Anatomy and Morphology of the Vegetative Organs of Sorghum ... · Technical Bulletin No. 957 • JUlie 1948 .Anatomy and Morphology of the V'egetative . Organs of Sorghum Vulgare

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MICROCOPY RESOLUTION TEST CHART NATIONAL BUREAU OF STANDARDSmiddot1963middotA

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MICROCOPY RESOLUTION TEST CHART NATIONAL BUREAU Of STANDARDS-1963middotA

22

Technical Bulletin No 957 bull JUlie 1948

Anatomy and Morphology of the Vegetative Organs of Sorghum Vulgare 1

By ERNSI ARTtlCHWAOBII Blnior 1)la1l1 (watolllist DillillOIl of Sligar Plant hlllcsshyligations Bureau of PllInt Indllstry Soils and Agricultural Engineering Agricultural Research Administrationl

CONTENTS Pugc Page

Prcvious investigations________ _ 1 Anatomy of the stem _________ _ shy-Internode _________________ _ Matcril~ls and methodsbull______ _ 3 27Gross morphology ____________ _ Growlh ring_______________ _ 3 33Root blLlleL ________________ _ Morphology of thc sttlk _____ bull __ 8 38

Internode __________ _____ _ NodlLl plate____ _ ______ _ 8 38Nodc _______________ _____ _ bull 1 Anatomy of the le~f _ _ ____ _ 41Growth ring_____________ _ j 1 ShelLth base ___ ____ ___ _ 41Root band_______________ _ Leaf sheath_______ ______ _11 43Bud ____________________ _ 12 lelLf blade____ -- ________ _ 4418 Dewlap _________ bull _________ _MorphologyHli the IcaL ______ _ 48LcGObllLdel ________ bull ______ _ I) Anatomy of the root__________ _ 40 ~

Ieiltshllnt)_______________ _ 20 Large laterals______________ _ - 4) 21 Butttmiddotpss root Hbullbullbullbull __ bull ___ _ 52- Bl~joint---- ----------shy______ Sl II e ~ ~ _ 21 Small Inteml _ 52

= ~lcGwlap -- - - - 23 LiterntlIrc eiLeei __ 55

~ J ~ gt 0 PREVIOUS INVESTIGATIONS SO~hU1i~Svlglmm llljate PP[s) is a plltut of It1I(i(m lillrugc nmiddotll lt Istublislwcr as all pcollomie CTOP (Yl1I ill Uw lHdi(st dilizH tiOllS The litelLtllll dpuling wiLhits history gtographi( disltib1ltioll alld 3 botanieal d(scription of vlLri(middotti(s is (xtellsiv( It hHs (1((IIUy b(tll

Tcviewcd in (ttuii by VillHll Stlpll(ns nnd ~lllrtill (14)1 TlHir tleatise gives the uotnlIienl ci(seliption of muny sOIghlllll vnlittits an(l a key for stpnmtillg th(m

in til( dps(Iiption of sllgnrenllc varitties minut1 mOlphologicul and eV(It anntomical (hn[lwt(rs UI( us(d hilt in sorghum taxonomy rcmiddotiiancc hns bN1l plltlNI chidly on thl SlructulC of illnOnS(PII(P spikehmiddott and sllmiddotd Tow SUIP sOlghulll shows a Iplativ pnueity of such ehnlll(ttrs ns Conn til( basis for suglu(Il1middot (lnssificlttioll but somc of the (hnruettls nle of glNt distin(tll(Ss unci of I(ILI vuluc in the des(ription of variCti(s and ill the annlysis of a hyblid populatioll

Th(I( is no litelutul-( dluling sppcifillIlty with thc millut(middot mOIpholshyogy nnd anatomy of SOlghll1ll pxcppt for a papPI 011 thl dlv(lopn1lntlli allntomy allCl homologi(middots in til( young R(middot((lling (J 1) but its stluClllr( I Rubmitlecl for Jlublication Oct 11 1)4fi

2 Credit is duo )Ii~s ROH~ Mary Schipman for the prppl1f11lion of till drawings 3 Italic nlllllbCls in parentheses refer to Literature Cited p ~5

78605iO-48-1 1

2 TECHNICAL BULLETIN 957 U S DEPl OF AGRICULTURE

is indirectly known through papers on the anatomy of corn (5 6 12 13) and the morphology und unatomy of sugarcane (1 2 3 4 7) Importallt differences do exist however us shown in trble 1 in which some of the morphological and anatomical cln lacters of the two crops are contrasted

lABLE I-Some contrasting characters in the morphology and anatomy of sorghum and sugarcane

Character Sorghum (vulgare) Sugarcane

Morphological

Color of internodes _____ A shade of green_ - _____ _ Green purple ivory striped

Wax band_ ____________ Part of genr-ral bloom U~lIaJly in the form of a rarely distinct distinct band

Bud furrow ____________ Often poorly dcveloped~~ Usually promillentBuds_______________ _ Oftln small 1nd uude- Always well developed

elopeeHair groups on prophyi- Few and similar_________ ~sually numerous amI

lum varyingRows of root primordia_ Usually one row_________ Almost always scveral

rows Hairs on root band _____ Often present _________ _ Almost always wanting Hairs on internode______ Alwny wmting________ _ Presen in a few varieshy

ties Persistence of leaves ____ Stems never self-cleaning_ StPIIlS usually selfmiddotcleanshy

ing Hairs on lamina ________ Never present _________ _ gtrcsent in many wild canCl)

Hairs on midrib ___ ____ Prominent in many varic- Always wlllting ticgt

Hairs on sheath ________ Always wanting__ ~ _____ _ Prominent in mlny varicmiddot tim

Hairs on sheath base____ AlwaYR present though Ofhll Wlnting when prc~

short flllt IIsualh long (ilia on sheath nmrgin __ Never preHenL _______ - Often preent Typical auricles________ _ _do __ bullbullbullbull -- bullbull - Often vpry promin(gtnL Ligular pattern _________ USUl1lly shallow crescent Crcscentiforlll arcuate

or rlcHoid Iigular fringe __________ Sometimes taller than USllally much shorter

ligule thln ligule Dorsal hairs on ligule bullbull Dense free or only Ofhn sparse lne partly

lightly arinatl or cOlllplct(gtly adnatc Buttress roots _____ __ _ Often inscrted yery high Kcver insertltd high

(4 feet above) lIsually wllnti1lg Anatomical

Bundle of lamina __ ___ _ rnner scl(renehyma Inner lclerenchyml1 Sheath wanting shenth wemiddot1l d((~loped

