(FOL Plant embryo development establishes a basic body plan. Establishing the Root-Shoot Axis In plants, three cimensianal shape ad lattag the amount and pattern of ell very first ecll division is resulting in wo dife [Spt cell pes: Eat in embrye development mint calls cm (ge ec ta wide ange of coll and org ype, incon Teaver Ac development procede the ext with multiple ‘petentids ae reriaed to regions called meristems. Many sc by re Bren the ‘ene adding cal the growing root and shoot tp afer sermination. Tho generate the large numbers of cll ‘needed te frm leaves, flawers, and al ether components of eee plane Aetel cried ol pes eee ple, divide every 17 hours, praducing hal milion cell 3 (Sey ivan actly growing com plant. Lateral meristems ‘Sn caure an increas inthe girth of some plants, while in {ercaliry meriseme within te sem allow for elamsion. Tn addition to developing the root-shoot sais im em bryogeness, cel differentiation eccurs and three asic t= sor systems are established. “These are the dermal, (gromal ul eecdir tae puiane eed Ve we coke yatcmed. Thee: tissue gytcis contain various ecltypox that can be highly difscntnted for specific fonctions. ‘These ttsuc sntems arc organized radially around the rookahoot ax While the embryo is developing, two ather exitical reste are cursing, A. Sood supply catabcea dat will sonpert dae cmbryo during germination w ie it gains pho fenynthotic-cxpacy. This starts with the second tina” sion event that produces endeqperm in angiosperms. Sec fondly, ovule tise (Irom the parental sporophyte) AiFerentists to form x hard, protective conering around the embeyo. The cad (orale containing the embryo) then enlers 3 dormant phase, sigmaling the end of embryogene sia Loreal grade Ger amigas ape: tur, and ligh) can break dormancy and tigger 2 cxcade Finer evens roweling in germination. Early Cell Division and Patterning ‘The firs dvsin of the ferulied cag in a flowering plant fe myrameitic and generate celle with two different fine (Ggure 402). One daughtcr ells smal, wath dense oyu ‘lam. That cell, which will become the embmyo, bine to (Bide peated in diferent planes, al of el, ‘The other daughicr cell divides repeatedly, forming an longated racure called 2 mepouar, which lis the em (pe eee te of toed "The gee aorinse min ae cg eye. The moat-shoot axis also forms at this time. Cells ES Does eiesaet a sear gcaeaaeeceea ancien. snc ag ee Sapo eisee ee eres treme Sorte gi Soe ee ae ioe naamn ior orats See Seer ip ining oe aegis oe Sie ean Stated omit errors Eiene arent eee phyte. This axis is first cxtablished by the point af sperm ies cg eaeaeree Se eee ere oee eed caoemcam gran eae TS Teirmiae ee Steen nasties Rietrairinge gerne cits Beige cca isin sece tae eis oereees etree errant =a rt leer tence eee pee peat ener nrs eee eee Sarees “The embryo suppreses development of dhe suspensor a second embry. Ths aspamor murano rails has eect (pear in eninge develope lloed yey te feo ‘dexclopracnt in the emibeyo followed by cmbryolke devel= ‘opment ofthe sspensr (figare 404). Frm this. one cn ‘snchude thatthe embryo nermally prevents the susponsor from developing ito a second emo ‘Eay in embryogenesis he rot soos ais is (Chapeer $0, Ely Plane Developer 797Establishing Three Tissue Systems Tare bk toes iFieeatite while the plane conbeys i sala ball of cally the globular stage (Ggure 40.5), but no ‘ell mowerents ar Involved The foto consists of the fete ol rx peat Selege ctl tere deel ‘isa. Thee col alone alays dvd with their ell plate \pespacer to le tees “Tae pepsi atk? ‘avir layer af cll, Dermal tess produces ccs chat pro- tect the plant Eom deication,incuing the somata that ‘epee opal lowe ty cea gp coche al alae: wate fos The balk of the ernbryonic Entcrior consists of gourd we cll tat cven tally fanetom in Food and water Storage. Lastly, preamiem at che core af the embryo is Sentinel tn for the future salen tue responsible For eevee eae eet Root and Shoot Formation “The root-choot are iecsabliched during the globular sage of development. The shoot apical merscm will ler give Saas iy eee calscreelly eopepeeetien aemetere= While huh the shact and race meristers are sical meris— tems; thelr formation ix controlled independently. This conclusion is supported by mutant analysis in Arbidons where the shutmerisemle sim) wntant falls i peoduce 2 Wiable shoot, but does prac a root (Figure 4049. Simi Ibex, root meristcm-specific gener have been identified. For example, monopteraus mutants af Arabidepa lack roots, ‘The hormone auxin may play a rc in roetshoot sis formation. Atti ison of si dane of armnomes that FIGURE 406 ‘Shooc-speatic genex specify formation of the