Biologia Del Maiz

The Biology of Zea mays L.
ssp mays (maize or corn)
Office of the Gene Technology Regulator
The Biology of Zea mays L. ssp mays (maize or corn)
Version 1: September 2008
This document provides an overview of baseline biological information relevant to risk assessment of genetically modified forms of the species that may be released into the ustralian environment!
For information on the Australian Government Office of the Gene Technology egulator visit !http"##$$$.ogtr.gov.au%
TABLE OF CONTENTS
"#$ %&'$!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!
Section 1 Section 2 2!1 2!2 2!2!1 2!2!2 2!2!, 2!, 2!,!1 2!,!2 2!,!, 2!. 2!.!1 2!.!2 Section , ,!1 ,!2 Section . .!1 .!1!1 .!1!2 .!2 .!2!1 .!2!2 .!2!, .!, .!,!1 .!,!2 .!,!, .!. .!5 Section 5 5!1 5!1!1 5!1!2 5!1!, 5!2 5!2!1 5!, 5!. 5!5 Section 7 ,
Ta(onomy!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! )rigin and *ultivation!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! *entre of diversity and domestication!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! *ommercial uses!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! %ai+e types and their uses!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! "rocessing of grain mai+e!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! -orld mai+e production!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! *ultivation in ustralia!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! *ommercial propagation!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! Scale of cultivation!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! *ultivation practices!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! *rop /mprovement!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! &reeding!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! 0enetic modification!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! %orphology!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! "lant morphology!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! #eproductive morphology!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! 1evelopment!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! #eproduction!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! se(ual reproduction!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! Se(ual reproduction!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! "ollen dispersal2 pollination and outcrossing rates!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! "ollen!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! "ollen dispersal and pollination!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! )utcrossing rates and isolation distances!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! $mbryogenesis2 fruit3seed development and seed dispersal!!!!!!!!!!!!!!!!!!!! $mbryogenesis!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! 4ruit3seed development!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! Seed dispersal!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! Seed dormancy and germination!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! Vegetative growth!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! &iochemistry!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! 6utrient components of the mai+e kernel!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! Starch!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! "rotein!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! 'ipids!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! To(ins!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! 6itrate poisoning!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! llergens!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! )ther undesirable phytochemicals!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! &eneficial phytochemicals!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! biotic /nteractions!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!
7!1 7!2 7!, 7!. Section 9 9!1 9!2 9!2!1 9!2!2 9!2!, 9!2!. 9!, Section 8 8!1 8!2 8!5 Section : :!1 :!2 :!2!1 :!2!2
6utrient re8uirements!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! Temperature re8uirements and tolerances!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! -ater!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! )ther abiotic stresses!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! &iotic /nteractions!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! -eeds!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! "ests and diseases!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! /nsects and other invertebrate pests!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! Vertebrate pests!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! 1iseases!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! )ther adverse associations with mai+e!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! )ther biotic interactions!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! -eediness!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! -eediness status on a global scale!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! -eediness status in ustralia!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! *ontrol measures!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! "otential for Vertical 0ene Transfer!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! 6atural intraspecific and interspecific crossing!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! *rossing under e(perimental conditions!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! /nterspecific crosses!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!! /ntergeneric crosses!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!
REFERENCES...........................................................................................................................................
PREA
BLE
This document describes the biology of Zea mays '! subspecies ;ssp!< mays2 with particular reference to the ustralian environment2 cultivation and use! /nformation included relates to the ta(onomy and origins of cultivated Z. mays ssp! mays2 general descriptions of its morphology2 reproductive biology2 biochemistry2 and biotic and abiotic interactions! This document also addresses the potential for gene transfer to occur to closely related species! The purpose of this document is to provide baseline information about the parent organism in risk assessments of genetically modified Z. mays ssp! mays that may be released into the ustralian environment! s mai+e is one of the best researched and characterised plants2 significant amounts of information are available for many aspects of the biology of mai+e2 and the reader is referred to the literature provided in this document as a starting point! /n this document the terms mai+e and corn are used to refer to Z. mays ssp! mays. )ther subspecies of Zea mays are referred to as teosintes! %ai+e is an annual grass growing up to . m tall! The female inflorescences2 the ears2 develop in leaf a(ils on the stalk2 which terminates in the male inflorescence2 the tassel! The broad leaf sheaths are overlapping around the stalk and the leaves are arranged in two opposing rows along the stalk! %ai+e has a multitude of uses and is used in the preparation of food or drinks2 as animal feed or for industrial purposes!
SECT!"N #
TA$"N"
The genus Zea belongs to the tribe ndropogoneae in the subfamily "anicoideae in the family "oaceae ;reviewed in )$*1 200,= >S1 2005<! There are currently 87 recognised genera within the ndropogoneae tribe ;>S1 2005<! *urrently2 there are five species included in the genus Zea. Species of Zea that have been e(amined2 largely have a chromosome number of 2n ? 202 e(cept for Z. perennis ;perennial teosinte with 2n ? .0< ;as reviewed in Tito et al! 1::1= $llneskog@Staam et al! 2009= Table 1<! The species Z. nicaraguensis was described by /ltis and &en+ ;2000<= it is closely related to Z. luxurians but currently2 the number of chromosomes and se(ual compatibility with other Zea spp! are unknown!
Table 1
Species 1 . 2 . Zea diploperennis HH Iltis et al Zea luxurians (Durieu & Asch.) RM Bird
Zea species and subspecies*

Chromosom e number 2n = 20 2n = 20 Subspecies Synonyms Euchlaena luxurians Durieu & Asch. Zea mays ssp luxurians (Durieu & Asch.) HH Iltis
3 .
Zea mays L.
2n = 20
Zea mays ssp huehuetenangensis (HH Iltis & D e!le") D e!le" Zea mays ssp mays Zea curagua M li#a Zea indentata $turte%.
Species
Chromosom e number
Subspecies
Synonyms Zea indurata $turte%. Zea japonica &a# H utte Zea mays c% alba Ale'. Zea mays c% leucodon Ale'. Zea mays var flavorubra Zea mays %ar i#de#tata ($turte%.) LH Baile" Zea mays var indurata ($turte%.) LH Baile" Zea mays %ar japonica (&a# H utte) Alph. ( d Zea mays %ar saccharata ($turte%.) LH Baile" Zea mays %ar tunicata Larra)a*a e+ A.$t.-Hil. Zea mays %ar vulgate , er#. & H (er#er Zea saccharate $turte%.
Zea mays ssp mexicana ($chrad.) HH Iltis
Euchlaena mexicana $chrad. Zea mexicana ($chrad.) ,u#t-e Zea mays %ar parviglumis Euchlaena perennis Hitchc.
Zea mays ssp parviglumis HH Iltis & D e!le" . . 1 . Zea nicaraguensis HH Iltis & B/ Be#2n =0 2n = .0 -
Zea perennis (Hitchc.) Ree%es & Ma#*elsd. 2$ urce3 4$DA (2001)
Z. maysa ssp! mays is the only cultivated species= the other species and subspecies are wild grasses2 referred to as teosintes! /n addition to the basic chromosome complement2 mai+e plants may contain one or more supernumerary chromosomes2 called & chromosomes2 which do not pair with chromosomes during meiosis ;as reviewed in Aones et al! 2008<!
a The name Zea is from the 0reek zea meaning cereal or grain! The specific epithet mays is thought to derive from the native rawak word mai+ or mahi+ used in the mericas to describe the plant= the word was adopted by the Spanish crew of *olumbusB first voyage who first collected the grain from the mericas and took it to $urope ;Cyam D "ankhurst 1::5= 1esEardins D %c*arthy 200.<!
The mai+e genome is large2 being somewhere between 2!, F 2!9 0bpb ; rumuganathan D $arle 1::1<2 with a total gene number of between .22000 and 572000 genes! *onsiderable work has been done on characteri+ing the mai+e genome ;see eg #abinowic+ D &ennet+en 2007= %essing D 1ooner 2007<! The mai+e genome is characterised by a high percentage of repetitive se8uences2 including transposons and retrotransposons ;Cake D -albot 1:80= 'iu et al! 2009<! To date2 the best characterised transposons are the Activator ; c< and Dissociation ;1s< elements2 first described by &arbara %c*lintock ;reviewed in 4edoroff 2000<! The presence of the autonomous Ac element is needed for transposition2 whereas Ds elements are non@ autonomous and trans@activated by an Ac element! %ai+e chloroplast and mitochondrial genes have been characterised in detail2 with a focus on cytoplasmic male sterility ;reviewed in )$*1 200,<! There is a programme initiated in 2005 to se8uence the mai+e genome and a dedicated website describes progress of the various mai+e genome se8uencing proEects ;http:33www!mai+egenome!org3<!
SECT!"N &
&.#
"R!'!N AN( C)LT!*AT!"N

Cen+re of ,i-ersi+y an, ,omes+ica+ion
The centre of origin of mai+e is the %esoamerican region2 probably in the %e(ican highlands2 from where it spread rapidly! rchaeological records and phylogenetic analysis suggest that domestication began at least 72000 years ago ;"iperno D 4lannery 2001= %atsuoka et al! 2002<! %ai+e spread around the world after $uropean discovery of the mericas in the 15th century2 particularly in temperate +ones ;"aliwal 2000d= 4arnham et al! 200,<! %ai+e is only known as a cultivated crop and its e(act genealogy remains uncertain! Zea mays ssp! parviglumis is hypothesised to be the progenitor of cultivated mai+e! This hypothesis is supported by the close genetic compatibility and relationship between the two sub@species ;reviewed in 1oebley 200.<! Cowever2 ribosomal internal transcribed spacer se8uence data suggest that Z. mays ssp! mays diverged no later than the ssp! mexicana and parviglumis ;&uckler D Coltsford 1::7<! The latter is supported by data on the distribution of alleles in organelles2 but conflicting with iso+yme data ;discussed in &uckler D Coltsford 1::7<! 1uring the domestication of mai+e2 every region in which it has been cultivated over the centuries has produced a selection of mai+e cultivars or landraces ;refer Section 2!.<! 4armers have maintained and improved these and they are adapted to local re8uirements and characteristics ;"aliwal 2000a<! %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 .0G South ;eg *hile<! 0enerally2 tropical mai+e is grown between ,0G6orth and ,0G South2 subtropical mai+e between ,0 and ,.G both 6orth or South2 and temperate mai+e beyond ,.G latitudes! /t can be grown in a range of altitudes from sea level up to ,2800 metres and with growing seasons between .2 and .00 days! This ability to grow in a wide range of environments is reflected in the high diversity of morphological and physiological traits!
b The amount of 16 in the nucleus of a eukaryotic cell is e(pressed as the total number of base pairs ;bp< in a haploid ;1*< chromosome complement!
&.&
Commercial .ses
%ai+e is one of the oldest cultivated grains and one of the most productive crop species 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 corn to mature grain! high proportion of mai+e produced is used as stock feed2 eg .0H in tropical areas and up to 85H in developed countries ;reviewed in "aliwal 2000g= 4arnham et al! 200,<! /t can be fed to stock as green chop2 dry forage2 silage or grain! Various fractions of milling processes can also be used as animal feed! Stover is the term used to describe the dried stalks and leaves of a crop used as animal fodder after the grain has been harvested! %ai+e can be processed for a range of uses both as an ingredient in food or drinks2 eg corn syrup in soft drinks or mai+e meal2 or for industrial purposes! %ai+e is the maEor source of starch world wide2 and is used as a food ingredient2 either in its native form or chemically modified ;-hite 1::.<! *orn starch can be fermented into alcohol2 including fuel ethanol2 while the paper industry is the biggest non@food user of mai+e starch! The oil and protein are often of commercial value as by@products of starch production and are used in food manufacturing ;&oyer D Cannah 1::.= "aliwal 2000h= Cobbs 200,= %c*utcheon 2009<! &.&.# aize +ypes an, +heir .ses
number of mai+e types can be discerned on the basis of endosperm and kernel composition ;"urseglove 1:92= "aliwal 2000c= 1arrah et al! 200,<! 4lint mai+e kernels are characterised by their high percentage of hard endosperm ;see 4igure ,< around a small soft centre! 4lint mai+e is grown predominantly in 'atin merica and $urope for food use! 1ent mai+e is the most commonly grown for grain and silage2 and is the predominant type grown in the >S ! Card endosperm is present on the sides and base of the kernel! The remainder of the kernel is filled with soft starch= when the grain starts drying the soft starch at the top of the kernel contracts2 producing the depression for which it is named! 4loury mai+e is being grown predominantly in the ndean region! /ts endosperm is mainly composed of soft starch2 making it easy to grind and process into foods! -a(y mai+e kernels contain almost entirely amylopectin as their starch ;rather than the normal 90H amylopectin and ,0H amylose<! -a(y mai+e is preferred for food in some parts of $ast sia and for some industrial uses= it produces a starch similar to tapioca! "op mai+e kernels are characterised by a high proportion of hard endosperm2 which is much higher than in any other mai+e kernel! "op mai+e is grown on a small scale compared to other types but popped kernels are consumed world@wide as a snack food! The ta(onomic relationship of pop mai+e with other mai+e is still under
discussion ;reviewed in Iiegler 200,<! Sweet mai+e is grown for green ears ;sweet corn<! The ears are harvested at appro(imately 18 to 20 days post pollination when kernel moisture is appro(imately 90H! The developing grain of sweet mai+e is higher in sugar content due to one or more recessive mutations blocking conversion of sugar to starch! &.&.& Processing of grain maize
/mportant ways of processing grain mai+e include
Traditional processing: 0rain mai+e is eaten by numerous people2 especially in 'atin merica2 the >S 2 frica and sia! vast number of recipes e(ist2 involving whole grains2 mai+e meal or mai+e flour! %ai+e grains may or may not be roasted before lye@cooking2 lime@cooking and3or fermenting to prepare traditional foods or drinks! n important way of preparing mai+e grains for cooking involves lime@cooking2 steeping and removal of the pericarp resulting in Jni(tamalB2 which can be used in the preparation of various soups or doughs ;masa<! %asa can be baked into tortillas2 chips etc ;reviewed in #ooney D Serna@ Saldivar 200,<! 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 particles2 germ2 tip cap and bran pieces! &y@products of this process include mai+e flour and hominy grits= or o Tempering@1egerming %illing "rocess by which moisture is added in the milling process ;tempering< to facilitate almost complete removal of the germ ;degerming< and bran fractions! /n addition to the prime grits2 meals and flours obtained from the endosperm fraction of the grain2 hominy feed for use in the manufacture of cattle2 swine2 poultry and a8uatic feed can be obtained! These descriptions are based on 1uensing et al ;200,<!
Wet-milling: 1uring the highly comple( processes of wet@milling the constituents of the mai+e grains F carbohydrates2 proteins2 oil and crude fibre F are separated and prepared for the use in food2 food manufacture2 animal feed and industrial uses! series of steps involving steeping the grain2 coarse and fine grinding2 centrifugation2 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 foods2 snack foods2 salad dressings2 paper products2 insulating materials2 paints2 tablet binders= modified starch is used in bakery products2 sauces and gravies2 icings and gla+es2 pastes and glues2 ceramics2 dyes and sandpaper= glucose and fructose ;in corn syrup< are used in beverages2 cheese spreads2 desserts2 fruit Euices2 fro+en seafoods2
e(plosives and shoe polish! The starch is also fermented to produce alcoholic beverages2 flavour enhancers2 industrial alcohols2 engine fuel and solvents! o The germ fraction can be separated into the oil fraction and meal fraction2 the latter of which can be used as animal feed! The oil can be used as cooking oil2 as a carrier for oil soluble vitamins2 and as an ingredient in mayonnaise2 shortenings2 soups2 insecticides2 linoleum etc! o &oth the fibre and protein fractions can be used in animal feeds! The steep water can be used for industrial purposes2 eg in the production of chemicals or yeast culture! The above descriptions are mainly based on Aohnson and %ay ;200,<! &.&./ 0orl, maize pro,.c+ion
/n 20002 6orth merica accounted for nearly 50H of the world mai+e production! The >S produced appro(imately .2H2 *hina appro(imately 18H and $urope appro(imately 10H2 whereas ustralia produced less than 0!1H ;data reviewed in 4arnham et al! 200,<! Table 2 shows the world production of mai+e! There are little data available on the area cultivated world wide for forage and silage! /n the >nited States of merica silage production accounts for appro(imately 10H of the total area planted to mai+e!
Table 2: World production of maize grain and green corn*
Year 1975 #ai$e %rain Area &ha' 4$A Austra lia Yie() &*%+ha' ( rld 4$A Austra lia ,ro)uc-i on &-' ( rld 4$A Austra ( rld 1215...5 1.1 2653775. 90 115381 25913 15.21 25183 3.157715 861 1.953715 062 1335300 13051035 611 305.3750 00 1025962 35620 65.06 25932 .9151265 301 2215.135 009 2815.30 1375.715 687 27539850 00 105218 35698 65123 .5927 11651385 961 19658795 882 2.25360 1..53675 .66 29518050 00 675000c .5911 85318 15000 78152295 290 27651895 000 3905000 1985 1995 !!"
c The figure for area sown to mai+e in ustralia is somewhat different from that supplied by & #$ ;2008< and discussed in Section 2!,! The & #$ estimate for area sown in ustralia in 2007309 was .:2000 ha and in 2009308 was 782000 ha!
lia .reen corn Area &ha' 4$A Austra lia Yie() &*%+ha' ( rld 4$A Austra lia ,ro)uc-i on &-' ( rld 4$A Austra lia 2695.70 35100 75397 1051.1 95.86 15172518 0 2592.500 0 2753.1 2135100 .5283 75861 125277 115.78 15872520 7 35108570 0 .85236 2975870 15.99 952.0 135813 135.8. 95093519 2 .500.510 0 6.5011 ( rld 9615011 9115213 8905821 15013503 9 27051.0 .5000 95636 115923 105000 85200592 . .5117527 0 .05000
2$ urce3 /i*ures ' r 1861 : 1881 (i#cl.) ;ere ta<e# 'r = i#' r=ati # supplied at the /A>$?A? ;e!site (http3@@'a stat.'a . r*@) i# 2007. ?he 2007 'i*ures ;ere ta<e# 'r = the sa=e site i# 2009.
&./
C.l+i-a+ion in A.s+ralia
/t is thought that although mai+e had been introduced by the "ortuguese to the southeast sian archipelago in the 17th century and that there were "ortuguese settlements on the island of Timor ;less than 500 km from the north west coast of ustralia<2 no agriculture was introduced to ustralia by the "ortuguese2 %alaysians or /ndonesians in the 17th or 19th centuries! /t is therefore likely that mai+e was introduced to ustralia with the first fleet in 19882 being grown initially on 6orfolk /sland and in Sydney ;"ort Aackson< ;1esEardins D %c*arthy 200.<! /n the 6orthern Territory it was grown by the first white settlers in 182. ;)K0ara 2009<! The first recorded systematic inbreeding of mai+e in ustralia began in the 1:20s2 firstly at 0rafton $(periment 4arm ;6S-< and later in other areas F Lueensland gricultural *ollege ;0atton<2 0len /nnes and &athurst $(periment 4arms ;6S-<2 Cawkesbury gricultural *ollege ;6S-< ;*olless 1:9:<! The maEor early cultivars were J-ellingroveB2 4unkBs Mellow 1entB2 J'arge #ed CoganB and J4it+royB! /t was not until 1:.9 that the first commercial hybrid mai+e crops were produced in ustralia2 based on a widely@grown >S *orn &elt hybrid2 and not until the 1:.8 F .: season that the first ustralian@bred hybrids ;0C:7 and 0C112 < were released ;*olless 1:9:<! %ai+e breeding in north Lueensland began in 1:71 and resulted in the release of the first tropical2 rust@resistant cultivar ;LN,9<! /n ustralia now2 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 1::9= 1"/D4 2007= 4arrell D )KNeeffe 2009= 1"/- @ Tasmania 2008<! /t is2 however2 grown mostly in Lueensland ; therton Tableland2 &urnett2 1arling 1owns< and southern 6ew South -ales ;%urrumbidgee2 %urray and 'achlan #iver Valleys< ;&irch et al! 200,<! ll mai+e crops in southern 6S- and Victoria are irrigated
11
compared to appro(imately half of the crops in northern 6S- and Lueensland ;#obinson D Nirkby 2002<! *haracteristics of some representative mai+e growing areas in ustralia are given in Tables , D .!
Table 3:
/epresen-a-i0 e si-e &1i-hin area'
Characteristics of some of the current maize growing regions in NSW