Stomates of stem _____ bull lUlTlcrOlls__ bullbull _ ______ Scarce OftPIl wlntingAlways present________Silica cells of stem epi- _ Often wlnting

dermis Starch in internode____ _ Abundant in many ari- ranting or ery parse

eties Silicl1 knobs on root enshy fI~Rive and irregular___ Small and uniform

dodermis Wanting __________ bullbull _ Well developed Sclerotic laver outside

root enriodcrmis Exodcrmis of root bullbull - - Outlr tan~cntial wall Tnner tan~(ntial wall thickened thickened

3 ANATOlIY AND llORPHOLorjy OF SOHGHLM

In the p((~sCnt contlibutioll the morphology ond ouotomy of tiltmiddot sOlghum plant is tlCatCd in somC tCtail since a knowledge of ther stlllctUIC of the maturc I)rgallS is (If villue to the g(n(ticist and the pothologist An occount of the Qntogtny of the tissu(s is omitted fOl til( most part however becaVde it displays no essential deviation from that of corn and sugarcllte

MATERIAIS AND METHODS

The mateinl fol stlldy WIIS gJOWlI at the txIgtClin1(utul fnlm at State Colltge N )llx The eolltdion ioeludtd Ethiopinn nud Indian sorghums sweet sorgos and milos nitog(th(I 152 diffc(ent valitties

All characters Nere studied and illustrations prepared f(om frCsh mntelinl On the whole the technique Cmployed by the writ(l in the I1lOlphologicol studics of sugllleunC vltlidi(s (3 4) was 11sed Sl1( shy

cCssfully in the prCsCnt illYtstigatiolls As in sugl11enllC some chamemiddot telS show to b(st ndvnntng( in young nmt(Iiitl sinN I(t(oglpssiyc chnnges or m(re injulips nssoeintec ith nging or due to disluses make Cvaluution difli(uit ~lost chnrn(tpIs OIP studied more Mlshyvantngeously in older morC mntuJe mawlilll hO(Vl1 Root band puhls(Cnce is flcqtlCntly limihd to the basal pflrt of the stnlk whCI(ns typical wax bnnds may bc found only in thc lIPpCI intlll1odes sillec suhsequCut all-o(( wux dlposits tIHS( til( boundulY b(tw((1l gtllPIHL bloom und wux bund Til( shmiddotm (pid(Imis is not so Cusily stlipppd as in SUgUT(lll1C and in ((ltain Ynlidi(middots surfnt( hund sletions mllst be emploYlC1 (Inn though thpy all not PIY sntisfndolY Ihl usul11 tlclmique of fixillg and stailling WIIS lIs(middot1I in till uniltomitHI studiNi hut was often abundOlltd ill fllOI of hand s((tiolls of fnsh mnPlinl [hin hand s(ptiol)s (sIwcinlly of hurd tiSSU(s 111( (qunl to and oftpn suplliol to mielOtoIllC PI(pl1rntions If PIoIwdy stnilled th(y show l)(tt(I tissuC difl(lI(ntiation in photogIll phs thun (ould bC obtuil1ld otIHIwisl 1lt01 that llIlS0n mnny of the illustlHtions in this bulletin ClC prepnlcd by this method

GROSS iIOHPJlOLOGY

Sorghum a 111(mh(I of til( tIih( Andlopogon(u( fomily Glnmin(u( is a IHlhnNous Hlll1Unl Ih(middot joillUmiddotd (ylindliclll s(IllS 01 (ulms YHly in h(igitt flom 2 to 15 f((t Ih(y (11( solid but (h( ((lIlml plll-t OftPll 1)((0I11(S pithy und fistulnl Both stlchls itnd sidt blllll(hshy(s nr( PlOdll(pd (fig 1) ]hc 11111(1(15 IIPIH111 only IIft(1 the maill sl(middotm hns 1HlIdpd ou t ant tiwil 01lt1(1 of ItPPNIIIlJ1(( is hnsi pptlll with lll( llppl1 bud producing th( fin bllI Illth TIl( sid( llIlwll(s 111( smnll(l than tilt muin stnlk lind thpy llWtlll( I1llwh 1111(1 Thp IlumblI of su(kpls 01 tillcIs d((wncls on elilllllt( splicing IIlld YIIligtnl tClHIfllcy in themiddot dtnlopnwll t of (IOWlI buds (Li J

Ill( 1(IIYI5 Ii 1( two-Iunk(d liS in OtlHI gliI5S(S ultpll1llting on oppositC sid(s nnd upp(l1ling npproximnlcIT ill n pitUll rIH 1l1lmblI of IlnYNi at mutlllitv Yl1li(s with til( vUli(I flom 7 to 28 En(1t ICnf (onsists of shllith nne bin dC Ill( hlldp is lilwnl Inne-polnt long nnd lint It is SPIHIli1tNI flom th(middot sh(lIth by thl SI)((iuI iZllL l(gions of tll( hlndp joint- middotltllwlu pS lind ligulP 111(1 Iii t ItI is It IIlembllI111WPOIIS npIWntlilg( thul doslly in t8ti th( (ulm lind tlds liS It

4 lECHNICAL BULLErrr 95 U S DEPl Oli AGRICULTURE

Fwnu L--Hubit k(gtt(h of sparH(-i(ufN s()r~hUlIl wjlh i(( hrUTleilts and lI1all Huckcr hoot Internod(l 10111([ than ~lIht(ndinj tNlthi

5 ANATOMY AND MORPHOLOGY OF SORGHUM

rain guard The sheath SUlTOUllds the inttmode as an open tube

r with overlapping margins At tlH base of tlH sheath where the l(af joins tIl( (ulm i a distinct sw(lling tht lenf sh(nth bns( 01 shenth node

