2

igure 2% +ature maize car-opses [photo credit: Steve Hurst @ USDA-N!S "#AN$SDatabase% "rovided b&AS S&stematic 'otan& and &coo*& #aborator&+ 'oivia%!ochabamba+ (http:,,pants+usda+*ov,ava,pro.ie/s&mbo012A)3

The pericarp ;ovary wall< and testa ;seed coat< are fused to form the fruit wall andbecause of this tight adhesion between fruit and seed the two structures actuallyappear to be a single structure! This structure is commonly referred to by a number ofinterchangeable terms F fruit kernel grain and seed! The kernels are composed ofthree main parts @ the embryo the endosperm and the fruit wall ;see 4igure ,

f The scutellum is the name given to the cotyledon of grasses ;#aven et al! 1

absorption of food by the embryo from the endosperm ;see Section 5!1!, P"TENT!ALF"R*ERT!CAL'ENETRANSFER

Vertical gene transfer is the transfer of genetic material from parent to offspring byreproduction! This type of gene transfer can occur by se(ual or ase(ual reproduction!This section deals with gene transfer to other plants of the same species or closely

related species by se(ual reproduction!

Successful gene transfer reuires that three criteria are satisfied! The plant populationsmust: 1< overlap spatially ;sympatry

subspecies ofZ. mays namely ssp! mexicana;central %e(icoperennial>.# Na+ral in+raspecific an, in+erspecific crossing

ll the various types ofZ. maysssp! maysfreely cross@pollinate and form fertile

hybrids ;"urseglove 192

)utcrossing has been discussed in Sections 2!,!1 and .!2!,! -ind pollination canoccur between mai+e crops over hundreds of metres but the relatively large weightand diameter of the pollen grains favours most pollen deposition within appro(imately70 m of the source plant ;#aynor et al! 192= 'una et al! 2001= ylor 2002= Aaros+ etal! 200,< and there is little or no cross pollination at ,00 m ;'una et al! 2001

et al! ;2005< have considered both time and distance components of pollen@mediatedgene flow in mai+e and concluded that under their e(perimental conditions 200 m


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was sufficient to reduce outcrossing to R 0!1H! lso of significance in consideringthe likelihood of cross@pollination in mai+e is pollen competition! The fact that asingle plant can produce millions of pollen grains ;see Section .!2!1< means that evenif there is long distance spread of pollen from a plant the pollen will be greatlyoutnumbered by JlocalB pollen ;&annert D Stamp 2009

reduce pollen production such as detasseling or cytoplasmic male sterility mayactually increase the likelihood of long distance cross pollination because they reduce

pollen competition ;&annert D Stamp 2009

The international seed certification standards provide a guide to the physicalseparation of crops to minimise gene flow ;see Section 2!,!1

genetically modified ;%< and non@% mai+e crops ;Cenry et al! 200,= $astham DSweet 2002= Stevens et al! 200.= oggi et al! 2007= %esseguer et al! 2007= Sanvido etal! 2008

Various incompatibilities e(ist both between and within subspecies ofZ. mays and arethought to have evolved to prevent indiscriminate hybridi+ation ;Nermicle 19

pre+ygotic barrier to crossing ;Schwart+ 150= Nermicle 19

The teosintes were formerly placed in the genus2uchlaena;*ollins 125=%angelsdorf D #eeves 1,1

%e(ico and uatemala where they share a common distribution ;-ilkes 199=oebley 10nusually the flow of genes has occurred in both directions;reciprocal introgression< ;-ilkes 199= oebley 10< although a number of factorstend to favour gene flow from teosinte to mai+e rather than from mai+e to teosinte;&alta+ar et al! 2005

populations ofZ. maysteosintes when teosinte is the female and mai+e the male

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parent and this has been linked to a teosinte gene or gene cluster known as Teosintecrossing *arrier' Tc*';$vans D Nermicle 2001= Nermicle 2007

suggested that the transfer of Tc*'into mai+e may be useful in avoidingcontamination of one commercial variety with another!

The likelihood of intraspecific or interspecific crossing occurring naturally inustralia is remote! Z. maysssp! mexicanaimported from South merica can now befound in 6orth Lueensland and -estern ustralia ;"heloung et al! 1a< but therehave not been any reports of introgression with commercial mai+e cultivars!

>.& Crossing n,er e3perimen+al con,i+ions

These crosses while possible under controlled conditions do not occur naturally andtherefore are of significance only in the conte(t of broadening an understanding ofunaided gene transfer ;)$* 2007

>.&.# !n+erspecific crosses

*rosses betweenZ. mays;2n?20< andZ. perennis;2n?.0< may normally yield only0!1 F 1H viable seed because of endosperm collapse after appro(imately 21 days post

pollination! Cowever hybrids ;with 2n?,0< have been obtained following embryorescue ;del *armen %olina D arcia 1

>.&.& !n+ergeneric crosses

There has been considerable speculation on the evolution of cultivated mai+eincluding suggestion that intergeneric introgression betweenZeaand Tripsacum;thatare both members of the sub@family "anicoidea tribe %aydeae< may have played arole Zsee e!g! discussion in de -et D Carlan ;19.