2uirin)i 3 Li0erpoo( ,(ains &Nor-hern 4n(an)' Lee-on &Sou-hern 4n(an)'' 4n0ere(( &Nor-hern Tab(e(an)s' Casino &Nor-h Coas-'
A0era%e )ai(y ma5+min -empera-ure 6C &Oc- 7 #ar'88 A0era%e )ai(y ma5+min -empera-ure 6C &Apr 7 Sep-'88 A0era%e mon-h(y rain9a(( mm &Oc- 7 #ar'88 A0era%e mon-h(y rain9a(( mm &Apr 7 Sep-'88 .ro1in% season8 E5amp(e o9 arab(e soi( -ype
28.7@13.1
29.7@1..1
29@12.3
29.9@17.8
18.1@..3
16.1@7.2
18.9@2.1
17.8@8.3
76..
33.9
83.6
112.7
.7.3
39.2
.0
10..
Au*ust : >ct !er Blac< %ert s l222
$epte=!er : A %e=!er 2222?ra#siti #al red-!r ;# earth
$epte=!er : A %e=!er Blac< earth (l a=)22222
Au*ust : Ba#uar" Red 'ria!le l a=
$ urces3 2 /arrell a#d >C,ee''e (2006)D 22 Ehttp3@@;;;.! =.* %.auFD 222 $c tt et al. (200.)D 2222 M sier et al (1897)D 22222 A$( G %er#=e#t (2009)D 222222 Ehttp3@@;;;.e#%ir #=e#t.#s;.* %.au@!i re*i #s@A rthH ast-La#d' r=.ht=F
Table :
Characteristics of some of the current maize growing regions in !"#

:in%aroy &Sou-h Burne--' 29.2@11.1 ;a(by &;ar(in% ;o1ns' 30.6@17.7 .a--on &#ore-on' 30.2@16.1 A-her-on &A-her-on Tab(e(an)' 29@16.9
/epresen-a-i0e si-e &1i-hin area' A0era%e )ai(y ma5+min -empera-ure 6C &Oc- 7 #ar'88 A0era%e )ai(y ma5+min -empera-ure 6C &Apr 7 Sep-'88 A0era%e mon-h(y rain9a(( mm &Oc- 7 #ar'8888 A0era%e mon-h(y rain9a(( mm &Apr 7 Sep-'88
21.2@6.1
23@6.3
23..@9.8
23.2@12.8
80.1
61
99.6
162.6
38.2
26.7
38..
12..
4)ea( p(an-in% -ime8
>ct !er Ba#uar"
Dece=!er
Au*ust Ba#uar"
A %e=!er Ba#uar"
E5amp(e o9 arab(e 'err s l %ert s l %ert s l 'err s l soi( -ype888 $ urces3 2 Hu*hes (2007d)D 22 Ehttp3@@;;;.! =.* %.auFD 222 Birch et al. (2003)
The area sown to mai+e in ustralia was greatest in 1:10 F 1:11 ;17820.0 ha< ;*olless 1:9:< and2 apart from a transient increase in 1:.0 F .1 ;1.92000 ha<2 gradually declined to around 502000 ha between the mid 1:90s F 2005! There is some indication that the area is increasing again with the 2008 F 0: forecast being 912000 ha ; & #$ 2008<! n indication of the demographics and farm si+e3yield associated with mai+e growing in ustralia is given in #obinson D Nirkby ;2002<! /n particular2 the average area grown to mai+e per farm is 250 ha in Southern 6S- and 100 ha in northern 6S- and southern Lueensland! verage yield ranges from 10 t3ha in southern 6S- to 9!5 t3ha in northern 6S- and 5!2 t3ha in southern Lueensland! /t is suggested that the differences between the regions reflects different degrees of irrigation between the regions! %ai+e is generally grown in rotation with other crops ;#obinson D Nirkby 2002<! *ultivars2 covering all end uses2 fall into two maEor categories ;*olless 1:9:<:
$arly38uick@maturing hybrids ;as low as :5 days *#%d< that are short ;appro(imately 250 cm<2 used mainly in inland areas and do not grow well in humid conditions2 have loose husks2 and tend to be susceptible to 6orthern 'eaf &light ;Drechslera turcica<! 'ate3slow@maturing hybrids ;up to appro(imately 1,5 days *#%< that are tall@ growing ;up to .00 cm<2 adapted to sub@tropical and tropical humid coastal regions2 have ears with well@covered husks2 and are generally resistant to D. turcica!
Cowever2 there is a continuum of differently maturing cultivars within these two e(tremes ;Cughes 2007d= 4arrell D )KNeeffe 2009<! lthough Z. mays subsp! mays is an important grain crop internationally ;see Section 2!2!,<2 its significance in ustralia is limited! 4or e(ample2 in 20082 only ,892000 t of grain mai+e were produced in ustralia ;as compared to 1,20,:2000 t of wheat grain<= appro(imately 122000 t were e(ported as compared to 727192000 t of wheat ; & #$ 2008<! 1etailed information on mai+e cultivation practices in ustralia is covered in publications by the 1epartments of "rimary /ndustries of every State and the 6T as well as in Eournal articles2 books and conference proceedings ;eg *olless 1:9:= "rice 1::9= &irch et al! 200,= 1"/D4 2007= 4arrell D )KNeeffe 2009= 1"/- @ Tasmania 2008= &irch et al! 2008<! The more important points are considered below!
d *#% ? *omparative #elative %aturity! /t is a measure of the time from planting to the stage when grain moisture content is suitable for harvest ;&irch et al! 200,= 4arrell D )KNeeffe 2009<! /t indicates the rate of maturity relative to a JstandardB hybridB and is used to determine where a hybrid should be grown!
1,
&./.#
Commercial propaga+ion
%ai+e is an annual plant and reproduces e(clusively by seed2 ie vegetative reproduction under natural condition does not occur! This is in contrast to other Zea spp! and the related Tripsacum spp!2 which may be perennial and reproduce not only by seed but also vegetatively by way of rhi+omes ;"aliwal 2000f<! Seed si+e varies from .2.00 seeds3kg to 22500 seeds3kg ;Cughes 2007d<! Seed for planting is usually treated with protective insecticides and fungicides ;*olless 1::2< ;see also Section 9<! /n developed countries2 including ustralia2 hybrid mai+e is usually grown! /mportant considerations regarding which cultivars3hybrids should be planted ;*olless 1::2= &irch et al! 2008< include:

maturity: length of growing season ;water availability2 temperature etc< end use ;grain vs silage vs sweet corn etc< yield lodging resistance disease resistance
%ai+e is an outcrossing plant species! /n order to be able to conduct directed crosses2 two re8uirements must be observed:
The female parent must be either male sterile or de@tasselede before pollen reaches maturity! /n mai+e2 three types of cytoplasmic male sterility are known: T2 * and S male sterility! &efore 1:902 Te(as@male sterile lines were used e(tensively! Cowever2 Te(as@male sterile lines are susceptible to southern corn leaf blight ;Helminthosporium maydis 6isikado and %iyake<2 race T2 which caused great losses to corn producers in the >S ! lthough * and S male sterile lines were discovered later ;and they are not susceptible to southern corn leaf blight<2 they are not important as growers have lost trust in male sterility ;reviewed in 1arrah et al! 200,= Sleper D "oehlman 2007<! *urrently2 detasseling is the preferred method to achieve controlled crosses! Seed breeding must be conducted in isolation from other mai+e plants: o 4eil and Schmid ;2002< reviewed the isolation distances re8uired or recommended by various organisations for seed breeding! /solation distances of up to 52000 metres ;>SS#2 seed of inbred lines< are listed2 with the maEority in the 200 F ,00 metre range! o The )$*1 re8uires 200 m isolation distance for the production of &asic and *ertified Seed for all mai+e varieties2 with purities of at least ::!5 and ::!0H2 respectively! 4or the production of &asic Seed of parental lines and single cross hybrids2 purities must be ::!:H! The production of *ertified Seed of hybrid varieties re8uires that the minimum varietal purity of both parents must be ::!8H ;)$*1 2008<! o /n the >S 2 an isolation distance of 201 m is re8uired for production of seed of ::H or higher purity! )ne option for reducing this distance is the planting of male border rows2 whereby
e Tassel is the term used to refer to the male infloresecence ;see Sections , D .<
one border row e8uals 5 m distance! Cowever2 even under those circumstances2 plantings for mai+e seed must be isolated by at least 1,. m from other mai+e! "redominant winds and storms in the seed breeding area are also to be considered ;1arrah et al! 200,<! ll recommended cultivars in ustralia are hybrids and originate from government or public breeding centres and private seed companies ;*olless 1:9:<! They are mostly produced under a voluntary seed certification scheme that ensures a minimum standard for purity2 seed germination and seed@borne disease! *ertified seed is classed according to its generation along the pedigree! &reeders seed is used to produce "re@ basic2 which is then used to produce &asic2 which in turn is used to produce 4irst 0eneration or *1 certified seed! %ost certified seed is *1 class grown from &asic seed ;Smith D &a(ter 2002<! %arket access to many countries2 including the $uropean >nion2 re8uires ustralian seed to be produced in accordance with the )$*1 ;)rganisation for $conomic *o@ operation and 1evelopment< Seed Schemes! The ustralian Seeds uthority 'imited oversees seed certification in ustralia and is licensed to administer the )$*1 Seed Schemes ; ustralian Seeds uthority 'td! 2007= ustralian Seeds uthority 'td! 2009a<! The figures for Z. mays certified seed production in ustralia under the )$*1 Seed Schemes are given in Table 5!
Table $% Zea mays certified seed produced in &ustralia under the '(C# Seed Schemes between 2))) and 2)) *
Year See) pro)uce) &*%' in -he 1 mon-hs en)in% <! =une 85810 105299 85801 15397 35693
!!! !!1 !! !!< !!>
2 Data ta<e# 'r = Australia# $eeds Auth rit" Ltd. (2006!)
&./.&
Scale of c.l+i-a+ion
Z. mays ssp! mays is grown as a grain crop for stockfeed2 food processing ;eg for breakfast cereals2 corn chips2 grits2 flour<2 industrial starch and popcorn ;4arrell D )KNeeffe 2009<! 0reen chop and ensiled mai+e may be used as supplementary feed in the dairy industry and also as feed for other animals2 including goats and horses ;#oche D 1alley 1::7= %oran 2007= "earson 2009= %c0regor 2009<! Sweet corn shows increasing domestic and international popularity in human consumption as a fresh2 canned or fro+en vegetable ;&eckingham 2009<! %ai+e plays a subordinate role in ethanol production in ustralia2 eg the maEor feedstocks for bioethanol production in Victoria are wheat2 sorghum and molasses ;%c*utcheon 2009<! Commercial ! maize in Australia *urrently2 no 0% mai+e is authorised for release in ustralia ;http:33www!ogtr!gov!au3<!
15
rain maize %ore than :5H of the grain mai+e produced is used domestically2 and the remainder for e(port! L'1 and 6S- account for most of the grain mai+e produced! & #$ ;2008< estimated that in the 2009308 growing season appro(imately 782000 ha were used for mai+e production! "ilage and green chop maize The si+e of areas used for silage and green chop mai+e production could not be obtained2 presumably because the planting areas per farm are not significant and most of the product is fed to animals on@farm! "#eet corn /n 20022 appro(imately 5.,. ha were used in sweet corn production in ustralia! ppro(imately :0H of this production area was located in 6S-2 L'1 and V/* ;&eckingham 2009<! &././ C.l+i-a+ion prac+ices
Z. mays ssp! mays may be grown as a dryland or irrigated crop ;&irch et al! 200,= 4arrell D )KNeeffe 2009<! /n ustralia2 mai+e is generally produced as a summer crop ;L'1: 1"/D4 2007= 6S-: 4arrell D )KNeeffe 2009<! %ai+e is suitable for mechanical cultivation during its entire development2 including harvesting! "lanting should commence when soil temperatures at planting depth reach 12G* or higher at :!00 am2 with an upward trend ;4arrell D )KNeeffe 2009<= for sweet corn planting at 1. F 17G* is recommended ;&eckingham 2009<! "lanting may also occur through to summer! Cowever2 planting times which will lead to flowering during the hottest period of summer should be avoided as pollen blasting ;damage to pollen< and conse8uently poor seed set may result! "lanting times should also try to avoid maturation during cool3cold weather as diseases and pests may cause poorer 8uality and3or lower yields! 4rost should be avoided during the entire life cycle of mai+e as should water stress2 the latter especially between two weeks to pollen shed and silking where it could cause disproportional yield losses! 0enerally2 mai+e is less water stress tolerant than other crops2 including sorghum ;4arrell D )KNeeffe 2009= &eckingham 2009<! %ai+e plants grow best in well@draining2 nutrient@rich soils with a pC ;*a*l2< between 5!5 and 9!0! %ai+e is not very tolerant of saline soils! The nutrient status of the soil is important for the productivity of mai+e and substantial amounts of nutrients are removed from the soil by harvesting mai+e cobs or plants! Therefore2 soil tests for various nutrients should be carried out to inform the grower before planting! 6utrient availability varies with soil type! 6itrogen is yield limiting in mai+e production! The amount of nitrogen that needs to be added to the soil depends on various factors2 including cropping history and yield target! "hosphorus2 potassium2 +inc and molybdenum application may also be necessary! &anded application of nitrogen and phosphorous may be carried out at sowing to reduce the chance of damage to the seedling by fertiliser burn and to make those nutrients available during the early stages of development2 when they are of greatest importance ;4arrell D )KNeeffe
2009<! /n 6S-2 mai+e was grown mainly in the 6orthern and Southern inland2 the 6orth *oast and the Tablelands in the 2007309 season ;4arrell D )KNeeffe 2009<! /n L'12 mai+e can be grown in 6orth L'12 *entral L'12 South &urnett2 the 1arling 1owns and %oreton ;Cughes 2007d<! 'ists of hybrids that can be grown in those areas are provided by the above authors! list of sweet corn hybrids for L'1 is provided in &eckingham ;2009<! /n Victoria2 mai+e production may be followed by green bean ;$haseolus vulgaris '!< plantings ;1imsey 1::.<! Sweet corn may be grown in rotation with %rassica crops2 including canola and lettuce2 or in rotation with lucerne ;!edicago sativa '! ssp sativa< and other vegetable crops2 in which case it may be followed by gra+ing oats ;&eckingham 2009<! rain maize 0rain mai+e usually takes appro(imately 1,0 to 150 days from planting to harvest! *rop density for irrigated mai+e grain crops is typically around 702000 F 802000 plants per hectare! 4or dryland crops density mainly depends on rainfall2 varying from appro(imately 202000 F ,02000 per hectare for dryer inland area up to appro(imately .52000 F 552000 per hectare for coastal or other high rainfall areas! Seeds are planted at depths of , F 5 cm in rows with spacing varying between 95 and 110 cm! /n dryland production systems2 single or double row skipping on 100 cm rows may increase yields! /rrigated mai+e may be irrigated pre@ or post@plant2 depending on soil characteristics! 1ryland mai+e should be planted into soil that is wet to a depth of at least one metre! %ai+e has low tolerance to water stress2 and high yielding hybrids may use up to 850 mm of water during the growing season! *rops re8uire continued water supply2 from irrigation or rain2 throughout their growth! -hen physiological maturity is reached2 the moisture content of the kernels is usually 28 F ,.H! 0rain storage is best if grain moisture is appro(imately 12H! Therefore2 dry down is re8uired! 1ry down of grains on the plant can continue as long as conditions are not too cool2 which could result in infection with mycoto(in producing fungi! Carvest usually commences at a moisture content of 18H and further artificial drying is re8uired! Typical grain yields2 depending on the region2 are appro(imately 2 F 8 t3ha for dryland mai+e crops and appro(imately 5 F 1, t3ha for irrigated crops! /nformation in this section is based on 4arrell D )KNeeffe ;2009<! "ilage and &orage maize Silage and forage mai+e is grown similarly to grain mai+e2 e(cept that planting density should be increased to 502000 F 702000 plants per hectare for dryland cultivation and 902000 F 1002000 plants per hectare for irrigated cultivation to obtain optimum yields! 'ater maturing hybrids can be grown as mai+e with ears at the dough stage of grain development is best for fodder and silage production! Silage is a moist preserved fodder2 in which anaerobic fermentation improves digestibility and preserves most of the nutrient content of the ensiled material! Therefore2 silage and 19
forage mai+e are ideally harvested when the dry matter content is between ,0 and ,5H ;*olless 1::2<! %ai+e silage production re8uires chopping2 compaction and careful packing of the plant material so as to e(clude air! Mield for silage is about si( to ten times that for grain ;*olless 1::2<! The information in this section is based on 4arrell D )KNeeffe ;2009<2 e(cept where indicated otherwise! "#eet corn Sweet corn differs from other forms of mai+e as the kernels have high sugar content at the milk and early dough stage of grain development due to mutation3s in the starch synthesis pathway2 eg the sugary' or shrun(en) allele is present in the sweet corn line! *ultivation is similar to that for grain mai+e2 although low seedling vigour2 due to reduced seed starch2 re8uires more precise early season management! *rop density ranges from appro(imately ,52000 plants3ha under marginal conditions to 902000 plants3ha for well managed irrigated crops! *obs are ready to harvest when the kernels reach the milk or very early dough stage! This occurs appro(imately 95 F 105 days after planting ;21 F 28 days after silk emergence<2 depending on variety2 climate and soil moisture! Carvesting may be by hand or machine! lternatively2 the crop may be harvested for baby corn2 usually by hand within two days of silks emerging2 when ears are . F 10 cm long! The early harvest of sweet corn ;ie prior to kernels reaching physiological maturity< allows two crops per season in warmer areas! Sweet corn is preferred to field mai+e for production of green ears for fresh consumption in ustralia and other developed countries! ppro(imately 80H of sweet corn produced in ustralia is processed ;canned or fro+en<2 with the remainder going to the fresh food market! /n 20022 appro(imately 50H of the national production2 ie .02000 t2 was in 6S-2 followed by Victoria producing ,02000 t and L'1 producing 92000 t! verage yield of sweet corn is appro(imately 19 t3ha in 6S- and L'1! The information in this section is based on &eckingham ;2009<! &.1 Crop !mpro-emen+
The fact that mai+e has high commercial value and is the main staple in the diet of millions of people in sub@Saharan frica and 'atin merica means that there is much to be gained from improvement programmes! number of international organi+ations and networks are involved in coordinating the improvement of mai+e growing in poorly developed countries! Significant among these are: */%%MT ;*entro /nternacional de %eEoramiento de %aO+ y Trigo< F which was established in %e(ico and now has regional offices in many countries! )ne of the aims of the 0lobal %ai+e "rogram is to use I=ai-e *e#etic res urces t pr %ide
di%erse5 hi*h-"ieldi#* %arieties that ;ithsta#d i#'ertile s ils5 dr u*ht5 i#sect pests5 a#d diseasesC (http:33www!cimmyt!org3english3wpp3afrPlivelih3inde(!cfm). MaJ r pr Jects i#clude II#sect Resista#t Mai-e ' r A'ricaCD IDr u*ht ? lera#t Mai-e ' r A'rica I#itiati%eCD I(ater K''icie#t Mai-e ' r A'ricaC. /A> Hr p & Grassla#d $er%ice (AGLH) - supports mai+e production improvement
through 4 ) 4ield "rogramme proEects in many countries2 including fghanistan2 &angladesh2 *hina and $ritrea ;http:33www!fao!org3ag3 0"3 0"*3doc3crops3.a@
2!html<! The Tropical sian %ai+e 6etwork ;T %6$T< F which was established in 1::, by 0"* to strengthen collaboration between national institutions in mai+e research and development and to increase mai+e production and productivity2 particularly through hybrids! /n 1::8 */%%MTKs regional office in &angkok took responsibility for coordinating technical activities2 including hybrid trials! The sian %ai+e &iotechnology 6etwork ; %&/)6$T< F which was established in 1::8 as a collaborative research2 training2 and information network2 aiming to help mai+e programmes in *hina2 /ndia2 /ndonesia2 "hilippines2 Thailand and Vietnam ;0eorge et al! 200.<!
$(tensive genetic and cytogenetic studies have contributed to an understanding of mai+e and are important tools in mai+e improvement! /n 20052 several years after launching the concept of the %ai+e 0enome Se8uencing "roEect ;*handler D &rendel 2002<2 the >S 1epartment of griculture2 1epartment of $nergy2 and 6ational Science 4oundation announced a Eoint >SQ,2 million programme for se8uencing the mai+e genome ;also see online information at Rhttp:33www!mai+egenome!org3S<! *onsiderable work laying a foundation for this proEect had already been completed ;see reviews in Caberer et al! 2005= #abinowic+ D &ennet+en 2007= %essing D 1ooner 2007<! The %ai+e 0enetics *ooperation ;http:33mai+ecoop!cropsci!uiuc!edu3< operated by the >S 1epartment of griculture gricultural #esearch Service collects2 maintains and distributes seeds of mai+e genetic stocks and provides information through the %ai+e 0enetics and 0enomics 1atabase ;http:33www!mai+egdb!org3<! large number of mai+e mutants have been identified and allow the identification of genes that are important in controlling mai+e plant development ;for e(ample2 mai+e morphogenesis mutants @ Sheridan 1:88<! &.1.# Bree,ing
wealth of information regarding the improvement of mai+e is available ;see "aliwal et al! 2000= Sleper D "oehlman 2007= 'ee D Tollenaar 2009<! The sections below indicate important trends in mai+e breeding! Conventional *reeding %ai+e is predominantly outcrossing ;cross@fertilising<2 which has its advantages and disadvantages for plant breeders and growers! 4or e(ample2 outcrossing species are characterised by a wider genetic base and are 8uick to adapt to changes in the environment2 whereas predominantly inbreeding species have the advantage of consistency and relative genetic uniformity within a given cultivar! %ai+e possesses diverse morphological and physiological traits resulting from a wide genetic base available for selection and2 therefore2 is adapted to a wide range of conditions ;"aliwal 2000a<! /t has been selected for desirable traits by farmer breeders for more than 72000 years2 and by professional breeders for appro(imately 150 years ;"aliwal 2000a= "iperno D 4lannery 2001= %atsuoka et al! 2002<! /ts long history of domestication has lead to the ;co<e(istence of a wide variety of primitive ;largely unimproved< races2 land races ;local varieties adapted for local conditions<2 cultivars improved and maintained by farmers2 and highly@improved professionally@bred open pollinated varieties ;"aliwal 2000a<! *urrently2 the highest yielding mai+e cultivars
1:
are various types of mai+e hybrids! &reeding has been2 and is being2 used to improve or alter traits such as plant height2 ear number2 yield2 maturity2 kernel properties2 and disease and pest resistance ;reviewed in "aliwal et al! 2000= Sleper D "oehlman 2007<! /n addition2 plant breeding is also aimed at increased nutrient content in cultivated field mai+e varieties ;reviewed in Ihu et al! 2009<! Speciality mai+e varieties are also being bred for sweet corn2 high@oil content2 high@8uality protein2 popcorn and silage! /n developing countries2 farmers select and maintain mai+e varieties adapted to specific local uses and conditions ;"aliwal 2000a<! /n the 1:.0s2 local mai+e varieties were collected in countries of *entral and South merica by scientists from the >S and %e(ico2 and those with similar morphological characteristics were grouped into land races! This classification allowed breeders to easily access mai+e germplasm with a particular trait of interest! Cowever2 many of these collections have been lost and new collections have been made! *urrently2 over 1,2000 germplasm accessions are stored at */%%MT in %e(ico2 with duplicate storage elsewhere ;reviewed in 1arrah et al! 200,= Sleper D "oehlman 2007<! +pen-pollinated cultivars )pen pollinated cultivars result from random pollination of ovules without pollen control! s each ear is pollinated by a random mi(ture of pollen2 it is possible that each resulting seed will grow into a plant that is genotypically and phenotypically different to the plants developing from the other seeds on the same cob! Therefore2 open pollinated cultivars are characterised by their hetero+ygosity and high genetic variability! )pen pollination and mass selection of mai+e has been and is being used by farmer@breeders to improve important visible plant characteristics2 such as maturity2 seed characteristics and plant height! "lants derived from open@pollination are generally not uniform and one of the aims in breeding proEects was to control the pollen source to increase plant uniformity ;reviewed in 1arrah et al! 200,= Sleper D "oehlman 2007<! /mproved mai+e breeding using open pollination includes ear@to@row breeding and variety hybridisation! /n ear@to@row breeding2 seeds are harvested2 labelled and stored separately from each ear! Some seeds from each ear are planted in individual rows= remaining seed is stored! $ach row is evaluated for improved characteristics and combined remnant seed from superior performing rows is planted in a single plot in the ne(t growing season allowing for open pollination to occur! $ar@to@row selection is repeated over a number of generations! Variety hybridisation includes planting of two different varieties of open@pollinated mai+e in one plot2 whereby the plants from one variety are detasselled! This breeding method results in improved yields in the following generation! Cowever2 it was never widely used in mai+e breeding ;reviewed in 1arrah et al! 200,= Sleper D "oehlman 2007<! Hy*rid *reeding Cybrid breeding was first used in the early 1:00Bs by professional breeders in the >S ;see discussion in 'ee D Tollenaar 2009<! n important re8uirement for hybrid breeding was that the source of pollen in crosses could be controlled by the breeder!
This was achieved by either detasseling of the female parent or by the use of cytoplasmic male sterile lines as the female parent in combination with a male fertility restorer line! /nitially2 the main goal of plant breeders was the selection for genotypes that would survive the inbreeding process that is followed by hybrid seed production2 as inbreeding generally leads to an accumulation of deleterious alleles within a given population! n important consideration in hybrid breeding is the combining ability of individual inbred lines as generally few combinations results in superior hybrids! /n single crosses2 two inbred lines are used in the production of hybrid seed ;line ( line & ? ( &<= the plants derived from the hybrid seed show increased vigour and yield ;heterosis<! s inbred lines generally showed low vigour and yield2 single crosses were substituted by double crosses2 in which four unrelated inbred lines are used in the production of the two parent lines! The two parent lines are then crossed resulting in double cross hybrid seed ;; ( &< ( ;* ( 1<<! n alternative to the double cross is the modified single cross involving the recombination of two related inbred lines in the generation of the female parent2 which is then crossed to another inbred line resulting in the hybrid seed ;; B ( BB< ( &<! Three@way crosses are similar to modified single crosses2 with the e(ception that three unrelated inbred lines are used in the generation of hybrid seed ;; ( &< ( *<! s inbred lines were improved over time2 single cross hybrids are now used widely in mai+e breeding! maEor difficulty in the evaluation of plant material is the separation of genetic and environmental effects on some ;especially polygenic< traits! This difficulty was overcome by conducting test crosses over a number of seasons in a range of environments ;reviewed in 1arrah et al! 200,= Sleper D "oehlman 2007<! *urrently2 mai+e hybrids play an important role in the mai+e production in developed countries! The entire process from initial hybrid cross to commercial release takes appro(imately 5 years ;1uvick D *assman 1:::<! The most important selection criteria used by commercial mai+e breeders in developed countries such as the >nited States are yield and yield stability and the relatively short time that hybrids remain on the commercial market ;less than 10 years on average< is due to their replacement by more high@yielding hybrids rather than any problems with disease or insect susceptibility ;1uvick D *assman 1:::<! Several mai+e traits have changed significantly in association with the selection efforts for increased yield and include increased grain starch: protein ratio2 decreased tassel si+e2 reduction in barreness at high density2 and reduced rate of leaf senescence during grain filling ;1uvick D *assman 1:::<! !utation *reeding %utation breeding in mai+e encompasses a number of approaches2 including spontaneous mutation2 cell@ or tissue@culture induced mutation ;also referred to as somaclonal variation<2 chemical mutagenesis2 transposon mutagenesis and irradiation mutagenesis! Spontaneous mutations occurring within plant cells may lead to new and useful characteristics2 which are employed in crop breeding! /n mai+e2 spontaneous mutations have been e(ploited commercially! 4or e(ample2 all the maEor types of mai+e2 including dent2 flint2 flour and pop mai+e as well as sweet corn were available to 6ative mericans by the time *olumbus arrived in the mericas ;reviewed in 21
&oyer D Cannah 1::.<! %utagenesis has also been used to increase the range of characters available for selective breeding! 4or e(ample2 somaclonal variation has been used successfully in the development of commercially available imida+olinone@ tolerant mai+e varieties ; nderson D 0eorgeson 1:8:= 6ewhouse et al! 1::1= 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 are discussed below! Singletary ;200,< has provided a detailed review on this subEect! %onocot plants such as mai+e are generally more difficult to transform than dicot species! Cowever2 both efficient transient and stable transformation protocols for mai+e have been developed and improved over time2 including Agro*acterium@ mediated transformation2 transformation through protoplast fusion2 particle bombardment and silicon carbide whiskers ;eg 0ordon@Namm et al! 1::0= 0ould et al! 1::1= 4rame et al! 1::.= /shida et al! 1::7= -right et al! 2001= 4rame et al! 2002<! The latest advances of genetic modification in mai+e include the development of circular mai+e minichromosomes ;%%*s<2 which can facilitate autonomous replication of the introduced 16 ! dvantages of this type of genetic modification include that the likelihood of position effects may be much lower than when using gene insertion methods and that larger 16 fragments can be delivered and e(pressed ;*arlson et al! 2009<! The maEor focus in the production of 0% plants has been on resistance to insects and tolerance to herbicides! 4or e(ample2 resistance to corn rootworm has been achieved by using cry@genes from %acillus thuringiensis and tolerance to the herbicides glufosinate ammonium and glyphosate using the pat@gene from "treptomyces viridochromogenes and the cp,epsps-gene from Agro*acterium spp!2 respectively! nother focus in early mai+e research was to increase lysine levels in mai+e2 which was achieved by transformation with the cordapA gene from Coryne*acterium glutamicum encoding lysine@insensitive dihydrodipicolinate synthase! These e(amples are approved for commercial release overseas and for use in human food in ustralia and overseas ;4S 6I 2002= "C/S3>S1 2002= "C/S3>S1 2005= "C/S3>S1 2007a= "C/S3>S1 2007b= "C/S3>S1 2009<! %ai+e grains2 like those of other cereals2 do not contain the amino acids lysine2 tryptophan and methionine at levels that are sufficient in the diet of humans and other monogastric animals ;as reviewed in 'ai D %essing 2002< ;see also discussion in Section 5!1!2<! *urrent molecular studies are aimed at addressing those deficiencies!
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
seasonal root system bearing a single erect stem ;culm< made up of nodes and internodes2 although some cultivars may develop elongated lateral branches ;tillers<! %any temperate cultivars are shorter than tropical ;and subtropical< cultivars! 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 is distichous ;reviewed in $sau 1:99b<! $ach leaf consists of a sheath surrounding the stalk and an e(panded blade connected to the sheath by the blade Eoint2 or collar! The mature plant can have up to appro(imately ,0 leaves2 with considerable variation in leaf number2 si+e and orientation between mai+e races! 0enerally2 tropical mai+e plants develop more leaves than temperate cultivars! The epidermis is the outermost layer that can be discerned in cross@sections of the stalk! Several layers of sclerenchyma tissue are underneath it and increase the strength of the stalk! The arrangement of vascular bundles is comple( and they appear scattered throughout the parenchyma tissue that constitutes the remainder of the cross@ section2 although they are arranged more loosely towards the middle ;reviewed in $sau 1:99b<! %ai+e F like many plants that evolved under tropical conditions F is a *. plant and2 therefore2 is more efficient at utilising carbon dio(ide than *, plants! These physiological characteristics are reflected in leaf morphology down to the microscopic level! 4or e(ample2 bundle sheath cells are richer in chloroplasts than mesophyll cells! The chloroplasts are also larger than those of mesophyll cells ;reviewed in $sau 1:99a<! /.& Repro,.c+i-e morphology
%ai+e is a monoecious plant: one or more lateral branches2 the shanks2 develop in the leaf a(is of the plant! They terminate in a female inflorescence2 an ear ;see 4igure 1<! The male inflorescence2 the tassel2 forms at the top of the stem! >sually one or two lateral shoots in the upper part of the plant develop into female inflorescences! The shank consists of nodes and short internodes2 the lengths of which vary between mai+e races! The ears arise from a(illary bud apices! The ear is covered in a number of leaves called husks! Those leaves differ in appearance when compared to those on the stalk: they surround and protect the developing ear! -here mai+e is left to dry in the fields2 more husks are generally desired to protect the grains from birds and insects! -here mai+e is harvested earlier2 it is often desirable for a cultivar to have a lower number of thin husks! The ear does not usually show any lateral branching! The thick a(is of the ear2 the cob2 bears an even number of rows ;between . and ,0< of ovaries2 each containing a single ovule! The number of ovules that will develop into kernels ranges from ,00 F 12000 and is dependant on the cultivar3variety as well as factors occurring later in development ;"urseglove 1:92<! The silks of the mai+e ear are the stylar canals of the mature ovaries!
2,
Silks ;long green strands< )varies containing aborted ovules ;under the silks<
%ature kernels
*igure 1% +ature female inflorescence of a sweet corn culti,ar (the husks have been removed to expose the cob)%
The apical meristem of the stem develops into the tassel2 a prominent branched structure at the top of the plant consisting of a central spike and a variable number of lateral branches ;up to appro(imately .0< bearing the male flowers! The peduncle of the tassel grows vigorously2 pushing the tassel out of the top of the plant!
SECT!"N 1
1.# 1.#.#
(E*EL"P
Repro,.c+ion
ENT
Ase3.al repro,.c+ion
>nder natural conditions mai+e reproduces only through seed! 1.#.& Se3.al repro,.c+ion
%ai+e is a 8uantitative short@day plant but some cultivars have low or no sensitivity to daylength ;Niniry et al! 1:8,<! /n those cultivars that are photoperiod sensitive ;mainly those that are late@maturing<2 flowering may be delayed when the photoperiod is greater than a critical threshold value ranging from 10 F 1,!5 h ;Niniry et al! 1:8,<! Those adapted to the tropics may show delayed maturity if grown in more temperate areas with longer days ;&irch et al! 200,< ;see also Section .!5<! >nless stated otherwise2 the following description is adapted from "aliwal ;2000f<! /nitially male and female inflorescences have primordia of bise(ual flowers! Cowever2 during their development2 primordia of stamens abort in the a(illary inflorescences2 and primordia of gynoecia abort in the apical inflorescence! The apical meristem elongates once the leaf primordia are initiated! /t is transformed into a reproductive meristem that develops into the tassel! "ollen is shed from the tassel continuously for a week or more as upper and lower florets in the male spikelets show developmental differences and the spikes mature asynchronously! The female inflorescences ;ears< arise from a(illary buds and bear flowers in rows along the cob! 1evelopment of the flowers and the ovules on the ear proceeds from
the base upwards ;acropetal<! 4rom each flower a style begins to elongate towards the tip of the cob2 forming long threads2 or silks! Silk development begins from the flowers near the base of the ear and proceeds towards the tip over several days! #eceptive silks emerge over the husks over a period of three to five days and can grow to more than ,0!5 cm length2 e(tending beyond the end of the husks! The silks have short hairs2 trichomes2 which form an angle to the stylar canals and help harbouring pollen grains! #eceptive silks are moist and sticky! %ai+e is often considered as protandrous ;anthers reaching maturity before the gynoecium<2 as anthers on the spikelets on the upper part of the central spike protrude out of the florets and start shedding pollen one or two days ;under optimal growth conditions< before the silks emerge above the husks! Cowever2 the gynoecium matures and the silks become receptive before they appear above the husk tips! >nder any stress ;especially water stress F see Section 7!,<2 the interval between pollen release and silk emergence increases! Selective breeding is aimed at producing many mai+e varieties with a reduced interval2 to ensure good seed set! 1.& 1.&.# Pollen ,ispersal4 pollina+ion an, o.+crossing ra+es Pollen
%ai+e pollen is relatively large when compared to other grass pollen! 4or e(ample2 &alta+ar et al ;2005< reported pollen diameters of :. to 10, M=! %ai+e pollen is surrounded by a double layer consisting of e(ine and intine2 and is2 therefore2 well protected ;reviewed in *heng D "areddy 1::.<! Cowever2 temperatures above ,5G* at pollen release may result in pollen blasting2 a maEor problem in some mai+e growing areas in ustralia that may result in poor seed set ;barrenness= 4arrell D )KNeeffe 2009<! single plant may produce up to 2 ( 107 pollen grains per day ;Aaros+ et al! 200,< and between 7 ( 107 F 25 ( 107 in total2 depending on the variety ;&annert D Stamp 2009<! t eight plants per s8uare metre ;at the high end for crop density<2 this would e8uate to appro(imately 200 million pollen grains released per s8uare metre! 'una et al! ;2001< reported that pollen viability as measured by the ability to produce seed decreased to 0H after atmospheric e(posure for 2 hrs near San Aose de Valle in %e(ico! /n that study2 80H of pollen lost viability within 1 hr! Similar results were obtained in a study by &alta+ar et al ;2005< that was conducted in Tapachula in %e(ico: 78H to 8.H of pollen was dehydrated after 1 hr of atmospheric e(posure! The study by 'una et al ;2001< also showed that high humidity prevented loss of viability: at high relative humidity2 pollen viability declined by 58H within 1 hr2 whereas it declined by :7H within 1 hr under lower atmospheric humidity! ylor ;200.< carried out in-vitro germination tests to investigate pollen viability under a number of environmental conditions in 6ew Caven in the >S ! /t was found that pollen viability was relatively insensitive to solar radiation and decreased most with loss of moisture! decrease of pollen viability to 50H was observed between 70 min and . hrs2 depending on environmental conditions! 4eil and Schmid ;2002< mentioned reports of mai+e pollen being viable between 20 min and 2. hrs! Various authors have reported that visible characteristics can be used as an indication of pollen viability: fresh pollen appears spherical in shape and white2 whereas non@ viable pollen is collapsed and yellow ;eg 'una et al! 2001<! 25
1.&.&
Pollen ,ispersal an, pollina+ion
%ai+e is normally outcrossing and the rate of self pollination is appro(imately 5H ;reviewed in Sleper D "oehlman 2007<! 1uring normal cross@pollination2 the pollen is shed mainly during mid@morning ;'una et al! 2001<! "ollen is shed continuously for a week or more from each plant2 starting appro(imately 1 to , days before silk emergence ;reviewed in "aliwal 2000f= Sleper D "oehlman 2007<2 with a crop as a whole producing pollen over a 1. day period! 1espite the short viability period of individual pollen grains2 the temporal spread in both pollen shed and silking of individual plants within a field means that2 on a whole field basis2 cross pollination between a donor field and a receptor field could occur over a 9 day period ;&annert D Stamp 2009<! The hori+ontal settling speed of mai+e pollen is in the range of 21 F ,2 cm3s2 depending on how much dehydration of the pollen has occurred ; ylor 2002<! /n plants where the tassels are at a height of about 2!5 m and the silks are at a height of about 1 m2 a settling distance of appro(imately 1!5 m is re8uired for pollination between adEacent plants and could take 5 s under ideal conditions ;&annert D Stamp 2009<! The vertical movement of pollen in thermals and air turbulence could e(tend dispersal distances but only in those areas where conditions do not favour hori+ontal dispersal ;&annert D Stamp 2009<! /nsect pollination in mai+e has not been reported2 although 4eil and Schmid ;2002< mention reports of mai+e tassels being visited by honey bees! Cowever2 as honey bees have not been observed on female inflorescences2 pollination of mai+e by honey bees has been ruled out! 1.&./ ".+crossing ra+es an, isola+ion ,is+ances
4eil and Schmid ;2002<2 &rookes et al! ;200.< and Sanvido et al! ;2008< have recently reviewed literature on mai+e pollen dispersal and outcrossing rates! These reviews as well as a number of other publications ;/ngram 2000= 'una et al! 2001= Stevens et al! 200.= Calsey et al! 2005= /reland et al! 2007= %esseguer et al! 2007= &annert D Stamp 2009< point out a number of biotic and abiotic factors that can influence outcrossing rates in mai+e2 including