The root system is fibrous composed of mnny sl(nder roots of about equnl (lianl(tcr Thetmiddotc is a single seminrti root that gows vertielllly downward giving off laterals throughout its length It may function during thC Cntilc Iif( of the plant but ((nslS to bc of

t

I r I

c

FTGCRB 2-A HOOl crown of 50rghulII plant Wilh thrce whorl of aclcntilious roots in urious stagcs of dcwloplllcnt fl Bu~c of young stalk with uclcnshytitious roots in cariy stagc of de(middotlopment Part of sheath in UppN iuternodc cut away to expose growlh ring and root band wHh rooL primordia C Basal intcrnodc with adjacent nodal regions Uoot band has two to three rows of root primordia bud furrow well denlopcd

imporlnn(C aft(r the plrn1illwnt root Systclll h(gins to fmktioll TIll pelmmwut roots nl( ficivcmiddotntitious nnd dnlop in sU(((ssion from thl bnsl11 nodeS nenr ground Ivel (fig 2 lt1 nnd lJ) Thl whods of ndnntitious roots in their CntirCtr (onstitute the root (rown Allla(llIls Csp((inlly thos 11(111 tilt sllrfnecmiddot of til( ground al( mllch brnnellPd illhIlneing thc soil in nil di)(ldions ~r1(il (0101 is white Ol~ rNldish brown dp(nding on agl lind stngC of dC(lopnwnt ~lfallY sorghum vnriCtils cllYllop buttlCss roots (fig a) flS high ns 4 f(Ct ahoY( ground ThC hultlCss roots 11)( (onsid(Inbly thiekcmiddotr thnn thC nOlmnl roots flnd usunlly fl dp(p gJllIl On tllt(ring th( soil thCy l)(illw(I ns ordinmT roots nncl dC(lrnsein sizC to tilc dilllTICtr of tilCmain lut(rnls (fig ~)

6 TECHNICAL BULLElIN 957 U S DEPT OF AGRiCULTURE

FlUnn 3~middot middotBllsal part of lt11111 of ILII Ethiopian lorghllIl1 ilh prolllirl(~lIt iJu(tns5 rOOl

Ihl suptrtilinl root s)sttll1 of sorghlllll J n(eonling to VPIIY(r(1 PIJ 192-195) is ext(nsivl Ih( roots nUllin a working dtmiddotpth of 3 to 4 fctt and u maximum d(pth of 6 f(middot(t ~Iill(gtr (8) sllltts thnt til( 100ts Ilrt mort fibrous than those of (Olll lind mil) form twi(( as mlmy Intlrnh itt I1ny slllgl of tlllir dlv()opmrnt milking till sorghum plttllt 111111k((11y drought-nmiddotsistnnl

The inflorest(rtet or IHi~dJ is with ftw ex(~(gtpliolls (brooIlHorn) n sOIl1twhnt cOlllpiwl or open pallid( Till 1111lt1 is USllnlly ertet (fig I)

ANATOMY AND MORPHOLOGY OF SORGHUM

but in certain groups it is inclined or pendent (fig 4) The main axis

r is angulor vmiously furmwed and usually hairy at the nodes The side bri1ncheg appear to be in whorls one above the other Each lateral may bmnch repeatedly The final branches bear one or several paired spikelets which are ellipitcal of varying length and diameter and usually compressed dorsiventrally In the case of paired spikelets one is sessile and perfect the other pediceled and staminate The fertile spikelet has two thick glumes of about equal length but the outer partialllY envelops the inner which is slightly narrower The glumes encloBe two flOWCIS the lower sterile the upper ~erfect The lOWe sterileloreurolt has a narrow lemma eleCted at the apex from which an awn arises The polea may be absent j when present it is small and

Cwtln middot1-middotmiddotHllbit skllh of sorghum nricty with 1XIHlcnt or in(rtld pllniclc

8 TEOHNICAL BULLETIN 957 U S DEPT OF AGRIOULTURE

thin There are two broad hairy lodicules three stamens fllut two plumose stigmas

The mature gmin is entirdy or pl[middottJy endosed by the glumes It is ovnl a little longel tlmn brond smooth awl tipped with the Imains of the style One sUIftLce is nlways more Ot less IlnttelHd nnd clUIies till eml)lYo at its base but in fctelita the flllHened sll1face and emblTo ale on ditTelcnt sidcs

MORPHOLOGY Ol~ THE STALK

INTEnNol)lo~

The htight of til( sbLlk 01 (ulm is dd(Imin(d bv the size nne llllrnbpr of illlcll1oltips composing it (titble 2) Ill( douhllt dWillf lllilos hlnt 1(I~lively fpw and sholt intelllocfs (fig 5 B) whilp in the tall EthiopilUl fOlms (fig 5 A) ILS mnny as 25 inlellloltis lllay dil1cleniintc beforc thc ndnnL of Iloweling

rile intelllotl(s llre iOligest nndllost unifollll in the middle of the stnlk imd shortest nt the base but lhe telminnl inl(lIlode thnt bellIS til( illflolPS(P1Ue is lhl long-pst of nil (tnblp ) The fOlln is cylindli( bobbin-shnpld 01 cOlloidid j lhl tIoss-sclionnl ann lOlllld 01 oval Th(gt bubbin-slmpNI int(lllodls ILljl I1sllally nssoeilLltmiddotd wHh swoll(IJI nod (5 jL fll1lulp of mnny thin-stpmmed Ynrictjes In the (onoililll jllllIlolie Ul(lc is n sharp constlielion below the shlnth bfiS~~ ilnd n glluillid inel(jlse in dil~n1ll(lr bllSipftally Il is n COlllmon type Muong thitk-stll11l11ld YIlrieti(s with swolll1l Ilod(s

Til the milo sllfLins (10) theinl(Illolaquo((s 1)((0111(gt PloglPssi (Iy longpr from til( glolnd up 111(1 unimodal forms hlLV(i On( sholl ililclIlodc bplWNn olhlIs lhtll ILIP 10ngcI above lId Iwlo while in til( IlLt (l vlLriptils thlIl is II dougt( fnlllIHllise in int(llllotie lpngth ending with II fi nal Iisl towlLrd til( pPliuncle

Thl surfllfl of lh( intlllllode hns n 1llllvy tonling of wnx tlmt oftl1I compldlly masks th(i grppn St~lll (oil) In thp young intlmodes of mltny vIIIietils nnd in lhl olc(I 1I0iks of n f(w thl WIIX dplOFlit is (OIl(llItlntNl bpi ow the shlnlh bast flWlllillg wax bllnds like those found in llllllly sugnnlle nllielils

On til( side nlxt the leuf (1)( illtlrnocc is ftllTowld tlw position of til fUllow nltll1Ltinp lit ln(h no( Ih( flllTOW is II IIIlTOW dl(((ssioll in th ill ((lIlode lxtlnd i liP Lolll tll( bud II pwurd (fig 2 f) In son( Yl1ridips lhl hud furrow is nllogrthtI wlnling ill otll(rs it (x(11l1s only Plllt wny lip till intllI1odp Ofl(n hut fUITOWS I1rc oigts((Cd only in lmnL iul(Inoc(s hut Illily nppllU sposhyIIldicully in otiHIS 1~1l1s( hud fwlows IU(I d((H(ssiOrts in the sLlllk not in line with lhl bud [Iwil d((loPIIHllt is Jllobably lellLted lo P(St1C dill((II(gt(S dUling ndin glO-th