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distance the Tripsacumpollen tube had to grow was reduced ;eg by shortening themai+e styles

backcross generations at least 5. chromosome combinations can be obtained ;Carlan

D de -et 199

been a report ofZ. maysbeing crossed with Coixlachryma-8o*i;Carada et al! 15.

Mork "eninsula area ;6orth ustralian 'and %anager 2008

%ai+e has also been crossed with another member of the "anicoideae sub@family! single intergeneric hybrid has been obtained between sugarcane ;"accharumo&&icinarumF tribe ndropogonae< with 2n ? 80 andZ. mayswith two additional &chromosomes using mai+e as the pollen parent ;Aanaki mmal et al! 192

)ther intergeneric crosses involving mai+e have all been with members of the sub@

family "ooideae! vailable information ;see eg Nynast et al! 2001< suggests that when"anicoideae are crossed with "ooideae chromosomes of the "anicoideae areeliminated soon after fertili+ation! $(amples include:

%ai+e ;pollen donor< crosses readily with he(aploid ;2n ? .2< wheat ;Triticumaestivum< although double fertili+ation is rare and development of the embryo

proceeds without the formation of endosperm ;Ihang et al! 17

i pomi(is is a genetically controlled mechanism in which an embryo is formed without union of male

and female gametes! iplospory is a form of apomi(is in which the embryo develops from a diploidegg that has derived from an unreduced megaspore mother cell! The resulting individual is thereforegenetically identical to the maternal parent ;Nindiger D Sokolov 18

59


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during early embryogenesis resulting in the formation of a wheat haploid ;'aurieD &ennett 188

/n controlled crosses of he(aploid ;2n ? .2< oat ;Avena sativa< with mai+e aproportion ;appro(imately ,1H< of the embryos may contain 1 F . mai+echromosomes in addition to a full complement of oat chromosomes and aretherefore referred to as partial hybrids ;#iera@'i+ara+u et al! 17= Nynast et al!2001

Ienkteler D 6it+sche ;18.< made a number of crosses within the cerealsincluding barley ;Hordeum vulgare= 2n ? 1.< and rye ;"ecale cereale= 2n ? 1.< asfemale parents withZea mays as the pollen donor! The crosses were made on in

vivo@grown plants and no attempt was made at embryo rescue! 6o embryosdeveloped in theH. vulgare(Z. mayscross but globular embryos formed in someof the ". cereale(Z. mayscrosses although these degenerated after 7 @ 10 days

presumably because of poor or no endosperm development!

evelopment of embryos from barley florets pollinated with mai+e has beenobtained following embryo rescue ;*hen et al! 11