synchrony in flowering time in pollen donor and receptor ambient temperature at pollen shed relative ambient humidity at pollen shed wind speeds ;and turbulences< and direction of prevailing winds topography of the terrain in 8uestion distance between pollen donor and pollen receptor competition between foreign pollen and pollen shed on the receptor plot ;the scale of pollen emission from the donor relative to the si+e of the pollen receptor plot as well as the absolute si+e of the receptor plot F the larger the receptor plot2 the more the foreign presence is diluted and therefore the lower the average outcrossing rate per plot ; *#$ 200,< ie outcrossing is highest at plot margins than at the plot midpoint< e(perimental design2 eg physical arrangements between pollen
donor and receptor ;eg2 are pollen donor and receptor arranged in a concentric fashion2 neighbouring or distant from each otherT<2 method of determining outcrossing rates ;/s pollen collected or are resulting seeds sampledT< agronomic practices such as detasseling2 use of border rows2 cytoplasmic male sterility
The dependence of outcrossing rates on a large number of factors may e(plain the range of results obtained in the scientific literature and the range of isolation distances recommended by various organisations with regard to seed production and food3feed production! ;eg see discussion in Sections 2!,!1 and :!1<! s an indication of cross@ pollination rates2 a recent Swiss study ;&annert D Stamp 2009< found that at distances of 50 F .2500 m between pollen donor and receptor fields2 the cross pollination rate in a field was never more than 0!02H! /n 8:H of cases2 cross@pollination was represented by a single fertili+ation event in an ear! /n ustralia2 the following isolation distances for the production of speciality mai+e currently are recommended:
/n L'12 the production of sweet corn re8uires either a .00 m isolation distance from all other mai+e or staggering mai+e plantings so that there are at least 1. F 21 days between pollination times to all other mai+e ;&eckingham 2009<! /n 6S-2 the production of speciality mai+e2 eg wa(y2 white and popcorn varieties2 re8uires 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<2 isolation distances of ,00 metres ;0ermany= sugar mai+e for consumption< to 12700 metres ;*anada= sugar mai+e for consumption< were recommended! /solation distances recommended for seed breeding are covered in Section 2!,!1! 1./ Em5ryogenesis4 fr.i+6see, ,e-elopmen+ an, see, ,ispersal
"ollen grains germinate immediately after settling on the silk! The pollen tube takes between 12 and 2. hours to reach and fertilise the ovule ;reviewed in Sleper D "oehlman 2007<! >pon completion of fertilisation the silk detaches from the ovary and dries out! 1./.# Em5ryogenesis
The following information is adapted from Sass ;1:99< and Sheridan and *lark ;1::.<! /n the first phase of embryogenesis2 irregular cell divisions result in a proembryo consisting of appro(imately 12 F 2. cells within 100 hrs after fertilisation! The initial basal cell divides into a number of large2 vacuolated suspensor cells! &y the same time2 the initial apical cell has given rise to : F 18 small cells that are densely filled with cytoplasm! Starting appro(imately 8 F : days after fertilisation2 the second phase of embryogenesis commences2 which leads to the establishment of meristems and the embryonic a(is by appro(imately 1, days after fertilisation ;transition embryo<! 29
ppro(imately 1. F 15 days after fertilisation2 the coleoptilar embryo is established2 characterised by the differentiation and growth of scutellumf2 coleoptileg2 coleorhi+ah and root and shoot apical meristem ;see 4igure ,<! &y appro(imately 17 days after fertilisation2 the first leaf primordium arises! fter this time2 the embryo is referred to as stage 1 embryo! /t is now appro(imately 1 mm long and weighs appro(imately 1 mg! 1uring the last stage of embryogenesis ;during ,0 F .0 days after the first leaf primordium appears<2 embryo growth continues2 more leaf primordia and a primary and one or more secondary root primordia develop2 and the whole embryo is enveloped by the e(panding scutellum! Storage products also accumulate within the embryo during the final phase of embryogenesis2 especially in the scutellum! t kernel maturity the embryo may have a fresh weight of appro(imately 50 mg ;Sheridan 1:88<! 1./.& Fr.i+6see, ,e-elopmen+
The fruit of mai+e is a caryopsis2 a dry indehiscent single@seeded fruit ;4igure 2<!
*igure 2% +ature maize car-opses [photo credit: Steve Hurst @ USDA-N !S "#AN$S Database% "rovided b& A S S&stematic 'otan& and (&co)o*& #aborator&+ 'o)ivia% !ochabamba+ (http:,,p)ants+usda+*ov,-ava,pro.i)e/s&mbo)012(A)3
The pericarp ;ovary wall< and testa ;seed coat< are fused to form the fruit wall and because of this tight adhesion between fruit and seed2 the two structures actually appear to be a single structure! This structure is commonly referred to by a number of interchangeable terms F fruit2 kernel2 grain and seed! The kernels are composed of three main parts @ the embryo2 the endosperm and the fruit wall ;see 4igure ,<! The
f The scutellum is the name given to the cotyledon of grasses ;#aven et al! 1:::<! /t functions in the absorption of food by the embryo from the endosperm ;see Section 5!1!,< g The coleoptile is a leaflike sheath that protects the shoot meristem h The coleorhi+a is a sheath that protects the root meristem
number of kernels per ear and the number of ears that develop is established at2 or shortly after2 pollination ;1uncan 1:95<!
*igure 3%
#iagrammatic representation o. a )on*itudina) section throu*h a mature mai4e kerne) ( aven et a)+ 5666)+ e.er to text .or de.initions o. the various parts+
The rate of cob development is dependant on environmental factors2 such as climate ;particularly temperature< and genetic factors2 such as cultivar ;see discussion in 6orman et al! 1::5= Sleper D "oehlman 2007<! The grain filling period in mai+e is appro(imately 8 weeks in length ;'ee D Tollenaar 2009<! s reviewed by 4arnham et al! ;200,<2 the following stages can be discerned during kernel development: blister stage2 milk stage2 dough stage2 ;dent stage in dent mai+e varieties< and physiological maturity! "hysiological maturity is generally reached appro(imately 55 F 75 days after silking!
4or the first 10 F 1. days after fertilisation2 the husks2 cob and shank develop rapidly and soon after2 nutrients are accumulated in the developing kernels ;4arrell D )KNeeffe 2009<! The small blister@shaped kernels are filled with a clear fluid! The milk stage is characterised by the presence in the kernels of a white fluid with high sugar content! 1uring dough stage2 the kernels are filled with a white paste and2 starting at the tip of the kernel opposite the embryo2 starch is deposited! This starch deposition is
2:
visible2 as a Jmilk lineB develops ;soon after denting in dent mai+e varieties< between hard ;starch@filled< and soft ;paste@filled< phases of the endosperm! milk line of 0 indicates that no starch deposition has occurred2 whereas a milk line of 5 indicates that starch deposition is completed! %onitoring the progress of the milk line towards the base of the kernels allows estimating the time to physiological maturity!
"hysiological maturity is reached once ma(imum dry matter has been accumulated in the kernels and their moisture content is appro(imately ,0 F ,8H! Jblack layerB is formed underneath the tip of the kernel that is attached to the cob! /t seals off the kernel from the remainder of the plant!
/nformation on harvesting and post@harvest can be found in a number of sources ;see *olless 1::2= 'afitte 2000b= 4arnham et al! 200,<! The moisture content of the grain at physiological maturity is usually above ,0H! fter physiological maturity is reached2 kernels continue to loose moisture and in moist tropical environments2 harvest should not proceed until moisture content has been reduced to appro(imately 25H ;"aliwal 2000e<! >sually2 mechanical harvesters perform best at a kernel moisture content of appro(imately 18 F 2.H! fter harvest2 mai+e grains must be dried artificially to no more than 1.H moisture in order to minimise infestation with pests and development of diseases during storage! rtificial drying should be carried out at temperatures below .:G*! The rate of development of mai+e is linked to temperature ;over 2. hours< rather than to photosynthesis which is governed by temperature only during daylight hours ;1uncan 1:95<! The concept of thermal time ;e(pressed in units of day degrees< can be used to categorise mai+e cultivars into early@ and late@maturing ;see also discussion in Section 2!,<! &asically2 thermal time is a measure of accumulated temperature that is re8uired for a phenological character ;such as flowering< to take place! 'afitte ;2000b< and 6orman et al! ;1::5< note that tropical mai+e cultivars have a lower yield than temperate cultivars because2 while temperatures are higher in the tropics2 the plants have a much shorter time to maturity! Xenia "ollen can have an immediate effect on kernel characteristics2 a phenomenon that is known under the name JUeniaB ;reviewed in Sleper D "oehlman 2007<! The underlying cause for it is the fertilisation of the diploid polar nucleus by the haploid vegetative sperm nucleus2 resulting in triploid endosperm cells! s the endosperm comprises appro(imately 80H of the mature mai+e grain ;reviewed in &oyer D Cannah 1::.<2 kernel characteristics depend on the genotypes of both female and male parent! $ndosperm characteristics that e(hibit Uenia include endosperm colour ;eg yellow vs white<2 wa(y vs non@wa(y endosperm2 aleurone colour ;purple vs colourless<2 starchy vs sugary endosperm and non@shrunken vs shrunken endosperm! 1././ See, ,ispersal
The mai+e cob lacks any abscission layers between its basic units and therefore the cob remains intact at maturity ;1oebley et al! 1::0<! Thus the tightly held grains are unable to be dispersed and confer a low survival rate to the mai+e plant in nature ;4edoroff 200,<! The cob itself usually remains on the plant until harvested but if left
on the plant or if damaged by insects or disease will eventually fall to the ground! This means that there may be locali+ed dispersal of grains around the base of the plant! Carvesting activities and grain transport result in more widespread grain dispersal ;see Section 8!1 for a discussion of volunteerism<! /ndications from the scientific literature would not suggest that dispersal of mai+e grain by animals2 including birds is significant although there is the possibility that intact grain may be spread as the result of the activities of vertebrate pests ;see Section 9!2!2<! 1.1 See, ,ormancy an, germina+ion
1ormancy is not associated with modern mai+e cultivars although it does occur in other Zea spp! ;Simpson 1::0<! Seeds from one mai+e crop can survive over winter and germinate in warmer weather! Cowever2 there is no or little inhibition of germination and mature seed can begin to germinate immediately under favourable conditions2 even if still attached to the cob ;1uncan 1:95<! The plant hormone abscisic acid ; & < has been shown to be important in the regulation of the onset and maintenance of dormancy in seed ;see discussion in &ewley 1::9<! Some mai+e plants show germination before seed maturity ;%angelsdorf 1:,0<! There is evidence that a balance between & and another natural hormone2 gibberellin2 determines whether precocious germination before maturation ;known as vivipary< occurs ;-hite D #ivin 2000= -hite et al! 2000<! Several mutants known as viviparous -vp.2 have been identified in mai+e and2 as a result of deficiencies in the & biosynthetic pathway2 do not show dormancy! %ai+e germination re8uires good soil moisture and a minimum soil temperature of about 12V*2 and does not occur in soil temperatures below 10V* ; 0&/)S 2005a= 4arrell D )KNeeffe 2009<! The rate of germination is dependent to some e(tent on temperature and2 for e(ample2 tropical mai+e may start germination within 2 F , days under high temperatures ;such as during tropical summer conditions< while it could take 7 F 8 days under lower temperatures ;such as during winter or at high altitudes< ;"aliwal 2000f<! There is some evidence that the pigment phytochrome may be involved in the control of germination in mai+e2 particularly under conditions of osmotic stress ;Thanos D %itrakos 200.<! %ai+e seed in usually planted to a depth of 2 F 10 cm if ade8uate moisture is available but can be planted down to 20 cm in arid regions ;"urseglove 1:92<! The ma(imum depth from which a seedling can emerge is limited by the ma(imum potential for elongation of the speciali+ed meristematic area ;known as the mesocotyl< Eust below the coleoptile node ;1uncan 1:95< ;see 4igure ,<! %ai+e germination follows a similar pattern to that of many grasses e(cept for differences in scale that occur as a result of the large endosperm and embryo in the mai+e seed! The cytohistology of germination has been described by Sass ;1:99<! &asically2 it is an ordered process that follows imbibition of water through the pericarp! %itosis and cell elongation start in the radicle ;root< appro(imately 2. hours after imbibition and then begin in other areas along the embryonic a(is! Seedling emergence is described in Section .!5 below!
,1
1.7
*ege+a+i-e gro8+h
Canway ;1:7,< proposed eleven stages of growth of mai+e2 with stage 0 being germination and emergence and stages 7 F 10 occurring after silking! /n terms of crop management it is more usual to condense the number of stages to si(2 with stage 7 incorporating dry@down and grain harvesting ;*olless 1::2<! The vegetative stages of growth2 together with their characteristics2 can broadly be described as follows ;Canway 1:7,= "urseglove 1:92= 1uncan 1:95= *olless 1::2= "aliwal 2000f<: Stage 1: 0ermination and $mergence ;appro(! 0 F 1. days after planting< F timing is dependent on factors such as soil temperature and moisture2 depth of sowing2 and surface hardness! The coleorhi+a and radicle ;root< emerge and 1@2 days later the coleoptile and plumule ;first foliage leaves< break through the seed coat ;refer to 4igure ,<! "lants develop primary ;seminal< roots2 a temporary root system2 until about the three@leaf stage of the seedling! Si( to 10 days after planting the coleoptile emerges from the soil2 splitting at the tip to allow the growth of the first foliage leaves! The shoot meristem itself remains Eust below the soil surface! Stage 2: $arly Vegetative ;appro(! 1. F .2 days after planting< F during this stage2 secondary ;adventitious< roots develop from the first node below the soil surface and from each successive node up to between seven and ten nodes2 all below the soil surface! These develop into a thick2 permanent fibrous root system reaching down typically to 1 F 2 m! Some adventitious roots may also emerge from nodes above ground2 and are termed brace roots! %ai+e races show marked differences in the mass2 branching and spread of lateral roots! The number of leaves that will develop on the plant ;up to about ,0< is determined ;/rish D Aegla 1::9< and the tassel begins to differentiate when about 5 leaves have emerged! The shoot meristem and the tassel primordium emerge above the soil surface by the si(@leaf stage and when eight leaves have fully emerged the shoot meristem is appro(imately 15 cm above the soil! 'ower leaves may start to senesce by the end of Stage 2! Stage ,: 'ate Vegetative ;appro(! .2 F 70 days after planting< F this is the stage of rapid growth ;linear dry matter accumulation< of both roots and leaves! 1uring development there is a basic repeating unit structure comprising leaf blade2 leaf sheath2 node and internode that makes up the entire vegetative shoot! /nternode elongation produces a new leaf every three to four days! $ventually2 the elongation of the lower internodes contributes to the formation of a stalk@like structure that rises up through the leaf sheaths! &y the end of Stage , the 17th leaf will have started to emerge2 the tassel will have reached full si+e although will not have fully emerged and the ears within the husks will be a few centimetres long! The first 5 F 7 lower leaves may senesce and cease to be functional! erial or brace roots usually emerge from the lower2 above@ground nodes! The e(tent of aerial root production is cultivar dependent as well as being influenced by rate of planting and nutrition! Tillering ;development of stalks from a(illary buds< is cultivar dependent2 with some cultivars forming few if any tillers under any conditions2 and some forming numerous tillers under all conditions! There is a correlation between the final number of leaves produced on a plant and the time between sowing and silking! The duration of vegetative development is linked to the thermal interval between the appearance of successive leaf tips ;called the phyllocron< and differs according to the temperature found in latitudinal +ones2 being
higher in tropical than temperate areas ;ToEo Soler et al! 2005< ;see also corresponding discussion in Section .!,!2<! /n terms of crop performance2 growth refers to biomass accumulation and is measured by parameters such as leaf area2 shoot3root weights and plant height! Techni8ues used to 8uantify growth are known as growth analysis and comprise a number of inde(es such as *rop 0rowth #ate2 #elative 0rowth #ate2 6et ssimilation #ate2 'eaf rea #atio2 and 'eaf rea /nde( ;' /< ;for a detailed discussion see 4ageria et al! 2007<! /mprovements in mai+e grain yields over time have been associated with delayed leaf senescence in newer hybrids leading to higher ' /s Eust after silking when the grain is filling ;Valentinu+ D Tollenaar 200.= 'ee D Tollenaar 2009<! The maEor contribution to ear development and final grain yield are made by the earleaf and all leaves below the earleaf ;Subedi D %a 2005<! %ai+e has a *. photosynthetic pathway2 which allows a continued response to increasing radiation up to full sunlight coupled with low levels of photorespiration! The ma(imum level of leaf photosynthesis per unit area occurs between full leaf e(pansion and silking ;'ee D Tollenaar 2009<! The growth of grain is dependent on total photosynthetic production only after flowering and is discussed in Section .!,!2 ;see also discussion in 6orman et al! 1::5= 'afitte 2000b= 'ee D Tollenaar 2009<! Thus2 vegetative dry matter continues to accumulate in mai+e after flowering ;4ageria et al! 2007= 'ee D Tollenaar 2009<! /f flowering is inhibited2 vegetative growth is significantly increased= tropical mai+e types that are grown in the long summer days of temperate regions such as the %idwest of the >S have delayed flowering and conse8uently produce more vegetative growth2 manifested as taller plants with high sugar levels in the stems ;>niversity of /llinois 2009<! /n a mature temperate mai+e plant appro(imately 50H of the total dry matter is allocated to the grain and 50H to the stover ;crop residue following harvest< F as shown by the C/ of appro(! 0!5 ;see Section .!,!2<! *ompared with other crops2 mai+e has a disproportionately high amount of stover ;over 50H of which is stalk< and it has been proposed that this could be utilised in the biofuels industry ;1hugga 2009<!
SECT!"N 7
7.#
B!"C2E
!STR%
N.+rien+ componen+s of +he maize 9ernel
The typical mature kernel as a whole is composed of 90 F 95H starch2 8 F 10H protein and . @ 5H oil ;&oyer D Cannah 1::.<! Cowever2 there are large differences in the relative concentrations of these components between different parts of the kernel ;see Table 7<! The two maEor structures of the kernel are the endosperm and the germ ;embryo<2 constituting about 80H and 10H of the mature kernel dry weight2 respectively! The endosperm is largely starch ;appro(imately :0H< while the germ contains high levels of fat ;appro(imately ,,H< and protein ;appro(imately 18H<! These differences become a significant consideration when mai+e is processed for consumption!
,,
Table .%
/elati,e content 012 of nutrient components in parts of the maize 3ernel*