Th( textulp of lIll fllsil is lilln lind jlliey ot dlY lind pithy In vltriCties with tit-Ill f1(h II fl(middotshly (lit sgt(tion is glllyish 01 Qli(

gllPIl Tn 30nH Yftli(lifS th(l ((llIttI lwcolllCs whill nlld pithy lind ill HIlilY Elliopilln nnd sonH Indilln Vlllillips till (Illil( (OSS s(dioll CXC(lpt the nl1(lo ouler cortex tnkes 011 It eollony-whit( npPNIIUII(l


r

~~~

~ lt7 ~ I~ tti~~~~~r

HB J~l Ifr

FWUltB i5middot~~A Habit sketch of YOllng IIcticy growing Ethiopiun sorghUIII lind 13 lIlature double dWllrf milo

lABLE 2-middotRelatite length in centimeters oj internodes and tmbtendin1 sheaths oj some sorghum varietus 1 ~

- ---- 1f N~~~2Tr~~ 640 2 1pound N 634l~lI N (i35 ~ -~IiI0 ---~~gar Drip M N 472 2 D~~rfb~iilo ~ Nurnheror illter- 1_-----1_ -_~~-I ___ __ _______ ~ nodes nlld sheaths I I I I Z

j Int(r- Shenthllnter- Shent11 IIIter-jShetlth flltr- Sheath Inter- Sheath Inter- Shenth Inter- Sheath Inter- Sheath I~I l~ ~ ~ ~ ~ ~ ~ 0gt-I --- - + --1-- -- -1- ---~-~ --_-- _ ---- ~- ---

i Cg Ci ~~r~ ~~ ~~~~ ~~ ~ ~~ ~~ ~~ ~~ ~~ ~~ ~ ~ 1~==============t ~ i l~ ----rj---if ====1==== ==== ===== ====== ===== === ====== ====== ===== ====== ====== = 11 ________________1 g -- I _-1 1------1_-------- ------ ------ ----_ ------ ------ ----- ------ ------ ------ c1 1tbull 13 I 1 1 t u- I I 3 31 I 27 ---------------- I - -) I Iv ------1 ----- ------ ------ ------ ------ ------ ~Ji________________ 12 1 21 l 3 2(i 0 24 0 I ) 1 17 18 340 bull ~ti ------ ------ ------ ---- shy14________________1 13 23 I 15 26 1 24 I 20 I 13 ]7 6 2(i 318 30 ______ ______ l 13 ________________ bull 15 23 22 1 25 2 I 23 I 23 I 18 16 9 22 11 18 ------ ------ Jl12 ________________ 1(i 22 9 26 13 t J7 4 24 I 20 18 13 22 10 ]6 ______ ______ bulll

J 1________________1 17 23 16 27 231 28 10 26 18 20 16 23 14 16 -_____ ______ t1 10________________1 191 22 20 2(i f 23 I 26 I 21 27 21 20 16 24 18 17 ______ ______ ~ H_________________ 21 22 15 2(i 22 21 21 30 20 19 12 23 20 18 328 28 Y 8_______________bull 23 23 18 281 23 28

1

22 30 19 19 15 22 19 19 5 17 7 ________________1 23 24 12 25 23 27 24 30 20 19 17 23 22 19 3 14 ~

(-----------------1middot 26 27 30 29 20 23 19 a23 14 18 28 19 15 22 135_________________ 24 26 14 27 18 30 ~o 27 18 18 13 23 23 20 2 13 gt 4________________ 25 26 t 13 24middot 14 30 28 27 16 18 )2 23 19 20 2 14 ~ L ____________ 24 25 7 24 11 27 21 l7 10 19 12 24 17 19 2 13 h 2 _ bull ___ 15 24 middot3middot 17 7 26 12 24 5 18 7 19 13 17 2 13 C1 1 bn~~ of sUllk)___ 4 24 I lt ~15 4 26 6 24 4 18 3 15 8 17 1 13 t Y

Totu________ 303 - 4731 197 439 --02 4121- 229 373 I 238 277~312m-235-W------UO sectlAvernge ______ 138 21 5110 lt1 I 23 1 125middot 257 153 24 l l 159 185 1191 223 166 181 25 131 tI

I 1 I

1M N=hl~i(iiII(hl) numher IlnllOrt~nce axis l1ullncluded In nlrnging t~ngth nllnternodes Still tlcticly growing Sheath suhlcnding lufiureStllntll llxi~ [wt included Innerogng iongth orshcaths

fill

r

ANAfOMY AND MORPHOLOGY OF SORGHUM 11

There is a slight correlation between color and texture of the flesh and stalch and sugar content of the variety as seen in table 3

T AIJLE 3-Relationship of color and texture offlesh to starch and sucrose content of some Ethiopian selections

I Starch 1

Color and texture of cross section of 1______1 Sucrose 2Variety Xo internode f racket Diffuse I

-----------------------------middotI------~------I------

Percent M N 620 ____ Gray green soli(L_______________ + + 107911 N 675 _________ do_________________ _______ +shy +-- 45811 N 688 ________ do_________________________ ++ 68411 N 698 ____ bull ____ do_______________________ _ +

-Ishy 482 M N 61L ___ Gray green with small white +- 834

center ~1 N 623 _________do_________________________ +shy + - 61711 N 624 _________ do__ ___ _ _ _ _ __ __ ____ __ ___ _ _ _ + + - 1098M N 672 _________ do_________________________ +shy 3 0711 N 673 ___ _____ do_________________________ + +- 407 M N 079 ____ ~t of 8tern cross section white_____ + +-- 30011 N 694 __ _______ do_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ + -M i26 ____ of S(1ll cross section whitc_____ -f + +

475 584

M N 619____ Flesh white spongy ami fislular_ +- +-- 7 71

I Drsilffintion of nmount of stnrch +=Ctlls pllrtly I1I1NI with slnrch ++=cllIs 1lI1ll1 with stllrch +-= starch scaret +--=itnrtll lry StfUCl- -=no stnrrh

t Fi~urrs Or$ucros contlll tnken from A ll SchlchutJlrs 1944 unpublished sorgo report In Illes oC DImiddot ision oC Sugnr Plnnt Investigntions

NODE

The node is citlwr flush with the int(rnode 01 somewhat thickened Ontogtndienlly it is limit(d to the r(gion just belo the inseltion of the leaf hut taxonomitally it includes growth ring and root bUIld

GltOWTfI IIING

The growth ring is a partially differentiated region that retains its cell growth potentialities Extprna11y it appears IlS a IlIlITOW band flush with the internode or protruding from it In lodged stalks one side of the growth rinamp is (Illite brond since it is throlJOh the acshyUdty of the cells of this region t lItt the stalk strllight(lls itself from the b(mt position lhpf( are IlO conspiellous Yllrietnl diffcIcnces as reshygluds height llnd conlOUl of growth Iing it has no taxonomic value

HOOT IIANll

The root band isintelmiddotpolated between sheath bllse and growth ring It contllins besides the bud one 01 severnl rows of root pIimollia The height of the rrtowth rillg varies from 4 to 15 mm Its colol is lighter than thllt of the internode above There is no visible coating of wax but the basal root bands of many Ethiopilln sorghums Ilre distinctly pubescent The hairs are usually limited to the J(middotgion below the root plimordia along the entire circlllnfelCI1(Cc of the Oot band or along certain sectors of it O((l1sio11n11y the hairs Ilre (onfined