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http://www.clw.csiro.au/publications/technical2002/tr13-02.pdfhttp://www.clw.csiro.au/publications/technical2002/tr13-02.pdfhttp://www.dpi.vic.gov.au/dpi/nreninf.nsf/childdocs/-89E7A8DAFEA417624A2568B30004C26A-BBB90E3F477D0B74CA256BC700811C92-B494B54DA7CA01F94A256DEA002753B4-CE2CED1252495905CA256BCF000BBD45?openhttp://www.dpi.vic.gov.au/dpi/nreninf.nsf/childdocs/-89E7A8DAFEA417624A2568B30004C26A-BBB90E3F477D0B74CA256BC700811C92-B494B54DA7CA01F94A256DEA002753B4-CE2CED1252495905CA256BCF000BBD45?openhttp://www.dpi.vic.gov.au/dpi/nreninf.nsf/childdocs/-89E7A8DAFEA417624A2568B30004C26A-BBB90E3F477D0B74CA256BC700811C92-B494B54DA7CA01F94A256DEA002753B4-CE2CED1252495905CA256BCF000BBD45?openhttp://www.dpi.vic.gov.au/dpi/nreninf.nsf/childdocs/-89E7A8DAFEA417624A2568B30004C26A-BBB90E3F477D0B74CA256BC700811C92-B494B54DA7CA01F94A256DEA002753B4-CE2CED1252495905CA256BCF000BBD45?openhttp://www.dpi.nsw.gov.au/__data/assets/pdf_file/0008/146690/err-20-Farming-Systems-in-the-Northern-Cropping-Region-of-NSW---An-Economic-Analysis-Region-of-NSW-An-Economic-Analysis.pdfhttp://www.dpi.nsw.gov.au/__data/assets/pdf_file/0008/146690/err-20-Farming-Systems-in-the-Northern-Cropping-Region-of-NSW---An-Economic-Analysis-Region-of-NSW-An-Economic-Analysis.pdfhttp://www.dpi.nsw.gov.au/__data/assets/pdf_file/0008/146690/err-20-Farming-Systems-in-the-Northern-Cropping-Region-of-NSW---An-Economic-Analysis-Region-of-NSW-An-Economic-Analysis.pdfhttp://www.dpi.nsw.gov.au/__data/assets/pdf_file/0008/146690/err-20-Farming-Systems-in-the-Northern-Cropping-Region-of-NSW---An-Economic-Analysis-Region-of-NSW-An-Economic-Analysis.pdfhttp://www.regional.org.au/au/asa/2006/poster/environment/4578_shavrukovy.htm?print=1%5Chttp://www.regional.org.au/au/asa/2006/poster/environment/4578_shavrukovy.htm?print=1%5Chttp://www.regional.org.au/au/asa/2006/poster/environment/4578_shavrukovy.htm?print=1%5Chttp://www.ruralsolutions.sa.gov.au/__data/assets/pdf_file/0005/43349/seeds_manual.pdfhttp://www.ruralsolutions.sa.gov.au/__data/assets/pdf_file/0005/43349/seeds_manual.pdfhttp://sgrl.csiro.au/aptc2003/21_staples_etal.pdfhttp://sgrl.csiro.au/aptc2003/21_staples_etal.pdfhttp://www.dpi.nsw.gov.au/agriculture/pests-weeds/weeds/publications/management/summerhttp://www.dpi.nsw.gov.au/agriculture/pests-weeds/weeds/publications/management/summerhttp://www.dpi.vic.gov.au/dpi/nreninf.nsf/9e58661e880ba9e44a256c640023eb2e/5f4bd3e13487dbe4ca256e720024dccd/$FILE/LC0009.pdfhttp://www.dpi.vic.gov.au/dpi/nreninf.nsf/9e58661e880ba9e44a256c640023eb2e/5f4bd3e13487dbe4ca256e720024dccd/$FILE/LC0009.pdfhttp://www.dpi.vic.gov.au/dpi/nreninf.nsf/9e58661e880ba9e44a256c640023eb2e/5f4bd3e13487dbe4ca256e720024dccd/$FILE/LC0009.pdfhttp://www.dpi.vic.gov.au/dpi/nreninf.nsf/9e58661e880ba9e44a256c640023eb2e/5f4bd3e13487dbe4ca256e720024dccd/$FILE/LC0009.pdfhttp://www.affashop.gov.au/product.asp?prodid=13796http://www.affashop.gov.au/product.asp?prodid=13796http://www.sciencedaily.com/releases/2007/10/071016101454.htmhttp://www.sciencedaily.com/releases/2007/10/071016101454.htmhttp://www.ars-grin.gov/cgi-bin/npgs/html/taxon.plhttp://www.ars-grin.gov/cgi-bin/npgs/html/taxon.plhttp://www.agric.wa.gov.au/content/pw/vp/rod/28_plague_rat.pdfhttp://www.agric.wa.gov.au/content/pw/vp/rod/28_plague_rat.pdfhttp://delta-intkey.com/grass/http://delta-intkey.com/grass/http://www.sgrl.csiro.au/aptc1998/16_webley_jackson.pdfhttp://www.sgrl.csiro.au/aptc1998/16_webley_jackson.pdfhttp://www.springerlink.com/content/ym84k246x6v57450/fulltext.pdfhttp://www.isb.vt.edu/news/2008/artspdf/feb0801.pdfhttp://www.isb.vt.edu/news/2008/artspdf/feb0801.pdfhttp://www.clw.csiro.au/publications/technical2002/tr13-02.pdfhttp://www.clw.csiro.au/publications/technical2002/tr13-02.pdfhttp://www.dpi.vic.gov.au/dpi/nreninf.nsf/childdocs/-89E7A8DAFEA417624A2568B30004C26A-BBB90E3F477D0B74CA256BC700811C92-B494B54DA7CA01F94A256DEA002753B4-CE2CED1252495905CA256BCF000BBD45?openhttp://www.dpi.vic.gov.au/dpi/nreninf.nsf/childdocs/-89E7A8DAFEA417624A2568B30004C26A-BBB90E3F477D0B74CA256BC700811C92-B494B54DA7CA01F94A256DEA002753B4-CE2CED1252495905CA256BCF000BBD45?openhttp://www.dpi.vic.gov.au/dpi/nreninf.nsf/childdocs/-89E7A8DAFEA417624A2568B30004C26A-BBB90E3F477D0B74CA256BC700811C92-B494B54DA7CA01F94A256DEA002753B4-CE2CED1252495905CA256BCF000BBD45?openhttp://www.dpi.nsw.gov.au/__data/assets/pdf_file/0008/146690/err-20-Farming-Systems-in-the-Northern-Cropping-Region-of-NSW---An-Economic-Analysis-Region-of-NSW-An-Economic-Analysis.pdfhttp://www.dpi.nsw.gov.au/__data/assets/pdf_file/0008/146690/err-20-Farming-Systems-in-the-Northern-Cropping-Region-of-NSW---An-Economic-Analysis-Region-of-NSW-An-Economic-Analysis.pdfhttp://www.dpi.nsw.gov.au/__data/assets/pdf_file/0008/146690/err-20-Farming-Systems-in-the-Northern-Cropping-Region-of-NSW---An-Economic-Analysis-Region-of-NSW-An-Economic-Analysis.pdfhttp://www.regional.org.au/au/asa/2006/poster/environment/4578_shavrukovy.htm?print=1%5Chttp://www.regional.org.au/au/asa/2006/poster/environment/4578_sha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