snc apical fears ignmiaron Amtannsry bars ay ‘emma eocmerinam, bur fo produces sho mera regulate plane development and funation that we will = ‘lor in mare dct ater inthis unit ‘At you uy the development of roots and shoots after gerinason, yom will oder that many ofthe mame pattems Stimus diferent tion seen inthe embryo ae eeerate in (he splcal merits Memember thet there are alse many crcits decal colic inthis chapter that re emg to cmbryogcnesis. For cramp, she LEARY COTYLEDON FIGURE 405 Early devcoprmernl sages of rabid bali, 2) Ex cll vison ha podced he ebay and supers (8| lable age (ei) Tear shaped mage 708 Pare XI Plane Growth nd Reprdacon(se in Araidops active in carly and Lae embryo devel- ‘opment and may be respanibl for maining ah eribey- cccoetorement Te pomle to tan de gee ere in development using rcoambinant DNA techniques (sc Ghapeer 3). that cae, mabeyor ean Form am lees! Morphogenesis ‘The lobular stage gives rc to a hear-shaped embry in cone groepof anpiowperm (the dicts sce Fpare 40.5) and = ball ith» bale or w ingle sic iv anather group (se monocot). The bulges are cotyledons (“irs leaves) and 1 produced by the embryonic cll, nat the shoot spit mmerintom that ioyins forming during the globular tags. ‘This proces, aed murphopnens (generation of form), re- sles fom changes in planes aed ras of ell iin. Be~ (aus plant cells carne more, the for of plat bodys largely determined bythe plone in which ells divide and Appel keeps be eA deg wt ecco Cally aftr they form. Both micratbule and actin play = rele in estlliching the portion ofthe cell plasewhich de~ ‘ermine the diretion of division, Plane growh-romuiators ‘sel other factors inducnee the orientation of undies of eaeroleten Hlc Se of te plana eee ‘Those microtubules also guide eclulose deposition as the cll ll See eel etl A nr a etc Jing its final shape. Fer cumple if you sart with x box and reinforce four ofthe six sides more heavily with cellulose, ‘he col will xqund aad grow in the direction of the sw sds with Im einforseraent. Mich bing learned a the ell bolewieal level about morphogenesis Frm reutants that dive, but cannot contol heir plane of cell division or the diresion of cel expansion, FIGURE 407 ‘scorn and bean Te ora ‘ere! ha endenpere thats peer Seances adnan ban hs deappered the beam ems cedonseate over fod menage oa Food Storage “Throughout embryogenesis there is the production of starch, lips, and proteins The aed storage prnicins are fo secede ht the pene coding fox them were the Bt Cloning target for plane molecular bislists. At noted Shaper 37, the cralutonary trend inthe plants kas been tuward increased protection ofthe embryo. Onc way this it accomplished fc through parental sporephyte Empat tel a he reepeteee Be eephirgree Or osperme the sarpcheor scrvcs only to pach the emirpe closer to the gamctophytic nutrient source produced by fisiple muclar divisions withowt cll diesem). This bape [pens concurrently with the development of the endorpersn (gros coy in aglonperms although donb Raeliration Fas een abnor in the gyemnorperm Epa) which may Be extensive or minimal, Endowperm In eocomat is the Srlb” and fet liquid ores. Tn corn the cachmperen fe slid and in popping cor cxpande with heat to form the clible part of popcorn. In peas and beans the endesper= i used up during cznbryo developement and nutrients are stored in thick, fleshy entyledans(Ggure 40,7). The pho- teaynthctic reacbinery i bil ix renporactolight. Soft ie (citral tha coi hare stored nutrients to aid im germina- on until the growing sporophyte can photosmthesze. A seed beri toe cpl wll ne op al fs morro before racking the surfser anid sunlight. “Aker the rotshoet ais established radial deve Sse meer, ad a eared food mappa formed ‘rough costo cell division and expansion.Plant Development ‘At the most baie level the developmental paths of plants aad animals share many key elements. However, the mech~ ‘alam ured to achieve body form are quite diffrent. ‘While wnimal col fallow an orchertnted secs of tore ‘ments during desclopenct, plant celle arc encase within sf elllone walls and therefore, canner move. Each cell ‘na plants fied iruo pition when it is created. Instead of ‘ming cll migration, plants develop by bulling thelr bod Fe outward) reading new ports Rom special groape af sl ‘renewing cells called meristems. As meric cll contin~ tally divide, they prodace cell that can diflerentise into. the tins of the plant. “Another major difference between animals and planes is that mont animals are mobile and can move away Fram ‘unfavorable circumstances, while plants are anchored in ‘poston and most simply endure whatever environment they