12 TECHNICAL BULLETIN ) 5 j U S DIoJPl Ol AdlUCULUHE

to the zona flanking the bud In some fOlfns the Nllim root bunel including tIta root primordia is covalcd with hail

The root bands arc usunlly cylind deal or tUll1cs(cn t-Obcolloidal bull Tha root primordia ala disposed in a single cOl)eentJ-ie ring but in bull many Ethiopian sOl]hums the lowet Dodes contain amp0 occasiollally three rings (fi~ 2 U)

rIta root p1lInordia of the basal intolnod(gts ato 1mgo and swollPII and usually glOW out into buttnss loOts whieh upon (gtntelil1g the soil establish a film SUppOIt for tho stem In SOI11( vlujetips tho buttl()sS roots develop at a considelahle lllicrh t abop Ioulld lj 3) but they coaso glowing nftel lttt11ining 11 l(~lgth of Sl~Vllll inches ~nd novel ent~r the soil If the root l)11l1ds IUC VltIY nllIIOW lht root primordill nro disposed centrnllYI but in (nil root bands thcy 1)lll close to the glOwth ring Root plimoldin may fnil lo dovelop in tho upper part of the stem at best they nre inconspieuous

lIun

Ev(ry node is nOlml1lly geml11iflrOIlS hCI1Iing n singlo bud In some vurictifs the best d(lvelopld 1)1lds Ille Iollnd in tho bllsnl int(fshynoells becoming progl(ssiyCy s111nll(I nnd npPl(ss(lc highpl up the culm Other IOII11S eM N (90) hnp n w(l-dlVloplIl bud on olwh node except the npicnl OIHS Still othols IUO chnIncl(Iiz((1 by having only one or two won-developed buds on nnv givln stalk In mnturo stalks in a dense plnnting the bnsnJ culm shonths 11111 withel nne the buds of the nodos thus llXpos(]1 oftnn nlso dry up lhis )rhaviol is unliko that OhSClVpd in stlgarenno in whieh the culms nm normnlly sulf-denning and exposll( docs not aneet the life of the buds

1he buds at glOulld loyel mny deyolop ns SlIek(ls while the u ppel- IllOst buds orbll clongnte to f011ll side bmllehes Both SlIck(lIs and sido branches develop hends thnt IllntllO much Intel thnntho I11nin hond

A bud primordiulll is initintcd nt the lInso of (neb inhnlOltic V(lI oarly in ontogfllY ItS dlll10nstmtod hy ShuIIllIUl (12) 101 t1w (Oln plallt As the int(IInode eongntes tlte bud retnins tlte Iwslt positioll It is thus nlwnys Ilssocinlod with the kaf above and 1I0t with tho leaf in whose axil it oceUlS

The buds 111( insCrtNI ill the tissue of th( root hllnd directly ahove t1w lonf S(IlI 01 SOIl1( distance above SOllwtimcs us high as tho middJe of the root halld The height of insOltion shows V111i ali on within (lIlm limits hut on the wholo it is n fNltlllO ehnladfIisliie of the vnJiely

The outer covpring of thl budfi is a plOphyllutn In its pntil()ly the pophylllllll of sOIghulI1 Iiko thnt of lHIgIlI(nIHl (SjirJ 6) forms 11

hood with th(~ fron t side (0111 pOSN of 1wo nsymmptlienl ovollnppillg hnlvcs and the bllek sifh (lnti( Jj flip bud deyolops into a side b1l)neh the plOph~rllUI11 (hngnlos ILnt splil Itnd the two sides form tho fist lett(s of the ))1Iwlh

Disl(gnrc1ing the nllll1(IOIlS nh(IIllut fOllns (fig G) two geneml typps of buds ILre distinguishnhlc-thc sorghum find thp sugnrelllw

The sorghum type chnlncUIistic of the sorgos nnd milos sotnewhnt 1(S(1l1hles an fLlIowtend in (IJid etw 10w(1I part hns 1 plonoullcNI cushionlikc ext(lnsion that Ipnebcs flom tltp bust of lhl plophyllul1l to the lln( selll (fig 7 11) The huds arc wdl dovlloped Inlge 01

medium ltugo nne symmetricnl

13 ANNrOMY AND lIOUPHOLOGY OF SORGHUM

r

FlOlU1~ G-A Prophylltllll of bud of illelisi illet cltLwfish (lnttltrll-right secondalY willg VCIY 1~1g( root bllnel Imiry H niJelrlLllt type with prominentsecondary wiugs

FlOenE 7-1 HOlghulI1 type of bud-obcolloidtLl root Imllltl with oue row of

root primordin IJ sugarcallc typc of bud with crossing-ocr of sides ill ncnr-ccntrnl position

14 TECHNICAL BULLETIN 95 U S DEPT OF AGRICULTURE

The anterior or fOnt side of the prophyllum is elongate-deltoid wing insertion basal with or without prominent auricular set-oft The two overlapping halves of the anterior side are symmetrical _ although frequently the overlying half is larger and may covei the entire front side The l1sal appendage of the ovellying margin is

FWGRE 8-A Jntermediate sorghum type of bud with crossing-over position neaf central X 6 B Sugarcane type of bud with large overl~ing side crossingshyover position apical X 10 C Typicl11 sorghum type of bud with hair groups 12 und 4 well developed X fl D Bud with large overlying side und one promishynent secondary wing X 6

15 ANATOMY AND MORPHOLOGy OF SORGHuM

never conspjouous as in sugarcane and is often wanting The two halves show frequent crossing-over i e the overlying half becomes the underlying half This interlocking of the two halves may occurmiddotr near the middle (fig 8 A) the upper part (fig 8 B) or the base of the prophyll (fig 9 B 0)

~~I~~L ItI)ft 1 rlilA

I ~J l)~

amp ~~~~~~ ~~~~f I~ ~ I bullI~ ~~ I t rl-~

i Ir

~~1 i~i ~~(( r IfJ~ If gtD -tlJl 1

1

FIGURE 9-A Prophyllum with small secondary wings X 3 B Prophyllum showing central crossing-over of overlying side X 6 C Overlying side covering entire basal part of bud X 12 D Prophyllum with two prominent secondarywings X 6

16 TECHNICAL BULLETIN 957 U S DEPT OF AGIUCULTURE

Groups on sides

Groups on wing

middotmiddot16

middotmiddot26