,ericarp &see) coa-' En)osperm 9.0 2.6 0.9 96.7 0.72 .erm &embryo' 19 . 9.9 33.2 9.3 10.9
,ro-ein Cru)e 9ibre Cru)e 9aS-arch Su%ar
3.6 97.6 1.0 6.3 0.3.
2 Data ta<e# 'r = /A> (1882)
Table 9 provides a more detailed breakdown of the nutrients that can be found in the whole kernel of sweet corn! "ellagra is a nutritional disease caused by a deficiency in niacin2 and may be associated with mai+e@based diets in the mericas and frica! -hile niacin is present in mai+e2 it e(ists in a bound form that is not biologically available to monogastric ;single@stomach< animals ;)koruwa D Nling 1::7<! Table 9! 6utrient content of sweet corn2 fresh2 boiledWNu-rien,ro5ima-es K#er*" M isture Aitr *e# Lr tei# /at Ash /ruct se Gluc se $ucr se $u*ars5 t tal $tarch A%aila!le Har! h"drate ? tal Dietar" /i!re #inera(s Halciu= H pper Ir # Ma*#esiu= Ma#*a#ese 3 0.070 0.7 .1 0.3.0 = * = * = * = * = * .39 6..0 0.76 ..2 2.9 1.0 0.3 0.. ..7 1.3 6.1 12.9 ..9 <B * * * * * * * * * * * * / late Dietar" / late KOui%ale#ts &ita=i# H Alpha Har te#e Beta Har te#e Beta Har te#e eOui%ale#ts Reti# l KOui%ale#ts Lipi)s H1730 0.21 * 31 31 . 12 7 12 2 Ri! 'la%i# Aiaci# Aiaci# deri%ed 'r = ?r"pt pha# r Lr tei# Aiaci# KOui%ale#ts 0.07 2.0 0.7 2.7 ?hia=i# ?i-amins 0.11 = * = * = * = * = * M* M* = * M* M* M* M* ?a(ue per 1!! % Lh sph rus L tassiu= $ diu= Ni#c 110 219 101 0.6 = * = * = * = *
H1930 H2030 ? tal $aturated /att" Acids H1931 H2031 ? tal M # u#saturated /att" Acids H1932 (u#di''ere#tiated) H1933 (u#di''ere#tiated) ? tal L l"u#saturated /att" Acids ? tal L #* Hhai# >=e*a 3 L l"u#saturates Hh lester l Amino Aci)s Ala#i#e Ar*i#i#e Aspartic Acid H"sti#e P H"stei#e Gluta=ic Acid Gl"ci#e Histidi#e Is leuci#e Leuci#e L"si#e Methi #i#e Lhe#"lala#i#e Lr li#e $eri#e ?hre #i#e ?r"pt pha#
0.0. 0.01 0.3 0.7. 0.01 0.7 1.26 0.02 1.3 0
* * * &ali#e * * * * * * Hitric Acid Malic Acid Or%anic Aci)s 1.0 0.2 22. ?"r si#e 111
* = * = *
* *
331 118 216 81 613 1.1 12. 111 .76 131 101 182 321 189 188 32
= * = * = * = * = * = * = * = * = * = * = * = * = * = * = * =
,5
2$ urce3 /$AAN (2007)
7.#.#
S+arch
Starch synthesis has been reviewed in a number of articles ;&oyer D Cannah 1::.= 6elson D "an 1::5= &oyer 1::7= Aames et al! 200,<! Starch is synthesised as part of a mega@molecular assembly2 the starch granule2 within the amyloplasts of mai+e endosperm cells and2 in most cases2 contains two maEor types of glucose homopolymers2 amylose and amylopectin! /n amylose2 the glucose residues are mai#l" li#<ed %ia Q-15. li#<a*es5 ;hich results i# a li#ear chai# ' *luc se u#its. I# a="l pecti#5 the maEority of linkages are Q@12. linkages2 with Q@127 linkages providing branching! Cighly branched regions alternate with regions devoid of branching! The latter regions make it possible for other linear glucose chains to align in the form of parallel helices in the gaps2 giving starch its semicrystalline structure! The primary source of the carbon skeletons for starch synthesis is sucrose translocated to the endosperm! The starch synthesis pathway in cereal endosperm re8uires en+yme isoforms that are not present in other cereal tissues or non@cereal plants! s endosperm cells mature2 the starch granules increase in si+e and in the proportion of amylose so that the mature cell typically contains the ratio ,: 1 amylopectin: amylose! -ithin an endosperm cell type2 all mature starch granules are of a similar si+e! mature granule may contain as many as 10: amylose and 109 amylopectin molecules! These molecules are arranged regularly2 giving the granule stability and crystalline properties! The mai+e mutants #axy -#x. and amylose extender -ae. are grown e(tensively for their uni8ue starches ;see discussion in 4ergason 2000<! /n the #axy mutant2 the starch is composed almost entirely of amylopectin and the mature kernel has an opa8ue phenotype! %utations at the ae locus result in an increase in the amylose content of the endosperm relative to its amylopectin content= such starch can be used to produce tough2 edible or biodegradable films and gels ;#obertson D Stinard 1:88<! 7.#.& Pro+ein
1iscussion of mai+e kernel proteins can be found in a number of articles ;&oyer D Cannah 1::.= -oo et al! 2001= %onEardino et al! 2005= %onEardino et al! 2007<! Storage protein ;a 9S globulin< is found in the embryo and in the endosperm! The relative amount of protein is highest in the embryo ;see Table 5< but2 because the endosperm occupies a greater part of the kernel2 it contributes the greatest total amount of protein ;4 ) 1::2<! The endosperm proteins can be divided into prolamins2 collectively referred to as +eins2 and comprising about 52H of kernel nitrogen= glutelins ;ca 25H of kernel nitrogen<= albumins ;ca 9H<= and globulins ;ca 5H<! Ieins2 which accumulate in the rough endoplasmic reticulum of endosperm cells2 contain large amounts of the amino acids glutamine2 proline2 leucine and alanine but are very low in the essential amino acids lysine and tryptophan! $mbryo proteins provide higher levels of tryptophan and lysine ;4 ) 1::2<! number of different
molecular weight +eins have been identified by S1S@" 0$ and have been further classified2 based on their solubility and structural relationships2 into Q- ;22 and 1: k1<2 R- (11 k1<2 S@ ;292 17 and 50 k1< and T@ ;10 and 18 k1< +eins! The Q@+eins are the most abundant at ca 80H of total +eins and are encoded by a large family of genes2 while the other classes of +eins are encoded by only one or two genes each! The opa/ue) locus ;o)< in mai+e regulates the e(pression of many members of the +ein multigene family and mai+e plants carrying an o) mutation have seeds with an opa8ue appearance that is attributable to a dramatic reduction in the +ein storage proteins and concomitant increase in lysine ;%ert+ et al! 1:7.< and tryptophan ;%isra et al! 1:92<! part from the desirable high lysine and tryptophan content2 these mutants have agronomically inferior characteristics including reduced yield and soft floury grain endosperm that increases disease and insect susceptibility and makes food processing difficult ;see review in>fa+ D 0alili 2008<! &reeding at the /nternational %ai+e and -heat /mprovement *enter in %e(ico in the early 1::0s saw the development of opa/ue) @ derived L"% ;Luality "rotein %ai+e< lines with better performance and nutritional values! These have now been adapted for use in many countries ;>fa+ D 0alili 2008<! L"% varieties with high levels of the amino acid tryptophan may also compensate to some e(tent for the lack of available niacin in mai+e ;see Section 5!1<! 7.#./ Lipi,s
'ipids ;oil< are found mainly in the embryo2 specifically in the scutellum! They comprise .0H of the dry weight of the scutellum and are used for gluconeogenesis to support the developing embryo following germination ;)aks D &eevers 1:7.<! The embryo contains appro(imately ,,H oil ;see Table 5< while a typical whole kernel contains appro(imately .H oil ;see Table 9<! Cowever2 the amount and composition of oil in the kernel is under genetic control and2 for e(ample2 selection in one line ;/llinois Cigh )il< after 107 generations2 over more than 100 years2 has continuously increased the percentage of kernel oil to over 20H ;1udley 2009<! 4atty acids in corn oil nearly always occur esterified to the hydro(yl groups of glycerol2 thus forming triacylglycerides! These triacylglycerides contain a mi(ture of saturated and unsaturated fatty acids distributed appro(imately as follows ;&oyer D Cannah 1::.<:

Saturated: 12H palmitic acid ;17:0<2 2H stearic acid ;18:0<2 >nsaturated: 25H oleic acid ;18:1<2 70H linoleic acid ;18:2<2 and 1H linolenic acid ;18:,<!
%ai+e oil is generally accepted as high 8uality2 as determined by the high linoleic acid and low linolenic acid content! There is a suggestion that the breakdown of esterified linolenic acid to the free form in stored mai+e meal may be associated with predisposition to carcinogenesis of the oesophagus ;Sammon D /puto 2007<! The genetics of lipid content in mai+e oil is comple(2 probably controlled by multigenic inheritance of a large number of genes ;&oyer D Cannah 1::.<! 7.& To3ins
To(ic reactions after feeding livestock with mai+e ;especially green chop< or allowing animals to gra+e on mai+e may result from nitrate or nitrite poisoning! 6itrite and ,9
nitrogen dio(ide are produced by bacteria associated with mai+e plants causing to(icity in humans and animals! lso2 mycoto(ins produced by fungi2 which may infect mai+e plants and3or grain2 may be present in mai+e! Since those to(ins are produced by plant@associated microorganisms rather than the plant itself2 they are discussed in Section 9!2!.! 7.&.# Ni+ra+e poisoning
6itrate is taken up and used in the biosynthesis of proteins and other nitrogen containing components by plants ;eg reviewed in "avelchak 1:::<! >nder certain environmental conditions2 eg after a drought breaks2 some plant species can accumulate relatively high levels of nitrate as it may be present at relatively high levels in the soil! %ai+e2 particularly the lowermost parts of the stalk ;R 20 cm<2 may accumulate nitrate to levels that can be to(ic to stock ;$verist 1:81<! #uminants are less susceptible to nitrate poisoning2 if their diet includes sufficient amounts of carbohydrates while they are fed nitrate@rich feed! %icroorganisms in the rumen convert nitrate to nitrite2 which is then converted to ammonia by other rumen@ inhabiting microorganisms! Symptoms of nitrate poisoning in stock are mainly abdominal pain2 diarrhoea and salivation ;Calpin D Cides 2002<! 7./ Allergens
lthough mai+e is not considered highly allergenic2 some allergic reactions have been reported in the scientific literature: mai+e can cause both food and inhalation allergies! 4or e(ample2 pollen and mai+e dust may cause allergies2 such as hay@fever and bakerBs asthma! The /nternational >nion of /mmunological Societies currently lists four mai+e allergens on its website ;http:33www!allergen!org3 llergen!asp(<: Iea m 12 Iea m 122 Iea m 1. and Iea m 25! /nformation on mai+e allergens is given below! "ollen allergens include the Iea m1 pr tei#5 als #a=ed KULB5 a Re+pa#si# ;ith a = lecular ;ei*ht ' appr +i=atel" 26 <D! Iea m 1 is a homolog of the 'ol p 1 allergen in ryegrass ;0olium perenne< ;&roadwater et al! 1::,<! The e(pansin is present in mai+e pollen in four isoforms2 the most abundant one being $U"&1 ;reviewed in Mennawar et al! 2007<! The allergen Iea m 12 is a profilin with the molecular weight of appro(imately 1. k1a! /n mai+e2 profilins are encoded by a multi@gene family ;Staiger et al! 1::,<! "rofilins have also been described in a number of plant species2 including alder ;Alnus glutinosa<2 ha+el ;Corylus avellana<2 timothy grass ;$hleum pratense< and rye ;"ecale cereale<2 and a profilin from birch ;%etula verrucosa< was characterised in a study by Valenta et al! ;1::1< as a maEor pollen allergen! /n one study2 "astorello et al ;2000< identified a maEor food allergen of mai+e as a : k1 lipid transfer protein ;'T"2 Swiss "rot 1ata &ank2 accession number "1:757< that shows homology to 'T"s from rice2 peach and apricot! /t is also known as the Iea m 1. cereal allergen! "re@incubation of immune sera from patients allergic to mai+e with either rice or peach e(tracts abolished antibody binding in immunoblots meaning that cross@reaction of the Iea m 1. allergen with rice or peach allergens occurs! The second most important food allergen in mai+e found in that study was a 17 k1
inhibitor of trypsin3activated Cageman factor ;U// = Swiss "rot 1ata &ank2 accession number "01088<! /t has been shown that members of this tr"psi# a#d Q-a="lase i#hi!it r pr tei# 'a=il" present in other cereals can cause &akerBs asthma! "re@ incubation of immune sera of allergic patients with Timothy grass pollen2 wheat2 rice or barley e(tract completely inhibited binding of antibodies to the 17 k1 allergen as well as other2 minor mai+e seed allergens! I# additi # t the pre%i usl" descri!ed aller*e#ic R-e+pa#si#s5 pr 'ili#s a#d L?Ls5 (eichel et al! ;2007< also found thioredo(ins2 such as Iea m 25 in enriched repertoires of /g$@binding se8uences of mai+e! "asini et al ;2002< also identified a 50 k1a protein as a maEor allergen2 which is both heat stable and resistant to peptic3pancreatic digestion! 'ee et al ;2005< described a 50 <Da S--ei# 'r = =ai-e ;ith str #* in-vitro cross@reactivity with the almond maEor protein ; %"<2 a maEor allergen in almonds ;$runus amygdalus<! /t is currently not clear if the two 50 k1a proteins correspond to the same allergen in mai+e! 7.1 "+her .n,esira5le phy+ochemicals
%ai+e contains low levels of some anti@nutrients2 chemicals that block the normal uptake or utilisation of nutrients: "hosphorous is mainly stored in plants in the form of phytic acid2 or phytate2 which is not easily accessible to humans and other animals! This has a number of conse8uences: humans may need to obtain phosphate from different sources2 animal feed needs to be fortified with phosphate and2 as a conse8uence of the latter2 phosphate is e(creted at high levels leading to environmental problems! "hytate also acts as an anti@nutrient as it chelates metal ions2 including iron2 and prevents those from being accessible to humans and other animals at sufficiently high levels ;reviewed in Currell 200,<! #ecently2 progress has been made in genetically engineering mai+e e(pressing phytase ;eg 1rakakaki et al! 2005= *hen et al! 2009<! nother anti@nutrient that is present in mai+e is the sugar raffinose! #affinose is non@ digestible2 and is considered an anti@nutrient due to gas production and resulting flatulence! /t can be removed from food and feed by soaking2 cooking2 and irradiation or by en+yme or solvent treatment ;)$*1 2002<! &oth trypsin and chymotrypsin inhibitors are present at low levels in mai+e ;)$*1 2002<! They are not considered important for human or animal nutrition! 7.7 Beneficial phy+ochemicals
%ai+e is considered an important food crop for humans and a high@energy feed for animals ;4 ) 1::2<! /n the human diet it is a good source of vitamin &12 vitamin &52 folate2 dietary fibre2 vitamin *2 phosphorus and manganese ;see Table 7< and as a staple it compares favourably with root and tuber crops and is similar in energy to dried legumes ;)koruwa D Nling 1::7<! The average nutritional content of various forms of mai+e is given in Table 8! "rocessing reduces the concentration of proteins2 lipids and fibre ;4 ) 1::2<!
,:
Table 4%
/anges for pro5imate anal-sis of maize grain6 sweet corn 3ernels and silage*
#ai$e %rain S1ee- corn 6..6 - 9. 11.3 - 11.7 ..97 - 9.61 2.19 - 3.97 62.1 - 68.19 11.2 - 11.7 (t tal dietar" 'i!re) .0 - .9.2 (#eutral deter*e#t 'i!re) 2.8 : 1.6 V 'resh ;ei*ht V dr" ;ei*ht V dr" ;ei*ht V dr" ;ei*ht V dr" ;ei*ht V dr" ;ei*ht 6 - 23 7 - 12 3.1 - 1.9 1.1 - 3.8 92.2 - 92.8 9.3 - 11.8 (t tal dietar" 'i!re) #ai$e si(a%e 72 - 69 ..6 : 8.2
#ois-ure ,ro-ein Lipi) &-o-a(' Ash Carbohy)ra-e Fibre
2$ urce3 >KHD (2002)
SECT!"N :
AB!"T!C !NTERACT!"NS
%ai+e shows a wide genetic base for abiotic stress tolerance2 which is mirrored by its ability to grow in a variety of environments2 although it is essentially a crop of warm climates with ade8uate moisture ;"urseglove 1:92< and is therefore not suited to semi@arid or wet tropical climates! /t is argued that all of the genetic improvement in mai+e yield over the past 90 years is the result of changes to physiological attributes that2 in turn2 have imparted enhanced abiotic stress tolerance ;'ee D Tollenaar 2009< ie selection of hybrids for high density planting has been accompanied by increased resistance to stresses such as drought and has permitted consistent performance across a range of variable environments ;1uvick D *assman 1:::= 1hugga 2009<! )ne way that abiotic stress affects the mai+e plant is by moving the source@sink balance ;'ee D Tollenaar 2009<! *lassic symptoms of e(cess source capacity are purpling of leaves2 sheath tissues and stalks during grainfilling while symptoms of e(cess sink capacity are premature senescence of leaves and stalks during grainfilling ;'ee D Tollenaar 2009<! :.# N.+rien+ re;.iremen+s
%ai+e grown commercially2 whether for grain or silage2 has a high demand for nutrients2 especially nitrogen ;6<2 phosphorus ;"< and potassium ;N< ;&irch et al! 200,<! s an e(ample2 Table : gives some figures for nutrient removal by a sweet corn crop!
Table 7% N6 8 9 : remo,al b- a 24 t;ha crop of sweet corn*
Ni-ro%en &*%+ha' /emo0a( by cobs /emo0a( by 0e%e-a-i0e par-s To-a( remo0a( 2$ urce3 Bec<i#*ha= (2006) 110 200 ,hosphor us &*%+ha' 17 2. ,o-assium &*%+ha' 70 110
310
.0
210
The micronutrients +inc and molybdenum may2 depending on soil type2 also be
important to apply to mai+e crops to prevent deficiency symptoms! Seedlings do not tolerate high levels of fertili+er and therefore starter fertili+er should be drilled at least 5 cm to the side of the seed during sowing ;Cughes 2007c<! /n the mai+e@growing areas of the tropics2 acid soils are widespread and plant growth is constrained by the associated aluminium to(icities that lead to stunting and impairment of root growth and subse8uent inability of plants to take up moisture from the soil ;'afitte 2000a<! 1itrogen /f water and temperature conditions are ideal productivity of mai+e is mainly limited by availability of nitrogen ;"urseglove 1:92= 'afitte 2000a= &irch et al! 200,<! /t has been suggested that in irrigated mai+e2 nitrogen deficiency caused by denitrification in irrigated soils that are slowly permeable to water may be the principal cause for low yield! Cowever2 in a study done in the %urrumbidgee /rrigation area of southern 6S- it was concluded that poor transport of nitrogen associated with ano(ic conditions in the root +one of water@logged plants may also play a significant role ;%osier et al! 1:87<! 6itrogen is a component of a number of compounds ;eg proteins2 nucleic acids2 chlorophylls< and has an important role in many plant physiological processes ;#aven et al! 1:::<! /n particular2 it is important in the efficient capture and use of solar radiation and therefore affects yield ;'afitte 2000a= &irch et al! 200,<! %ai+e begins to rapidly take up nitrogen ;and other nutrients< during the middle vegetative growth period with the ma(imum rate of nitrogen uptake occurring near silk ;Canway 1:7,<! 6itrogen deficiency is indicated by leaf yellowing ;first in the lowest leaves< that starts at the tip and then e(tends along the mid@rib2 stunted plants2 delayed flowering and short2 poorly filled ears ;*olless 1::2= Cughes 2007c<! %ai+e can utilise nitrogen in both the ammonium and nitrate forms but2 because of the ready conversion of ammonium to nitrate by soil microbes2 most nitrogen is taken up as nitrate ;*olless 1::2= 4arnham et al! 200,<! /f nitrogen is supplied via irrigation water2 urea is the best source ;&irch et al! 200,<! $hosphorus "hosphorus is a component of energy@carrying phosphate compounds such as T" and 1" ;#aven et al! 1:::<! "hosphorus deficiency reduces the leaf area inde( ;' /< of mai+e2 thus reducing the amount of photosynthetically active radiation absorbed by the canopy and leading ultimately to lower biomass accumulation ;"ellerin et al! 2000<! The negative effect of phosphorus deficiency on ' / also adversely affects adventitious root emergence and therefore may further e(acerbate phosphorus uptake ;"ellerin et al! 2000<! Symptoms of phosphorus deficiency are slow growth2 late maturity2 a reddening of leaves2 poorly developed root systems and small ear si+e ;*olless 1::2= 'afitte 2000a= Cughes 2007c<! /n the %urrumbidgee and *oleambally irrigation areas of southern 6ew South -ales where mai+e was grown in rotation with flooded rice crops2 it was noted that growth of the mai+e crop was significantly reduced because of phosphorus unavailability .1
associated with increased iron o(ides that immobili+ed the phosphorus ;-illett et al! 1:98<! 4or this reason mai+e is now rarely grown in rotation with flooded rice! Some mai+e growing soils in ustralia ;eg the vertosols of the 1arling 1owns2 'ockyer Valley and 'iverpool "lains< have inherently high phosphorus status ;&irch et al! 200,<! $otassium The potassium re8uirement of mai+e2 particularly if it is used for silage2 is high ;&irch et al! 200,<! "otassium has a crucial role in a number of plant physiological responses2 such as stomatal opening and closing2 driven by osmosis and ionic balance! /t is also an activator of many en+ymes ;#aven et al! 1:::< and influences photosynthesis2 nutrient and assimilate transport and en+yme activation for protein synthesis ;see discussion and references in Aordan@%eille D "ellerin 200.<! "otassium deficiency causes a reduction in ' / probably through altered plant@water relationships that reduce leaf elongation rate ;Aordan@%eille D "ellerin 200.<! 1eficiency symptoms include yellowing and death of leaf margins particularly in the lower leaves2 the tendency of the crop to lodge and development of small ears that fail to fill at the tip ;*olless 1::2= Cughes 2007c<! Zinc %ai+e grown on alkaline soils such as the vertosols can show severe +inc deficiency symptoms ;*olless 1::2= &irch et al! 200,<! These symptoms include light streaking of leaves in from the leaf margins ;leaf edges2 midrib and tip remain green<2 and stunted growth of the crop ;*olless 1::2= Cughes 2007c<! !oly*denum cid soils2 particularly in high rainfall coastal regions of ustralia2 can be deficient in molybdenum! 1eficiency symptoms include yellowing and eventual death of leaf tips2 and stunting and sometimes death of plants ;*olless 1::2= Cughes 2007c<! :.& Tempera+.re re;.iremen+s an, +olerances
%ai+e is a summer@growing crop re8uiring warm daytime temperatures of between 25o * @ ,0o * and cool nights ;*olless 1::2<! Temperatures below 8o * ;or 0o * after silking< or above appro(imately .0o * usually cause cessation of development ;&irch et al! 200,<! 1ifferent mai+e cultivars have different optimal temperature re8uirements and2 for e(ample2 tropical cultivars derived from JhighlandB mai+e are better able to grow and develop at lower temperatures than those adapted to JlowlandB or Jmid@altitudeB areas! Temperatures that are outside the range of adaptation of a cultivar may impact negatively on factors such as photosynthesis2 translocation2 and pollen viability ;see discussion in 'afitte 2000a2 summarised in Table 10<! /n particular2 high temperatures have a negative impact on kernel growth2 kernel mass and protein accumulation ;Aones et al! 1:8.= Aones et al! 1:85= %onEardino et al! 2005= %onEardino et al! 2007<!
Table 1)% (ffects of temperature on 3e- processes in maize*
,rocess E99ec-
@i%h Tempera-ure
Lh t s"#thesis R t h r= #e pr ducti # (A!scisic acid a#d c"t <i#i#) L lli#ati #
Reduced a! %e .0 H due t =e=!ra#e da=a*eD irre%ersi!le da=a*e a! %e .1 H Reduced h r= #e pr ducti # restricts chl r plast de%el p=e#t a#d ph t s"#thetic acti%it" L lle# %ia!ilit" reduced a! %e 31 H especiall" i' there is l ; hu=idit". Ma" reduce *rai# set. Grai#'illi#* durati # is reducedD *rai#'illi#* rate is i#creased. >%erall "ield is usuall" reduced. K+acer!ated !" ;ater stress. Reduced due t reduced e#-"=e 'u#cti # a#d =e=!ra#e da=a*e Reduced due t reduced e#-"=e 'u#cti # a#d =e=!ra#e da=a*e Reduced due t reduced e#-"=e 'u#cti # a#d =e=!ra#e da=a*e i# r ts Reduced5 especiall" i# tr pical l ;la#d culti%ars at less tha# 11 H Reduced5 especiall" i# tr pical l ;la#d culti%ars at less tha# 10 HD
Grai# "ield
Lo1 Tempera-ure
Lh t s"#thesis
Lea' e+te#si #
(ater & #utrie#t upta<e Grai# "ield
?ra#sl cati # 2$ urce3 (La'itte 2000a)
%ai+e varieties grown in ustralia are mostly adapted to tropical2 sub@tropical and warm temperate regions and are therefore not grown in areas where frost may limit early development! "rior to emergence plants may survive frosts for as long as their growing point is below the soil surface2 ie at least until four@leaf stage ;4arnham et al! 200,<! Mields in areas with high temperatures ;such as Natherine and Nununurra< are generally lower than in areas with more moderate climates ;such as 0atton and the #iverina< ;discussed in &irch et al! 200,<! The high temperatures affect yield by both shortening the time to maturity ;thus limiting grain filling< and e(acerbating water stress ;&irch et al! 200,< ;see also discussion in Section .!,!2<! :./ 0a+er
Water de&iciency -orldwide2 the average yield losses in mai+e crops due to drought can be high2 particularly in the tropics ;Srinivasan et al! 200.<! #ainfall is a limiting factor to dryland production of commercial mai+e crops and irrigation is essential in areas with a winter dominant rainfall pattern or where the amount of summer dominant rain is highly variable ;&irch et al! 200,<! %ai+e is particularly susceptible to water stress at the flowering stage when yield potential is being set ;*olless 1::2= &irch et al! 200,= Srinivasan et al! 200.< especially as this coincides with the high evapotranspiration rates of mid@summer ;4arnham et al! 200,<! Stress at this time can reduce grain yield by 7 F 8H for each day of stress ;*olless 1::2<! 0rant et al! ;1:8:< did a study to determine the timing of sensitivity of mai+e yield components to water stress and concluded that kernel number was sensitive to water stress between 2 F 22 d after silking and kernel weight was most sensitive 12 F 17 d after silking! The reduction in .,
kernel number has been proposed to be due to a number of factors including lack of pollination2 abortion of ovules prior to fertili+ation and abortion of fertili+ed kernels ;see discussion in 0rant et al! ;1:8:<= &assetti et al! ;1::,<= Srinivasan et al! ;200.<<!The latter is the most probable and is likely to be due to decreased assimilate flow to the developing grain ;0rant et al! 1:8:= Iinselmeier et al! 1::5<! Waterlogging /n the maEor mai+e growing areas of sia and merica2 significant annual losses can occur because of waterlogging ;Sus+kiw 1::.= Srinivasan et al! 200.<! "lants growing for prolonged periods in waterlogged soils show stomatal closure2 reduced leaf area growth2 chlorosis2 reduced root growth2 root death and ultimately plant mortality ;'afitte 2000a= Srinivasan et al! 200.<! 1amage to roots is due mainly to the accumulation of to(ic products ;such as lactic acid< as a result of anaerobic respiration! Tropical and sub@tropical cultivars are most susceptible to waterlogging at the early vegetative stage and at Jknee@highB stage2 ie prior to tasseling ;Srinivasan et al! 200.<! %olecular and cellular changes to mai+e seedlings under short@term waterlogging has been widely studied as mai+e typically shows an Janaerobic responseB in which some 20 proteins2 mostly associated with glycolysis or sugar@phosphate metabolism2 are synthesi+ed ;Sachs et al! 1::7= Subbaiah D Sachs 200,<! #oot porosity2 that facilitates o(ygen diffusion from the above ground parts to the submerged roots2 is increased by the selective death of root cortical cells and helps to prolong the survival of plants ;Subbaiah D Sachs 200,<! #oot tip death2 also characteristic of response to ano(ia2 may enhance survival and subse8uent recovery2 when waterlogging is removed2 by eliminating an area of metabolically active tissue ;Subbaiah D Sachs 200,<! 1evelopment of adventitious roots on the soil surface is also important in conferring flooding tolerance ;%ano et al! 2005<! :.1 "alinity Salinity is regarded more as a problem of irrigated rather than dryland crops! %ai+e is classed as a salt@sensitive or moderately salt@tolerant plant ;Naddah D 0howail 1:7.= 'afitte 2000a< although there is wide variation between cultivars in response to salinity ;#ao D %c6eilly 1:::<! /t is a Jsalt e(cluderB rather than a Jsalt accumulatorB and therefore shows rapid productivity decline with increasing salinity ;Mensen D &iel 2007<= salt e(cluders2 because they cause salt to accumulate around the roots2 find themselves in an increasingly to(ic environment! 4or mai+e growing in a moderate to slow draining soil in an inland 6S- climate it is estimated that a 10H reduction in yield would occur if the crop were irrigated with water of electrical conductivity 1!9 dS3m compared to water of 1!1 dS3m ;$vans 2007<! Naddah D 0howail ;1:7.<2 in a study of mai+e grown in $gypt2 noted the following with regard to the effect of salinity ;up to :2000 ppm of a mi(ture of 6a*l and *a*l2< at various stages of growth:

"+her a5io+ic s+resses
Salinity delayed germination but had no effect on final emergence! Salinity applied at seeding time resulted in reduced vegetative growth2 delayed
tasseling and silking2 smaller cobs and fewer developed kernels ;i!e! reduced yield<! Salinity applied 21 days after seeding had less deleterious effects "lants in which salinity was applied at the commencement of visible tasselling did not show any specific sensitivity to the salt concentrations that were tested!
SECT!"N <
<.#
B!"T!C !NTERACT!"NS
0ee,s
lthough mai+e is a vigorous and tall@growing plant2 it is susceptible to competition from summer weeds2 particularly at the early stages of crop growth and particularly because the wide spacing between rows provides opportunity for weed establishment ;"ritchard 1::5= Aames et al! 2000= 4arrell D )KNeeffe 2009= %orris 2008<! -eeds may directly lead to yield reduction by competing with the mai+e plants for nutrients and water and need to be controlled within , weeks of crop emergence ;Aames et al! 2000<! %inimising weeds through to crop maturity is also important in terms of ease of harvest and reducing grain contamination! The latter is a significant consideration for mai+e grain imported into ustralia ;"heloung et al! 1:::a<! range of annual grasses ;eg barnyard grass F 2chinochloa colona= crowsfoot grass F 2leusine indica= $ennisetum spp!= Digitaria spp!= %rachiaria spp!< and broadleaf weeds ;eg caltrop F Tri*ulus terrestris= senna F "enna o*tusi&olia= nutgrass F Cyperus rotundus= pigweed F $otrtulaca spp!< occur in mai+e crops ;see eg Cughes 2007e= )K0ara 2009<! Some weeds such as Cardheads ;Acroptilon repens< and -itchweed ;"triga spp!< are of concern because they occur generally in cereal growing areas and are difficult to eradicate ;1"/ Vic 2009= Aohnson 2008<! -eed management can take a number of forms with an integrated approach being recommended ;4arrell D )KNeeffe 2009< and including crop rotations2 planting into weed@free seedbeds2 pre@plant cultivation2 inter@row cultivation2 use of pre@plant3pre@ emergence3post@emergence herbicides and use of conventionally bred imida+olinone tolerant mai+e hybrids ;Cughes 2007e= 4arrell D )KNeeffe 2009= )K0ara 2009<! range of herbicides offer fle(ible options for weed control and often two herbicides can be used in combination to give a broader spectrum of control ;)K0ara 2009<! *ommonly used herbicides include atra+ine2 flumetulam2 dicamba2 metolachlor2 fluro(ypyr2 pendimethalin2 propachlor2 tribenuron methyl and triclopyr ;Storrie et al! 2005= 4arrell D )KNeeffe 2009= )K0ara 2009<! The *learfieldX production system provides weed management through the use of two imida+olinone@tolerant mai+e varieties and imida+olinone herbicides such as ima+ethpyr Y ima+apyr ;)K0ara 2009= gricentre 2008<! similar system using resistant mai+e seed dressed with ima+ypyr Y pyrithiobac has been developed by */%%MT for use in subsistence farming areas where "triga spp! cause significant losses ;Nanampiu et al! 2001<! <.& <.&.# Pes+s an, ,iseases !nsec+s an, o+her in-er+e5ra+e pes+s
%ai+e is most susceptible to damage by insects during the establishment phase when soil insects can cause up to ,0H losses and necessitate replanting of the crop2 and
.5
from tasselling to harvest ;0oodyer 1:85= 4arrell D )KNeeffe 2009= )K0ara 2009<! /nvertebrate pests of establishing crops damage germinating seeds and seedlings and include ;Table 11< black field earwigs2 wireworms2 false wireworms2 cutworms2 mai+e stem borer2 and beetles such as the frican black beetle! The recent development of insecticidal seed treatments ;eg imidicloroprid and thiametho(am< has helped to control soil pests and some above@ground insects ;)K0ara 2009< while more traditional methods such as in@furrow spraying and agronomic practices at sowing are also effective ;Cughes 2007b<! /nvertebrate pests of developing and maturing crops include ;Table 11< common armyworm2 corn earworm2 mai+e leafhopper2 redshouldered leaf beetle2 corn aphid2 two@spotted spider mite2 green vegetable bug2 mai+e weevil and thrips! /nsect pests multiply and develop faster in the tropical north of ustralia than in southern environments and can represent a significant problem there ;)K0ara 2009<! Specific pesticides have been developed for control of specific pests ;see Certel D #oberts 2009<! /n irrigated mai+e2 JchemigationB ;application of pesticides through irrigation water< is an option ;)K0ara 2009<! The cereals are susceptible to insect attack during grain storage and mai+e weevil ;"itophilus zeamais< is a particularly serious pest of stored mai+e as well as infesting maturing cobs ;0oodyer 1:85= *ollins 1::8<!
Table 11%
C(assi9ica-ion Arachni) Acari#a3 ?etra#"chidae Tetranychus urticae ?; sp tted spider =ite Ma" cause e+cessi%e lea' da=a*e a#d he#ce reducti # i# "ield.
Common in,ertebrate pests of maize in &ustralia

Ta5onomic name Common name 4mpac-
4nsecH le ptera3 ?e#e!ri #idae Gonocephalum spp.; Pterohelaeus spp. /alse ;ire; r= /eeds # *er=i#ati#* seeds a#d sh ts. I# dr" c #diti #s ca# 'eed # dr" seed. /eeds # seed5 !ut = re usuall" attac< u#der*r u#d ste=s. Hhe;s i#t the ste=s ' " u#* pla#ts5 <illi#* the *r ;i#* p i#t. /eeds # ' lia*e5 tassels5 sil<s a#d the hus<s ' the c !. I#Jur" t the sil<s =a" i=pair seed set. La"s e**s i# r # =aturi#* *rai#s a#d the i==ature sta*es the# 'eed # the *rai#s. I#'estati #
Klateridae $cara!aeidae
Klateridae 'a=il" eteronychus arator
(ire; r= A'rica# !lac< !eetle
Hhr"s =elidae
!onolepta australis
Redsh uldered lea' !eetle
Hurculi #idae
"itophilus #eamais
Mai-e ;ee%il
C(assi9ica-ion
Ta5onomic name
Common name
4mpacc #ti#ues i#t *rai# st ra*e.
Der=aptera3 La!iduridae He=iptera3 Hicadellidae %icadulina bimaculata Mai-e lea'h pper Apart 'r = 'eedi#*5 the lea'h pper i#Jects a t +i# ;hile 'eedi#* ;hich causes I;alla!" earC. /eeds # the !ases ' c !s causi#* *rai# da=a*e. ?ra#s=its tra#s=it =ai-e d;ar' = saic %irus5 !ut is c #sidered a pr !le= i# 'resh e+p rts. /eeds # " u#* lea%es a#d ste=s. ?u##els i#t ste=s ' " u#* pla#ts causi#* the ce#tral lea%es t ;ither. Ha# cause seri us da=a*e duri#* tasseli#* a#d sil<i#* !" reduci#* p lli#ati # a#d seed set. I#sect da=a*e 'acilitates e#tr" ' 'u#*al diseases. /eeds # lea%es a#d =a" cause se%ere de' liati # at sil<i#* that reduces seed "ield. $ala lividipes Blac< 'ield ear;i* /eeds # sh ts5 r ts a#d ste=s.
Le#tat =idae H = ptera3 Aphididae
$e#ara viridula
Gree# %e*eta!le !u*
&hopalosiphum maidis
H r# aphid
Lepid ptera3 A ctuidae 'grotis spp. (atytricha truncata Hut; r= Mai-e ste= ! rer
elicoverpa armigera
H r# ear; r=
!ythimna convecta
H == # ar="; r=
?h"sa# ptera3 ?hripidae M st c == #l" ' u#d i# ;h rls5 tassels5 ears5 r # the u#derside ' lea%es. $uc<s 'luid 'r = cells. At ear ' r=ati #5 thrip i#Jur" pr %ides e#tr" ' r i#'ecti # !" )usarium spp. $ urces3 G d"er (1891)5 Hei#richs et al. (2000)5 Hu*hes (2007!)5 /arrell & >C,ee''e (2006)5 >CGara (2006) a#d Hertel & R !erts (2006) )ran*liniella +illiamsi Mai-e thrip
<.&.&
*er+e5ra+e pes+s
1amage to crops can be caused by a variety of birds and animals! ustralian native parrots such as Sulphur@crested cockatoos ;Cacatua galerita<= .9
0alahs ;2lophus roseicapilla syn! Cacatua roseicapilla.3 'ittle corellas ;Cacatua sanguinea<= and Scaly@breasted lorikeets ;Trichoglossus chlorolepidotus< can all cause damage to germinating grain and will also pull back husks of cobs on mature plants to remove the grains ;Temby D %arshall 200,= "rice 1::9= Tracey et al! 2009= )K0ara 2009<! 6ative crows and ravens ;Corvus spp!< commonly consume grain during sowing ;Tracey et al! 2009<! "igs reduce yields in grain crops by consuming grain or trampling plants to form bedding or to gain access to the centre of the crop ;*ho8uenot et al! 1::7= "rice 1::9= )K0ara 2009<! #odents such as mice ;!us domesticus< and long@haired rats ;4attus villosissimus< will chew on the stems of young plants to feed on sap ; nimal *ontrol Technologies 200.< or will eat newly sown grain ;Staples et al! 200,<! 'ong@ haired rats caused significant damage to mai+e in the )rd #iver Valley in 1:8, ;- 0overnment 2009<! Iinc phosphide can been used to control rodent infestations in mai+e crops in ustralia ;Staples et al! 200,<! (iseases
<.&./
list of the more common diseases of mai+e in ustralia is given in Table 12! To some e(tent the impact of a disease is related to geographical3climatic factors that may favour the growth and spread of the causal agents2 eg i**erella zeae is common in the therton Tablelands of Lueensland but much less serious in southern Lueensland ;Cughes 2007a<= wallaby ear virus can be a significant problem in the 6orthern Territory because of the abundance of leafhoppers ;)K0ara 2009<! Cybrids have now been bred that show degrees of resistance to a variety of diseases ;see 4arrell D )KNeeffe 2009 for a listing of cultivars used in 6S-<! -hile chemicals are available for control of fungal pathogens2 disease management is mainly through hygienic practices such as regular crop inspection= washing down e8uipment= controlling weeds and volunteers that may spread the disease= controlling insects that may be associated with disease spread or cause damage that increases plant susceptibility to disease= post infection practices such as paddock 8uarantine2 burning of infected plants3stubble= and agronomic practices such as seed treatment2 crop rotation2 avoiding moisture stress in plants2 minimising damage to plants during cultural operations2 harvesting before kernels start to split ;Stovold 1:8.= &eckingham 2007= 4arrell D )KNeeffe 2009<!
Table 12%
,a-ho%en c(assi9ica-ion Bac-erium K#ter !acteriales Fun%us H"p creales )usarium verticillioides (' r=erl" ). moniliforme, ). proliferatum; ). subglutinans /usariu= c ! r t a#d stal< r t L r seedli#* r t de%el p=e#t. Mature pla#ts ;ith r tted stal< tissues5 l d*e easil". (hite 'u#*al *r ;th c %ers <er#els r the e#tire c ! causi#* "ield reducti #. ? +i#s pr duced !" Er+inia spp. $ 't r t 4pper stal<s r t a#d 'all %er.
Common diseases of maize in &ustralia*