experience. Plants compensate for this restriction by ‘elating the rules of devclopment to accommodate Iocal (Greumstances. Instead of creating 2 body in which every [Bart opeclicd to have fied ser and location, plant ‘stmbles its body fam afew sypes of modules, such a5 eaves, roots, branch nods; and flowers. Each module ‘hus a rigidly controlled structure and organization, bt how the modales are tized is quite Geuible. Ae plant develops i simply adde more modules, with the eniton ‘mcat having 1 major influence on the type, number, sae, ‘zed location of what is added. In this way the plant i= able to adjust the path of its development to lecal Early Cell Division “The fi division of the fertilized emg im fawering plant fs ofcerter, os that ame of the dauphter cli smal, wit emwc eytoplanm (Bigure 17-4). That col the Fate era Layo, begins tp divide repeal, forming a ball of ecil. ‘The ocher daughter eel slo divides epestedly, forming clongated stracture called a suspensor, which links the ccnbrye to the nutrient tise of the sed. The suspeasor ‘abe provides = route for nutrients to reach the developing ‘sey, Jas asthe areal cmbryo acquires its iil ans 35 {Fall mace formed during cleavage divisions, so the plant ‘Galeye Forms i root-shoot acs a this time. Celle near the ‘mipetoor ae destined to form 2 root, while those at the ther end of theses lemately becomes shoot ‘Tissue Formation “Thrce basic tnmes differentiate while the plant crbryo is still ball feels (gure 17.80, analogous tothe formation fof the three germ layers fn animal embryos, although in plants, no cell movements arc involved. The eutcrmost ‘ellen a plane creye become epidermal cells. Tae ball (SF the cuyone inevor comsite of ground tissue coll SBE Part V Molecular Genetce that eventually fimcion in food and water storage. Lastly, cells tthe core ofthe embryo are destined ws form the fa. spent of the plant bedly- lant hormones cx chapter 4) arc the tools that allow plant development to adjust to the M i ‘The form af plant body is largely determined by con- trolled changes in ell shape as they expand osmotically her they for Gee Bre 1786) lant grew tregulating hormones and ether factors inflcace the orientation of bundles of microtubules om the interior of the plasma ‘membrane. These micronubelcs sccm to guide clllone de postion asthe cell wal forms rsa the outs of new (ol The orientation of the elluone Bhery, in turn, deter sinc: haw the ecl wll elongatc a ic inercse in volume, and sadetermine the cell's final shape, developing plan, ears, flowers and branches a= sede vo the growing body in way hac are stony fnflacnced by heFIGURE 178 "The pa of plane development. An ilssracon ofthe dewlopmensal sages of Arabs lia (2) Fay el vision (@) Teeu frmaion (3 Seed formation, 2) Germination.) Mersamiicdevelopnent snd morphogeas Chapeer 17 Caller Machaninmscf Developmen 330‘The Flowering Plant Scheie me nly begioning to eave the melcenar Hil ‘ogy of plan development, langcly through inti recent sly of 2 omall wendy zeative of the mustard plan, the Sw ee dpe Heinen Bay geen te Coo wed with a shart generation time, Arabidopsis makes an ical ‘model for inenigating plane developmen. Ie is able to self-ferlic, lke Mendel's pea plants, making genetic analysis canvenicnt, Arabudapeir can he grown indoors in fetches, 0 Engl pla prodaing domes of cllgriog, after only twa months, Its gcname is approximatcly the js ey hee Ns acetic: Cereb epee ‘aod the fut fy Drphle melempater An der brary (Arabi gene clones ua made available to researchers fn 1987, and the fll genome sequence war completed 1999, bayo, the patter of Haoace lad dawn asthe embeyo Best Forms, Mutations in over 50 different gener that alter ‘puticm formation in rabidipeisconbryos arc now known, ‘fecing every tage of development While work in thi aca fs ll yery preliminary, it sppcars that the mechay ‘alms that exablish patterns in che early Arabidoes ert- tae broly dlc i: rs ci animal development. (Organ Formation nmpprecely she weber sderchapmicat of eng a Acabidopis sso scems to parallel organ development in ‘ssimals, and ssmllar et OF regulatory genes comtral de- velopment in reidepis Drape, nel rice. Arabidoe flowers, far camp, are modified levee formed as four whorl in specific onder, and homeotie mutations have been isnsfied shat convert one part f the pattern to ‘another, just as they de in the body scaments of 3 fly Gxsee 17.28), are only beginning co understand the coed teks eet nepeeree se dering ‘uml. The genes thar devrait pater formas nud organ development, for exam, operse inthe ‘sme way in plants and sna 386 Paik Moder GeneticChapeer 17 Caller Machaninmscf Developme 387