FJOR~ 1O-Dingrnmmntic drawing of prophyllull of ugnrcllnc nplicntcd thnc timc8 showillg locution of hnir groupil (A) on ici(s (13) 011 wing und (G on JUHcturc bctwccn sides nlld willg (Legelld cOIIUllucd 011 fllcing puge)

17


Hair groups on aTlerior ilido (fig 10 A)

(1) Lateral groups on overlapping halves of prophyllum Hairs are white long and coyer the baso of the bud to a greater or lesser extent

(2) Basal groups on overlapping halves of prophyllum These strips of short hairs are often intenpcrsed with groups of longer hairs alternating with the veins alld often covcring the short hairs

(5) Groups of straight or wavy vcry apprcssed white hairs on one or both sidcs of the gcrm pore and usually associatcd with long buds

(6) Bands of short brown hairs betwccn the veins (17) Groups of long oftcn wavy apprcssed white hairs bctccn the vcins

giYing the side a silky appearance (24) Short lashcs along the upper half of the mcmbraneous margin (28) Incurvcd and downward-pointing hairs immediately above the gcrm

pore on thc posterior sidc of thc prophyllum (30) Short inward-pointing lashcs on edge of germ pore (31) A group of long white dowllward-poiilting hairs imphwtcd at the point

of insertion of basal appendage (32) Short brown hairs on the surface of basal appendage

Hair groups on posterior side (fig 10 tt)

(10) Group of lone llairs implantcd on the tip of thc side betwccn the vein Rnd possibly extending into rcgion of wing Hairs mlv or may not protrudeabove the bud

(18) Strip of long white hair~ along basal cdgc Thc hairs may be long or vry short rcstrictd to the ccntr or distributed ovcr cntire baal curvc The hairs may form a compact row or sraall groups

(20) Short brown hairs bctwecn thc veins (23) Short brown hairs on tip often maskcd by hairs of group to (25) Long whitc hairs IWLween the CillS

Hair groups on anterior wing (fig 10 B)

(3) Iong white hairs on basc of win (4) Lon whitc oblique or appresscd lashcs on edge of wing limitcd to basal

rcgion or occurring oer entire edge (12) Small brownish hairs on surfacc of wing (13) Iollg appresscd hairs On surface of wing (15) Lashcs at tip of willg may be considered a part of group 4

Hair groups on posterior wing (fig 10 B)

(14) Tong appresscd white hairs on surface of wing rarely found in Saccharum officinanun

(21) Short brown hairs on surfac of wing (29) Long white hairs on base of wing Groups 4 and 15 shared with anshy

tcrior of wing II-zir group~ on aTlterior jllltctllrc (fig 10 C)

(7) Groups of shori brown or long white hairs above celltral plgtrm pore (8) Long hairs appnsscd ill regard to the will or projecting abovc thc wing (I) Long wavy white apprcssed or protruding hairs on juncture of wing

just ben(ath the willg tip (Hi) Groups of long white hairs implantcd on a broad base in the basal dcprcsshy

sion bctwcen sides and wing (26) Long lashes in corncrs of thc wing

Hair groups on posterior jl1Iclllre (fig 10 (J)

(19) Groups of long whitc and appressed hairs inserted in the basal cornrs and sometilll(s prntrudin abovc wing

(22) A narrow balld of appressed long hairs often forming a connection heshytwecn grouP$ 19 and 10 Group 2li is sharcd ith antcrior jUllcture

18 TECHNICAL BULLElIN 95i U S DEPT OF AGRIGULlJRE

Tile pubescence of t110 prophyllum is promiIHnt or inconspicuous There is never the wealth of individual hair groups found in sugarcane and for that reason prophyll pub(scence will I1(ver be an impmtant character in sorghum taxonon1Y The anterior side of the prophyllum has hair groups that are roughly equivalent to groups 1 2 4 12 and 26 in sugarcane (fig 10)4 Groups 4 and 12 arc usually present (fig 8 G) while the others may be wanting Frequently groups 1 and 2 are faidy massive and often constitute a continul1tioll of til( hairs of the root bane IIll anterior side of the prophyllum shows an all-over pubescence of wing and sides No distinct hnr groups an recognizable

The basal bud cushion that characteriz(s tho sorghum typl of bud is most prominent in buds that all inserted high or fairly high in the tissue of til( root bond This cushionlike swelling (fig 7 4) is always paler than the adjll(middottnt root-bnnd tissue The swelling is thickest near the insprtlon of the pr-ophyll wlHnce it gradually tapers ill the dir(ction of the Iraf scar If the bud is inserted nry low the cushion is in(onspieuous but seldom aetunlly wanting

The sugarcane type of bud (fig 7 B) is found in many Ethiopian varhmiddotlies In its typical form it is similar to that of sugarcane except for the paucity of hair groups The prophyllum is usually deltoid with wing inslrtion lHar-e(ntral or sonwwhat lower The overshylapping half of the two sid(middots is always til( IIIImiddotger Tll(re is frequently a w(II-deyeloped apical appendagpound The interlocking of the sideS already dlscribCd for th( sorghum type of bud is n velOY (ommon feature Ivln in prophylls whpre the overlapping half is vpry large (fig 9 0) The puiws((nee is parse

A feature that is conspieuous in lh(~ sllgar(anc typl of bud but not entirely absent from the sorghum type is tht formlllion of s((ondlll-Y wings that are outgrowths from the flanges of Lhcpormnl wing (fig 8 B D and fig 9 A D) As a rule their location IS basnl but thC)T mny deyelop along any part of the wing Oftpn one s(condnry wing is suppressed OI undeIdevploptd (fig 8 D) gimiddoting the bud an asymshymetrical nppeamnce In plump buds with high wing instrtion the addition of secondary wings may prodtl(( the cmwfish typemiddot (vtlriety M i 60)) h ieh is (specinlly conspieuous whnevr the basal cushion (fig 7 A) that (OIllltctS the bust of the prophylluHl to the knf sear is very promincmiddotnt Plump buds with promin(nt basal Illlli(ulnr set-offs of the wing but lacmiddotking slcoIHinry wingsilluy show a distill(t dover-Ieaf paltelll This type is usually restricted to basal internodes with swolleu root primordia