,a-ho%en name ;isease 4mpac- o9 )isease
,a-ho%en c(assi9ica-ion
,a-ho%en name
;isease
4mpac- o9 )isease the 'u#*us =a" !e pr !le=atic (see $ecti # 1.1.2).
Gibberella #eae -ase+ual state = )usarium graminearum,
Gi!!erella stal< r t r Gi!!erella ear r t r pi#< ear r t
Ha# ccur as seedli#* !li*ht. Lla#ts =a" die pre=aturel". $tal<s r t a#d !rea< easil". Kar r t causes *rai# "ield l ss. Ha# pr duce a t +i# (Nearale# #e) har='ul t li%est c<. Blac< =asses ' sp res replace the ears a#d@ r tassels. Br ;# sp ts # lea%es5 =a" als cause r t ' the <er#els # the ear. Gre"ish-*ree#5 ;ater-s a<ed sp ts ;hich =a" c %er = st ' lea' a#d reduce ph t s"#thetic area. $tu#ted5 "ell ;5 thic<e#ed lea%esD tassels usuall" d # t de%el p r sh ; a!# r=al de%el p=e#t. Lustules ' r= # lea%es5 =ass ' red!r ;# p ;der" sp res. M re seri us # s;eet c r#. Lustules e%e#l" spread %er lea' sur'ace. Ma" cause de' liati # starti#* 'r = the !ase. Attac<s a#" a! %e*r u#d *r ;i#* part ' the pla#t t ' r= !listers r *alls c #tai#i#* !lac< sp res. Ha# sur%i%e i# the s il ' r =a#" "ears. $eedli#* disease. $eedli#* =a" # t e=er*e r =a" "ell ; a#d die a'ter e=er*e#ce. Li*ht a#d dar< *ree# = saic5 ri#*-sp t r
Micr ! tr"ales
"phacelotheca reiliana %ochliobolus heterostrophus -(ipolaris maydis, "etosphaeria turcica -Exserohilum turcicum,
Head s=ut
Lle sp rales
Ma"dis lea' !li*ht r $ uther# lea' !li*ht
?urcica@?urcicu= lea' !li*ht r # rther# lea' !li*ht
$cler sp rales
"clerophthora macrospora
D ;#" =ilde;
4redi#ales
Puccinia sorghi
H == # rust
Puccinia polysora
L l"s ra rust r tr pical rust r s uther# c r# rust
4stila*i#ales
.stilago maydis
B il s=ut
Oomyce-e L"thiales Pythium spp. L"thiu=
?irus Mai-e d;ar' = saic@ B h#s # *rass
.:
,a-ho%en c(assi9ica-ion
,a-ho%en name
;isease = saic
4mpac- o9 )isease
"ell ;i#* ' lea%es causi#* "ield reducti #. $pread !" aphids. $;eet c r# %arieties5 especiall" supers;eets5 are = st suscepti!le. Lea%es ha%e erect (alla!" Kar *r ;th a#d r ll up;ards a#d i#;ardsD there are #u=er us *alls # the lea' %ei#s. 2$ urces3 $t % ld (189.)5 A"all (1888)5 Bec<i#*ha= (2007)5 Hu*es (2007a) /arrell & >C,ee''e (2006)5 (ats # (2006)
<.&.1 1itrite poisoning
"+her a,-erse associa+ions 8i+h maize
6itrite can be generated by microbial organisms from nitrate and may be taken up by animals through their feed2 such as mouldy hay2 or2 in ruminants2 it may result from the conversion of nitrate! /f2 in ruminants2 microbes cannot convert nitrite to ammonium2 it may accumulate! 6on@ruminants have no mechanism to deto(ify nitrite2 so they are generally susceptible to nitrite poisoning! 6itrite is absorbed into the blood2 leading to a reduction in its ability to transport o(ygen ;o(ygen starvation<! Symptoms of nitrite poisoning are breathing difficulties in staggering3weak animals! /n severe cases2 the animal may convulse and die2 in less severe cases2 cows may abort their foetus ;Calpin D Cides 2002<! "ilo &iller5s disease 0aseous o(ides of nitrogen2 including nitrogen dio(ide ;6)2<2 are released from mai+e after the conversion of nitrates through microorganisms after ensiling ;reviewed in &rightwell 1:92= $verist 1:81= %aw et al! 2002= 'eavey et al! 200.<! The -orld Cealth )rgani-ati # has de%el ped *uideli#es that rec ==e#d # t e+ceedi#* a# a##ual =ea# ' .0 M*@=, (ie 0.02 pp=) a#d 200 M*@=, ;ie 0!1 ppm< for short term e(posure ;-orld Cealth )rganisation 2005<! Cowever2 in the >S 2 measurements of over 1002000 ppm have been documented in mai+e silos ;reviewed in $verist 1:81<! &oth humans and stock may be affected by those o(ides2 which may lead to dyspnea2 pulmonary oedema and death within a few days in severe cases ;$verist 1:81= 'eavey et al! 200.<! /n other ;human< cases2 respiratory distress over a period of 2 F , weeks may be followed by chills2 breathing difficulty2 cyanosis and death within si( weeks after e(posure ;$verist 1:81<! !ycotoxins %ycoto(ins are low@molecular@weight natural products produced as secondary metabolites by filamentous fungi and are to(ic to vertebrates and other animal groups in low concentration ;&ennett D Nlich 200,<! The fungi that produce them typically grow on agricultural products ;eg cereal grains2 rice and cottonseed meal2 groundnuts
and other legumes< before or after harvest or during transportation or storage ;4 ) 1::8<! %ai+e has one of the most serious mycoto(in problems of all crops ;%unkvold 200,<! %ycoto(ins are stable compounds that are not destroyed by processing and may also be present in the meat2 milk or eggs of animals that have ingested them! /n most instances the main source of mycoto(ins for humans is contaminated cereals and legumes rather than animal products ;4 ) 1::8<! The occurrence of mycoto(ins in grain harvested and stored in ustralia is generally not significant because the moisture content at harvest is below that permitting fungal growth ;-ebley D Aackson 1::8<! %any nations have ma(imum tolerated levels of mycoto(ins in food and feed and mycoto(in contamination can lead to reEection of mai+e imports! *ommon mycoto(ins occurring in mai+e are given in Table 1,!
Table 13%
#yco-o5in Ca-e%ory A'lat +i#
+-coto5ins associated with maize grains*