MORPHOI~OGY OF TilE LEAF

T111 leaf of sorghum liS in all gmss(s (onsists of two purts-the sheu th and the blnde A t the j tHHtll of sllPn th and bllld is tht blnde joint with its spNmiddotinliztd Ilgions the ligule und tilt dcwlaps

1ht lPuves vnry in numbcr frolll 7 to 25 Ilnd un IUTllngNI nlttlIlal(ly middotwith n divrgpn( of 180deg Endy vllri(tils hux 7 to 9 ItIH(S aboo th l (ron while lull vuri(llips may proltiu(p us muny ns 2$ Yaridi(S that a vpmgc 10 01 feWlI ]Paves a bovc the (TOWn are dassed IlS sparsely

bull Fig 10 i fig 4 of (8) lind is T(prOdUlNJ h(rmiddot to uid the fluder

19

r

ANATOlIY AND llORPHOLOGY OF SOHOHUlI

leaved those with 11 to 13 as midleafy and those with 14 01 more as leafy (14)

LEAF BLADE

The blades of young leaves are stiff and erect on maturing they spIend funlike in a gentle curve

There is little ynriation in the length uf the hlade among difftlent varieties The majority of sorgos have medium-long blades (100 Clll) wherens the milos have short blndes (60 em) The width of the blnde vtries between 5 and 13 cm )[ost blndes arc linear lanceoshylnte widest neal the center or uppltr central ZOlle In certaill varieshytiCs the blnde is Inllceoillte widest neilL the bllse and tapeling gradushyully in the dirCetion of the apex )Iatum leayefJ am smooth-middotedged 01 SCITlltC hu t the margins of young lellvcs am ulwuys scabrous

The color of thC blade is medium 01 dark green and the sUlface is shinr The busul pllrt of the upper midrib stlface is often conspicushyously fllrinose The powltiCry wax deposit that chnlu(middottCrizes this condition forms a broad zone just aboye the hlnde joint narrowing acropetally und fading out some 20 em highel up

The hlude is dhrided by the midrib into symIlletrical or EloIllCtin1(S slightly usymmetricnl hnlns TIl( spacing of the principal Y(ins is similar for most varilties Cx(ept thn t in the brond-ltnyed forms the veins nlt set fnrthtr npnrt thltn in plants with IHUTOW(r bladCs someshytimcs nnITOW nncl m(diuIll-wide leaycs have theil veins (xtra far apart

rhe midrib of thl I(nycmiddots of most IlImiddotiemiddotties is very promirwnt It is nlwllYs ery wide Ilt th( bnsC gradul1lly tup([ing townrd thC npex and disnpplnring nltogelhCr blfore renching thC tip of the lumina The upper sUlfllce is chnnn(led the lowel connx A conspi(IIQlIS chnlshynrtpr is the difJlrence in color of the two sides of the midrib While the lower slIrfnte is nlwnys green except for n light stripe thnt (uns nlong the center the upper stll-fnce muy be vuriously colored dependshying on the Yilriety and the stnte of maturity of the orgnn The npppr midlib surfllee of all young lellns is cloudy 01 light Qlin in cololmiddot As the leaf nges the surfllce tUIl1S completely white or rClllllillS (loudy except for a lIillTOW or medium-wide (eJltml zone thnt is either pure white 01 somewhllt pllllr thlln the flanges The milos hnye a y(lIow or olin-co(orld midrib caused by a yellow pigment in the suiJCpishydlIIll111 pllr(ncbyma cells

Yari(ti(s IlIlly be dnssifiNI as haying whitl cloudy or yellow midshyribs (14) 1he cil)ssifkn tion wh i t(-ribbed is nppliNI only to those Yllli(ties in wbi(h the entirC midrib of mature 01 huH-grown l(1Y(s is pure white (fig ] 1 A-D) Yaritti(s with a ctntrul white streak nnd wi th the flunges 01 margins cloudy (fig 11 E F) are classified as doudy (14)

The midrib of the IcnY(s of many YnrietiCs is sCllbrous espccil1lly in the lowel part nnd mlly enn fpel hairy to tbe tOIl(h In some forllls the puhescncc extellds upwfll(l for only lor 2 em in othcr Yfuilties two-tbirds of the midrihis hairy rhe hairs fire nlwflYs dens(st find longest in the hnsnl Plllt of the midrib Her they covel the (~Jltire width of th( midrih whil( highltl up they thin out und I)((orne loculized along the fiunges In many forms the midrib pubescenee is dense only in the very busal zorw with some sefitlCred

20 TECHNICAL BULLETIN 95i U S DEPT OF AGRiCULTURE

FWlJIU ll-~ridrib~ with adjaCll1t middotblnde tiRSIIC A-f) White-rihbed lLricti(l R midrib wHh brQll(i white c(ntJr llml cloudy flan~(~ f lIIidrib cloudy All lt l~

hni( ioeniiz(d medinlly 0( intllully Ilu(hing lip ns higb ~IS 10 (m ~rany yulitmiddottjmiddots including the milos hl1Y( 11 smooth rnidlih

Midlib pu)(s((1(e is nn important eitnluet(l lIs(rlli rOtmiddot group classifi(ation Ilnd impoltllnt fol pUlposes of identiJielltioll d)Nl lIs(d in (onnection with other ehtulIet(ls

L~F SIIETII

11( shcnth is Ilttnched to th nodI nlld (nCilels til(l uim It fOlms 1111 OPtlI tuhe extending Ilbout Ollt Ilncl n thild tinHs IIlound the bllse of tlw internode The out(l rilillgin of th(~ shNlth OY(Jinps (hp

21

r


inner the overlapping margins being altell1ately the right and the left in successive nodes The sheath is thickest medially grad ually becoming thinner toward the malgin whele it is membranaceous The outer surface is glabrous the inner surface white and glistening

The length of the sheath valies comparativcly little avemging ahout 25 cm for most varieties Howevee there ale valicties (~L N 628 and 694) with rather long sheaths attaining a length of 33 ern or relatively ShOlt ones C~L N 448 462 and 463) with a length of only 16 cm Within culm limits the lengthis faidy constant (table 2) except that the bnsal and apical sheaths are shOlter and the lowe I