,ro)uce) by Commen-
'spergillus flavus; '. parasiticus
?he 'u#*i als cause c ! r t. A'lat +i#s ca# ccur # =ai-e prehar%est ('a% ured !" hi*h te=perature a#d hi*h hu=idit" at *rai# =aturati # a#d@ r !" stress) !ut are usuall" ass ciated ;ith = uld *r ;th i# st ra*e. >ccurre#ce i# Australia is =uch less c == # tha# that ' a'lat +i#. >ccurs i# Wuee#sla#d ccasi #all". /u= #isi# pr ducti # ccurs prehar%est ;he# pla#t de'e#ces are ;ea<e#ed !" stress. De%el p=e#t ' 'u=i#isi#s i# st red *rai# is u#li<el" !ecause ' the hi*h = isture reOuire=e#ts ' )usarium spp. >' the 110 t +i#s i# this *r up5 de +"#i%ale# l (D>A) a#d #i%ale# l are the = st si*#i'ica#t i# Australia. ?he latter ccurs # =ai-e *r ;# # parts ' the Athert # ?a!lela#d. D>A has ccurred i# the Li%erp l Llai#s re*i # ' A$( ass ciated ;ith c l5 = ist c #diti #s duri#* *rai# =aturati #. /u#*us *r ;s # the *rai# !e' re har%est. Larticularl" ' u#d i# =ai-e *r ;# i# c l5 ;et upla#d re*i #s such as the Athert # ?a!lela#d.
>chrat +i# A
'spergillus ochraceus; '. niger )usarium verticillioides
/u= #isi#
?ric thece#es
). graminearum
Nearale# #e
). graminearum
2$ urces3 (e!le" & Bac<s # (1889)5 , e##i#* (1888)5 Bla#e" (200.)5 /arrell & >C,ee''e (2006)
Cigh or abnormal fluctuations in temperature and drought stress predispose pre@ harvest mai+e to fungal growth and subse8uent mycoto(in accumulation! /nsect infestation is also considered to facilitate mycoto(in contamination because insects act as vectors for fungal spores and insect damage leads to wounds in the plant through which fungal coloni+ation can occur ;%unkvold 200,<! $vidence would suggest that2 in areas where there are insect pests2 mai+e lines genetically modified with the gene coding for the %acillus thuringiensis ;&t< to(in2 that protects plants from insect pests2 have lower levels of mycoto(ins than non@0% lines ;-u 2008<!
51
<./ <./.#
"+her 5io+ic in+erac+ions ycorrhizae
rbuscular mycorrhi+al fungi ; %4< coloni+e the roots of most crop species and mai+e is a mycorrrhi+al host ;%o+afar et al! 2000= Aansa et al! 200,= 0rigera et al! 2007<! /n general2 it is considered that coloni+ation of roots by %4 improves crop productivity for a variety of reasons ;see discussion and references in Sylvia et al! 1::,<! 1ata on the significance or otherwise of mycorrhi+al associations with mai+e are not always clear and2 at times2 are conflicting! There would appear to be some interaction between mycorrhi+al effect and nutrient ;particularly phosphorus< and water status and there is an obvious re8uirement for further research in this area! /n trials using artificially inoculated plants it has been shown that %4 improve plant growth and yield of mai+e by improving uptake of phosphorus and copper particularly under conditions of water stress ;Sylvia et al! 1::,<! There is uncertainty about the specific effects of mycorrhi+ae on root development with Nothari et al! ;1::0< noting a decrease in root length per plant and )sonubi ;1::.< finding an increase! /noculated plants show higher transpiration rates than non@inoculated plants possibly due to the total greater leaf area of inoculated plants ;Nothari et al! 1::0<! >nder optimal growing conditions carbon may be reallocated to the fungi from the plant2 a factor that potentially may reduce grain yield in the crop2 but there is evidence that this is offset by increased uptake of phosphorus by the crop as a result of mycorrhi+al contribution ;0rigera et al! 2007<! Soil tillage affects the relative prevalence of different genera of mycorrhi+al fungi in the mai+e rhi+osphere ;%o+afar et al! 2000<!
SECT!"N =
=.#
0EE(!NESS
0ee,iness s+a+.s on a glo5al scale
-eeds are plants that spread and persist outside their natural geographic range or intended growing areas such as farms or gardens! -eediness in ustralia is often correlated with weediness of the plant2 or a close relative2 elsewhere in the world ;"anetta 1::,= "heloung et al! 1:::b<! The likelihood of weediness is increased by repeated intentional introductions of plants outside their natural geographic range that increase the opportunity for plants to establish and spread into new environments2 eg escapes of commonly used garden plants ;0roves et al! 2005<! *haracteristics in plants that are generally associated with weediness include prolonged seed dormancy2 long persistence of seeds in the soil2 germination under a broad range of environmental conditions2 rapid vegetative growth2 short lifecycle2 very high seed output2 high seed dispersal and long@distance seed dispersal ;Neeler 1:8:= Neeler et al! 1::7<! 1uring development of the mai+e plant the main vegetative growing meristem is transformed into a reproductive meristem which is then removed at harvesting! *oupled with the fact that mai+e does not resprout or reproduce vegetatively ;see Section .!1<2 this means that harvesting effectively destroys the plant! There is2 however2 a significant mass of plant material ;stover< that is left in the ground following harvest and this may be removed2 retained as is for oversowing with another crop2 incorporated into the soil after tilling or sometimes burnt ;#obinson D Nirkby 2002= 1hugga 2009<! s discussed in Section .!,!,2 corn seeds generally remain tightly attached to the cob
and are collected during harvest or may be transferred to the ground if the ear becomes detached prior to harvest ;eg as a result of insect or disease damage<! /n the latter instance2 the hundreds of mai+e kernels inside can geminate in the following season but2 because of competition for light and soil2 very few are able to reach maturity ;1oebley 200.<! Volunteer plants Zself sown plants derived from seed of a previous crop ;&rookes et al! 200.<[ can also occur as a result of harvesting or transport! lthough mai+e volunteer plants occur consistently2 they are not regarded as a serious weed problem e(cept perhaps when grown in rotation with soybean ; ndersen et al! 1:82= *4/ 1::.= $astham D Sweet 2002= )wen 2005<! Carvest grain losses can be as great as 207 kg3ha of mai+e grain and occur when2 for e(ample2 harvest machinery does not efficiently gather ears or shell the grain from the cob2 or when weeds plug grain sieves ; 0&/)S 2005a<! Volunteerism is a greater problem in no3low tillage systems when the dropped seeds remain near the soil surface! /t is suggested that volunteer plants rarely produce viable seed for the ne(t growing season ; 0&/)S 2005b<! /n areas with cold winters2 frosts will normally kill volunteers ;$* 2000< and feral populations of mai+e have not been observed in $urope ;*)0$% 2008<! -ith the recent adoption of genetically modified ;0%< herbicide resistant ;glufosinate ammonium= glyphosate< mai+e cultivars ;see Section 2!.!2< there has been concern with management of potential 0% mai+e volunteer plants! The fact that Zea mays ssp! mays has no vegetative means of spreading combined with its complete dependence on humans for survival2 means that it represents a minimal weed risk! Cowever2 some of its relatives may have some weed potential eg:
&alsas teosinte ;Zea mays ssp! parviglumis< can invade fields in %e(ico and is therefore considered weedy in places ;reviewed in 1oebley 200.<! /n central %e(ico2 Iea mays ssp! me(icana is a troublesome weed of mai+e fields where2 until flowering2 the plant can be difficult to distinguish from domesticated mai+e ;reviewed in 1esEardins D %c*arthy 200.<! #andall ;2002< notes that Z. mays ssp! mexicana and Z.mays ssp! parviglumis are listed as naturalised weeds in the >S = Z. luxurians is a cultivation escape in 0uyana= Tripsacum andersoni is a weed in 0uyana and "eru= T. lati&olium is a cultivation escape in "uerto #ica= and T. laxacum is a cultivation escape in several countries! 0ee,iness s+a+.s in A.s+ralia
=.&
0roves et al ;200,< assigned a rating of J1B to Zea mays ssp! mays and Zea mays ssp! mexicana in natural ecosystems indicating that they are \naturalised and may be a minor problem but not considered important enough to warrant control at any location]! /n agricultural ecosystems2 the two species are also not considered a problem ;0roves et al! 200,<! Z. mays ssp! mexicana can be found in north Lueensland and -estern ustralia ;"heloung et al! 1:::a< and is also cited in #andall ;2002< as being naturalised in ustralia! Tripsacum dactyloides has been collected in ustralia ; ustraliaKs Virtual Cerbarium2 http:33www!cpbr!gov!au3avh!html< and is listed by #andall ;2002< as a naturalised weed there!
5,
=./
Con+rol meas.res
-hile mai+e is not a weed problem as such2 it may be necessary to remove volunteer plants that arise in a subse8uent non@mai+e crop following a rotation! 4armers have traditionally used herbicides ;eg glyphosate or graminicides such as diclofop ; ndersen et al! 1:82<< and tillage to control volunteers! -ith the increase in the use of glyphosate@tolerant mai+e cultivars ;especially in rotation with glyphosate@tolerant soybeans< herbicides such as setho(ydim2 clethodium2 feno(aprop@p@ethyl2 flua+ifop@ p@butyl and 8ui+alofop@p@ethyl can also be used to control volunteer mai+e ;&eckie D 0ill 2005= Soltani et al! 2005<!
SECT!"N >
P"TENT!AL F"R *ERT!CAL 'ENE TRANSFER
Vertical gene transfer is the transfer of genetic material from parent to offspring by reproduction! 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 re8uires that three criteria are satisfied! The plant populations must: 1< overlap spatially ;sympatry<= 2< overlap temporally ;including flowering duration within a year and flowering time within a day<= and ,< be sufficiently close biologically that the resulting hybrids are fertile2 facilitating introgression into a new population ;den 6iEs et al! 200.<! s discussed in Section 12 Z. mays ssp! mays is the only cultivated species within the genus Zea and the other species and subspecies Zoriginally classified as races ;-ilkes 1:99<[ are wild grasses2 referred to as teosintes! Three of the annual teosintes are subspecies of Z. mays2 namely ssp! mexicana ;central %e(ico<2 ssp! parviglumis ;southern and western %e(ico< and ssp! huehuetenangensis ;western highlands of 0uatemala< ;1oebley 1::0<! )ther species of teosinte are Z. diploperennis ;a diploid perennial<2 Z. perennis ;a tetraploid perennial< and Z. luxurians ;an annual<! s many publications tend to discuss crossing between teosintes and mai+e generally2 it is sometimes difficult to be able to distinguish between intraspecific and interspecific crosses within the genus2 so these have been considered together! >.# Na+.ral in+raspecific an, in+erspecific crossing
ll the various types of Z. mays ssp! mays freely cross@pollinate and form fertile hybrids ;"urseglove 1:92<! Since mai+e is predominantly outcrossing2 intraspecific crosses are highly likely between nearby plants if flowering time overlaps and other factors ;see Section .!2!,< are favourable! Volunteers are not likely agents of gene transfer in mai+e since their occurrence is minimi+ed by the fact that the mai+e cob cannot shed seed naturally ;see Section .!,<! )utcrossing has been discussed in Sections 2!,!1 and .!2!,! -ind pollination can occur between mai+e crops over hundreds of metres but the relatively large weight and diameter of the pollen grains favours most pollen deposition within appro(imately 70 m of the source plant ;#aynor et al! 1:92= 'una et al! 2001= ylor 2002= Aaros+ et al! 200,< and there is little or no cross pollination at ,00 m ;'una et al! 2001<! Calsey et al! ;2005< have considered both time and distance components of pollen@mediated gene flow in mai+e and concluded that2 under their e(perimental conditions2 200 m
was sufficient to reduce outcrossing to R 0!1H! lso of significance in considering the likelihood of cross@pollination in mai+e is pollen competition! The fact that a single plant can produce millions of pollen grains ;see Section .!2!1< means that2 even if there is long distance spread of pollen from a plant2 the pollen will be greatly outnumbered by JlocalB pollen ;&annert D Stamp 2009<! /ronically2 measures to reduce pollen production such as detasseling or cytoplasmic male sterility may actually 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 physical separation of crops to minimise gene flow ;see Section 2!,!1<2 and a separation distance of less than or e8ual to ,00 m is standard worldwide for production of certified seed while basic seed has a higher re8uirement ;eg .00 m in 4rance ;%essean et al! 2007<<! Separation distances have also been discussed in considerations of gene flow between genetically modified ;0%< and non@0% mai+e crops ;Cenry et al! 200,= $astham D Sweet 2002= Stevens et al! 200.= 0oggi et al! 2007= %esseguer et al! 2007= Sanvido et al! 2008<! The variation in results reflects the range of factors that can be taken into account when defining the distances ;eg see discussion in Section .!2!,<2 and especially the level of gene flow that might be tolerated2 but it would appear that for gene flow rates of R 0!1H2 recommended distances are similar to those used in non@ 0% certified seed production ;Cenry et al! 200,= 0oggi et al! 2007<! comprehensive discussion of the factors to be considered in gene flow between 0% and non@0% mai+e is given by %essean et al! ;2007<! )f importance in restricting gene flow between 0% and non@0% corn is surrounding the 0% corn with non@0% corn so as to reduce the density of any 0% pollen ;0oggi et al! 2007<! Various incompatibilities e(ist both between and within subspecies of Z. mays and are thought to have evolved to prevent indiscriminate hybridi+ation ;Nermicle 1::9<! )ne system that has been particularly studied is concerned with pollen@pistil incompatibility affected by si( or more alleles at the gametophyte-' ;ga'< locus and involves the arrest or retardation of ga' pollen tubes within a'-s6- silks resulting in a pre+ygotic barrier to crossing ;Schwart+ 1:50= Nermicle 1::9<! The teosintes were formerly placed in the genus 2uchlaena ;*ollins 1:25= %angelsdorf D #eeves 1:,1<! %ai+e and all of the teosintes ;e(cept Z. perennis< are se(ually compatible2 and it is known that they naturally produce fertile hybrids in %e(ico and 0uatemala where they share a common distribution ;-ilkes 1:99= 1oebley 1::0<! /n particular spontaneous hybridi+ation occurs between Z. mays ssp! mays and both mexicana teosinte and parviglumis teosinte ;$llstrand et al! 2009< with introgression being most common with ssp! mexicana ;4ukunaga et al! 2005<! Cybrids of ssp! mays ( ssp! mexicana crosses have statistically significant heterosis compared to the wild teosinte but not when compared to the cultivated parent ;0uadagnuolo et al! 2007<! 1espite the ease of crossing2 gene flow occurs at low fre8uency and all the subspecies still co@e(ist as genetically separate entities ;&alta+ar et al! 2005= 4ukunaga et al! 2005<! >nusually2 the flow of genes has occurred in both directions ;reciprocal introgression< ;-ilkes 1:99= 1oebley 1::0< although a number of factors tend to favour gene flow from teosinte to mai+e rather than from mai+e to teosinte ;&alta+ar et al! 2005<! There is also evidence of a restriction to crossability in some populations of Z. mays teosintes when teosinte is the female and mai+e the male 55
parent and this has been linked to a teosinte gene or gene cluster known as Teosinte crossing *arrier' -Tc*'. ;$vans D Nermicle 2001= Nermicle 2007<! The Tc*' locus occurs within the chromosome@. cluster of teosinte known as Teosinte /ncompatibility *omple( and represents a barrier to crossing that occurs before fertili+ation! /t is loosely linked to the a' locus! $vans D Nermicle ;2001< have suggested that the transfer of Tc*' into mai+e may be useful in avoiding contamination of one commercial variety with another! The likelihood of intraspecific or interspecific crossing occurring naturally in ustralia is remote! Z. mays ssp! mexicana imported from South merica can now be found in 6orth Lueensland and -estern ustralia ;"heloung et al! 1:::a< but there have not been any reports of introgression with commercial mai+e cultivars! >.& Crossing .n,er e3perimen+al con,i+ions
These crosses2 while possible under controlled conditions2 do not occur naturally and therefore are of significance only in the conte(t of broadening an understanding of unaided gene transfer ;)$*1 2007<! >.&.# !n+erspecific crosses
*rosses between Z. mays ;2n?20< and Z. perennis ;2n?.0< may normally yield only 0!1 F 1H viable seed because of endosperm collapse after appro(imately 21 days post pollination! Cowever2 hybrids ;with 2n?,0< have been obtained following embryo rescue ;del *armen %olina D 0arcia 1:::<! Z. perennis is not found in ustralia! >.&.& !n+ergeneric crosses
There has been considerable speculation on the evolution of cultivated mai+e2 including suggestion that intergeneric introgression between Zea and Tripsacum ;that are both members of the sub@family "anicoidea2 tribe %aydeae< may have played a role Zsee e!g! discussion in de -et D Carlan ;1:9.<= $ubanks ;1::9<[! *onse8uently there have been a number of attempts to hybridi+e Zea mays and Tripsacum spp! although all of these have been under controlled conditions and there is no suggestion that hybridi+ation occurs naturally! The Tripsacum genus comprises rhi+omatous perennial grasses distributed from northern >S to "araguay in South merica! The base chromosome number is (?18 ;$ubanks 1::9< compared to the base chromosome of mai+e ;(?10< and therefore the cytogenetic interactions in the hybrids have been of interest! Tripsacum dactyloides has been collected in ustralia ; ustraliaKs Virtual Cerbarium2 http:33www!cpbr!gov!au3avh!html< and is listed by #andall ;2002< as a naturalised weed! Tripsacum with a basic chromosome number of 2n?18 is the genus most closely related to Zea and crosses between these two genera are possible! T. dactyloides and T. andersonii are used as fodder plants in the tropics ;*ook et al! 2005a= *ook et al! 2005b<! T. dactyloides ;$astern gamagrass< has been collected in ustralia ; ustraliaKs Virtual Cerbarium2 http:33www!cpbr!gov!au3avh!html<! *urrent research addresses the value of Tripsacum grasses and their hybrids as fodder plants in ustralia ;eg mai+e ( T! dactyloides hybrids as in Shavrukov et al! 2007<! $arly crosses ;using Tripsacum as the pollen donor< ;%angelsdorf D #eeves 1:,1< showed that some degree of successful fertili+ation could be obtained only if the
distance the Tripsacum pollen tube had to grow was reduced ;eg by shortening the mai+e styles<! /t appears that genetic transfer from Tripsacum ;e!g! T. &loridanum7 T.dactyloides7 T. pilosum7 T. lanceolatum< to Zea mays is comple( and that2 depending on the choice of Tripsacum parent and the events occurring in early backcross generations2 at least 5. chromosome combinations can be obtained ;Carlan D de -et 1:99<! The hybrids are sterile or somewhat female fertile and repeated backcrosses to mai+e eliminate the Tripsacum chromosomes! "olyploids in the genus Tripsacum show diplosporous apomictic reproductioni ;0rimanelli et al! 1::8< and this has been e(ploited commercially in mai+e3Tripsacum hybrids to introgress diplosporous apomictic reproduction into a mai+e background= this would eventually allow the establishment of immortali+ed commercial lines of apomictic mai+e ;Nindiger D Sokolov 1::8<! )ther traits that have been obtained in mai+e3Tripsacum hybrids include high silage biomass ;Shavrukov et al! 2007<! *rosses between Tripsacum and Z. mays teosintes have not been successful but fully fertile hybrids ;2n ? 20< have been achieved between Z. diploperennis and T. dactyloides using Tripsacum as the female parent ;$ubanks 1::5= $ubanks 1::9<! These hybrids2 referred to as Tripsacorn ;$ubanks 1::2< can be crossed with cultivated mai+e and offer a pathway for the transfer of useful genetic material into commercial mai+e varieties ;$ubanks 200,<! The tribe %aydeae also includes five siatic genera ;Coix7 "clerachne7 $olytoca7 Chionachne and Trilo*achne< ;-atson D 1allwit+ 1::2< but of these there has only been a report of Z. mays being crossed with Coix lachryma-8o*i ;Carada et al! 1:5.<! /n this instance2 hybrid seed was obtained in about 7H of crosses but only when Coix was used as the female parent! number of Coix spp! are permitted imports to ustralia ; L/S 2008<! C. gasteenii is considered a threatened species in the *ape 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 ;"accharum o&&icinarum F tribe ndropogonae < with 2n ? 80 and Z. mays with two additional & chromosomes2 using mai+e as the pollen parent ;Aanaki mmal et al! 1:92<! The individual2 whose cells contained chromosome numbers varying from 52 F 582 survived ,0 years through clonal propagation and was induced to open its inflorescence by application of gibberellic acid! )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 "ooideae2 chromosomes of the "anicoideae are eliminated soon after fertili+ation! $(amples include:
%ai+e ;pollen donor< crosses readily with he(aploid ;2n ? .2< wheat ;Triticum aestivum< although double fertili+ation is rare and development of the embryo proceeds without the formation of endosperm ;Ihang et al! 1::7<! The embryo will therefore abort2 unless cultured in vitro2 because of a lack of nutrition normally supplied by the endosperm! The mai+e chromosomes are eliminated
i pomi(is is a genetically controlled mechanism in which an embryo is formed without union of male and female gametes! 1iplospory is a form of apomi(is in which the embryo develops from a diploid egg that has derived from an unreduced megaspore mother cell! The resulting individual is therefore genetically identical to the maternal parent ;Nindiger D Sokolov 1::8<!
59
during early embryogenesis resulting in the formation of a wheat haploid ;'aurie D &ennett 1:88<! This occurs because of a failure of the kinetochores of the mai+e chromosomes to attach to spindle microtubules of the metaphase plate during cell division ;%ochida et al! 200.<!
/n controlled crosses of he(aploid ;2n ? .2< oat ;Avena sativa< with mai+e2 a proportion ;appro(imately ,1H< of the embryos may contain 1 F . mai+e chromosomes in addition to a full complement of oat chromosomes and are therefore referred to as partial hybrids ;#iera@'i+ara+u et al! 1::7= Nynast et al! 2001<! )at3mai+e hybrids represent useful material for mapping the mai+e genome ; naniev et al! 1::9= Nynast et al! 200.<! The hybrid plants can only be grown following embryo rescue! Ienkteler D 6it+sche ;1:8.< made a number of crosses within the cereals2 including barley ;Hordeum vulgare= 2n ? 1.< and rye ;"ecale cereale= 2n ? 1.< as female parents with Zea mays as the pollen donor! The crosses were made on in vivo @grown plants and no attempt was made at embryo rescue! 6o embryos developed in the H. vulgare ( Z. mays cross but globular embryos formed in some of the ". cereale ( Z. mays crosses although these degenerated after 7 @ 10 days presumably because of poor or no endosperm development! 1evelopment of embryos from barley florets pollinated with mai+e has been obtained following embryo rescue ;*hen et al! 1::1<! "lants grown on from the embryos were largely sterile haploids ;2n ? 9< and there was no cytological evidence of retention of entire mai+e chromosomes!
REFERENCES?
http:33www!abareconomics!com3publicationsPhtml3cr3crP083cr08PAune!pdfhttp:33www !defra!gov!uk3$nvironment3acre3advice3pdf3acrePadvice28!pdfhttp:33www!agbios!com3 cstudies!phpT book?$S Dev?%)6810Dchapter? ppd(1Dlang?http:33www!agbios!com3cstudies! phpT book?$S Dev?%)6810Dchapter? ppd(1Dlang?http:33www!agricentre!com!au3co ntent3*4%ai+e!pdfhttp://www.animalcontrol.com.au/mice1.htmhttp://www.aqis.gov. au/icon32/asp/ex_querycontent.asphttp://www.grainscouncil.com/See s/!S!/!S!_ "orporate_#lan.p $http://www.as$.asn.au/p $/!S!%2&'(')*%2&!++,!%2&./#0.)%2&2&&12&3%2&'inal %2&1&!ug&3.p $http://www. pi.nsw.gov.au/agriculture/horticulture/vegeta4les/ ise ases/ iseases2an 2 isor ers/4oil2smut2o$2 cornhttp://www. pi.nsw.gov.au/agriculture/horticulture/vegeta4les/commo ity/swee t2 cornhttp://espace.li4rary.uq.e u.au/eserv/,5:6167/!gronomy_o$_8ai9.p $http://w ww2. pi.ql .gov.au/pigs/:::3.htmlhttp://www.inspection.gc.ca/englis h/plaveg/bio/dir/dir9411e.pdfhttp://affashop.gov.au/PdfFiles/ PC12791.pdfhttp://www.coge .net/ContentFiles/!"!#2$% !2&2!advies&2!hernieuwing&2!i port&2! ais &2!'t11&2!eng.pdfhttp://sgrl.csiro.au/aptc199"/1!(collins. pdfhttp://www.tropicalforages.info/)e*/Forages/+edia/,t l/ -ripsacu (andersonii.ht http://www.tropicalforages.info/)e */Forages/+edia/,t l/-ripsacu (dact*loides.ht http://www .nal.usda.gov/research/ ai.e/http://www.dpi.vic.gov.au/dpi/ nreninf.nsf/childdocs/% 71/"!91F#770#202412#2"'$!!!4F$'2% F$#'1C1C4!#4922'C12#2'C"!!!29104% 142$"'2/210!4"19412#20/1!!27$00F% /9"2F!9/C29'/1!0C12#2'CF!!!''/F23 openhttp://www.dpi.vic.gov.au/dpi/nreninf.nsf/v/4aac7ca!f4c b4e#1ca2#74!e!!7ee$92/4file/hardheads.pdfhttp://www.dpi .5ld.gov.au/cps/rde/dpi/hs.6sl/22($491(/71(,-+8.ht http:// www.dpiw.tas.gov.au/inter.nsf/9ebPages/CP1:%#;<==-3 openhttp://reports.eea.eu.int/environ ental(issue(report(2! !2(2"/enhttp://ec.europa.eu/food/fs/sc/scp/out77(g o(en.pd fhttp://www.freepatentsonline.co /PP!7977.ht lhttp://www .freepatentsonline.co /2217492.ht lhttp://www.dpi.nsw.go v.au/agriculture/resources/soils/salinit*/crops/tolerance% irrigatedhttp://www.fao.org/docrep/t!$9#e/-!$9#/!!.ht >C ontentsftp://ftp.fao.org/es/esn/food/9"9!1/.pdfhttp://www. dpi.nsw.gov.au/pubs/su er%crop%production% guidehttp://www.foodstandards.gov.au/(srcfiles/Final &2!7?--1'&2!2!!2&2!Food&2!Co position&2!-ables &2!%&2!+a* &2!2!!7.pdfhttp://www.cropscience.org.au/icsc2!!4/poster/
E ll websites cited in the #eference 'ist were current as at September 2008
5:
$/4/1/4$9(george lc.ht http://www.regional.org.au/au/asa/ 2!!2/poster/agrono */42"2(dri@berrd.ht >-opAfPagehttp:// affashop.gov.au/PdfFiles/PC127"1.pdfhttp://www.dpi.vic.gov .au/0PB/nreninf.nsf/9e#"221e""!ba9e44a2#2c24!!2$eb2e/1 421$"f7#ac##e$fca2#717f!!!d7$92/4FB8//1C!7!1.pdfhttp:// ip world.u n.edu/chapters/ ai.e.ht >-hripshttp://www.dp i.nsw.gov.au/agriculture/field/field%crops/protection/insect% ite% cropshttp://www.dpi.5ld.gov.au/cps/rde/dpi/hs.6sl/22($247(/ 71(,-+8.ht http://www.dpi.5ld.gov.au/cps/rde/dpi/hs.6sl/2 2($24#(/71(,-+8.ht http://www.dpi.5ld.gov.au/cps/rde/dpi /hs.6sl/22($72!(/71(,-+8.ht http://www.dpi.5ld.gov.au/cp s/rde/dpi/hs.6sl/22($#!2(/71(Print.ht http://www.dpi.5ld.g ov.au/cps/rde/dpi/hs.6sl/22($27"(/71(,-+8.ht http://www. nature.co /hd*/@ournal/v2"/n1/abs/hd*19721$a.ht lhttp://w ww.w*ong.nsw.gov.au/environ ent/9eeds(categor*(one(9i tchweed.pdfhttp://www.ci *t.org/english/docs/proceeding s/africa/pdf/$7(;ana piu1.pdfhttp://www.ci *t.org/abc/ge neflow/geneflow(pdf(engl/geneflow(cross.pdfhttp://www.fre epatentsonline.co /#71!$27.ht lhttp://www.ces.ncsu.edu/d epts/pp/notes/Corn/corn!!1.ht : / / . . . . / / . / / 7 771 12 8 47 2 5 7 4 0900829 / / 1 3 14_ 0 7 . : / / . . . . / / . / / 1 7 4 4 6964 3 9 2 5 7 4 1 1 0002 4 8 / / _ _ _ _ _ _ . :// . . . . / / . / / 89 7 8 4 1 7 6 2 4 2 5 6 8 3 0 0 0 4 2 6 9 0 3 4 7 7 0 7 4 2 5 6 7 0 0 8 1 1 9 2 95 19 3 5 4 9 3 6 6 4 2 5 6 0 0 2 7 7 6 6 2 5 1 4 8 2 8 3 5 2 5 6 0 0 0 3 ? :// . . . . / / / / 2 0 . :// . . . / / 1
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= 1 3 7 9 6 : / / . . / / 2 0 0 7 / 1 0 /071016101454. :// . . / / / / . /28_ _ . . / / . :// . . . . / / / /
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/content/* "4)24262v#74#!/fullte6t.pdfhttp://www.isb.vt.edu/ne
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71

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 mays L. ssp mays (maize or corn)

Version 1: September 2008

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

Zea species and subspecies*

Zea mays ssp mexicana ($chrad.) HH Iltis

Zea perennis (Hitchc.) Ree%es & Ma#*elsd. 2$ urce3 4$DA (2001)

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

"R!'!N AN( C)LT!*AT!"N

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

/mportant ways of processing grain mai+e include

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

Characteristics of some of the current maize growing regions in NSW

Au*ust : >ct !er Blac< %ert s l222

$epte=!er : A %e=!er 2222?ra#siti #al red-!r ;# earth

$epte=!er : A %e=!er Blac< earth (l a=)22222

Au*ust : Ba#uar" Red 'ria!le l a=

Characteristics of some of the current maize growing regions in !"#

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

4)ea( p(an-in% -ime8

>ct !er Ba#uar"

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

!!! !!1 !! !!< !!>

2 Data ta<e# 'r = Australia# $eeds Auth rit" Ltd. (2006!)

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

Pollen ,ispersal an, pollina+ion

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

N.+rien+ componen+s of +he maize 9ernel

/elati,e content 012 of nutrient components in parts of the maize 3ernel*

,ro-ein Cru)e 9ibre Cru)e 9aS-arch Su%ar

3.6 97.6 1.0 6.3 0.3.

2 Data ta<e# 'r = /A> (1882)

The Biology of Zea mays L. ssp mays (maize or corn)

Office of the Gene Technology Regulator

0.0. 0.01 0.3 0.7. 0.01 0.7 1.26 0.02 1.3 0

2$ urce3 /$AAN (2007)

The Biology of Zea mays L. ssp mays (maize or corn)

4mpacc #ti#ues i#t rai# st rae.

Gree# %eeta!le !u

Ma"dis lea' !liht r $ uther# lea' !liht

?urcica@?urcicu= lea' !liht r # rther# lea' !liht