near-centml ones longer than the Itvemge Thele is often n certain deglee of overlap because of valintion in lenoth of internode In many of the medium-tall splllsely leilved sorgluuns the sheaths nrc shorter than the intelllodes they envelop so that the nodal region of tile next highee leaf is plainly visible (fig 12) In othel fonns the neUl-centml slHaths and their respective internodes are of equnl length (fig 12 B) while the basal and apical shenths sho- a celtnin degree of on~rlap The geeatest extent of overlap is found in the dOlble dwarf milos whose internodes aee only a few centimetels long

The plineipnl veins of the leaf sheath are much closer together thnn those of the blnde giving the outer smfnce n finliy ridged topoglRphy lhc venation is tesselate with tmnSVlIse veinlets clearly visible

The inner smface of the sheath is white and glossy often purplish at the base The outer surface is dalk green or whitish depending on the degree of waxiness fhe wax deposit is usunlly yely hell7 in the upper part of the culm and mOle conspicuous in some yarictils thnn in othCls Occasionally wax deposits 1l1a)T be very scant OJ altogether wanting

The surface of the sheath is always smooth The only hail is found on the sheath base the I(middotgion hpl( thl shenth is nttulhld to the node The hails that il1est the sheath buse arc short alJ(l white TIllY occupy a concentric band of varying width cOVlring the entire slHntll bllse and partly investing the stem tissue imnwdiatply bdow the insershytion of the ICIlf The slHath base region is also sCt ocr fom the I(st of the sheath by a slightly difr(I(nt colo genClI1lly a light olive

JlLAlm JOINT

At the junetur( of sll(ath and blade is n speCinlized region the blnd( joint The inside of the blade joint is delimited bllsipetally by the ligule Thefiang(s of the blade joint are formed by two deltoid 01

curved stll1p-shnped nleas known as d(wlnps 01 joint triangl(s 111 (S(

areas arc soft-textured and often glulfrlIld to give the leaf mobility

TIll ligule is fOlmed ontogel1ttitnlly from (pidermal ((115 and is mndc 1I p al together of ((ongnt(middotd pannehyma ((Us rill sidl Iwxt to the leaf is coveled with free or slightly adnnte hnir The UPIHr free cdgc is ciliate the cilia sometilll(s eXC(Nling in lcmiddotngtll the hcmiddotigh t of the ligule In its immature state thp ligul( is trllnslucent IflU it I)(com(s dry loenlly discolorpd especially below the uttaclunent of the cilia and sometimes torn

22 TECHNICAL BULLETIN 95i U S DEPT OF AGIUCUUrlHE

(shy I shyt~

0

F(()tJn~ 12-1 Leuf h(~ath (middotx[tnris b(yolld no(e of IHxl )taf d(wnps deltoid H I(af heath (n with node C leaf sheath shorttt thun iUl(Irno(e edge of dewlnp tLnd baoe of laminIL folded btlck D internode longer thlln shClLth (eoll1shypMe with fig J)

23

r

ANATOlIY AND lIORPHOLOGY OF SOHGHUlI

Fundamentally there is only one IigUIUI pnttern the shallow crescent Most Iigules ale about 2 mm high somewhat taller Ilt the middle and gradually thinning out toward the margin of the leaf In a few forms the ligule is considelllbly higher at the center thlln 11CUl the margin of the leaf Such types may be designated as shallowshydeltoid On the other bund thc ligule mllY be depressed in the ccntel giving rise to a shallow-arcunte type In one variety (~L N (83) the base line of the ligule in the region of the midrib wns found to be thrust upwnrd like a lllllTOW horseshoe or inverted V

SinCe the outel margin of the dcwlnp curves sharply inwurd the ligularflanges with thcir conspicuous dorsul pubesccnce ure cieurly visible Jrom the outside (fig 13 I-G) The f1angei) commonly abut squurely on the narrow-deltoid extension of the shenth Illurgin (fig 13 F) Occusionully they thin out almost to a line nnd extend to the V(Iy margin of the sheath (fig 13 E) or may terminate eadier (fig 13 G)

DEWLAP

There are two COmmOl dewlup patterns The curv(d ligulnte (fi~ 13 B D) and lhe dlllOld (fig 13 A)rhe form of elthcr type IS somewhat inflUln(ed by the slope of the ligule that mllrks the lowel houndnlS The shnpe of the dewlap is usually (ha1llcteristic of the yuliety but it may vnry within (ulm limits Young lenyes commonly have a nalTow-ligulute dlwlnp WI1(I(I1S in olelel ones thc dlltoi( type pIcdominat(gts ~Iany forms l(lnin the juY(nile typc through matUlily howcver The two dewlaps of a given leaf often show conspicuous varintion in (egaId to slope und form (fig 14) The dewlap in conshytinuity with the onllying slHlIth margin is apt to be ligulllte and stNply sloping the other dcltoid tlnd less sloping but some lenls 011 thl snme stalk may LHINe very symmctlicnl dewlaps (fig 14 Ill

rhe older ((wlnps of many vUrilti(s Hwe a pronoun(pd tpndcll(Y to fold bn(k (fig 130) whl1lns in others YCrtieal folds and guuflcling gi n the haf 111 0 bili ly

Young dewiups ar( usuully (on(olorous wi t h their 1lSlwdiTe lel1es rpon nging the dewlnp (0101 fadps be(omes ivory iyory-yellow 01

a shade of olive Thc oulpr mnrginal zonc of th( (ewlnp is nlwnys lighlCI in color than thc r(gion next to tht midrib which oft(11 rcluins the nllturul grctn of the udjl((nt leuf tissue

The degrcp of wnxiness of Lhc dwlnp Ylllips from it n)(IC bloom to n htmiddotayy WflX dcposit thnt often musks the nntulIlI (0101 of lhc oulpl dpwlup surfllc( This surillc( is sometimes glnbrous but usuully has sonw sort Qf pu)es(pne( Th(~ huils nmy (onr the entir( sulfn(e unishyformly or mny be restricted to((rLuin rcgions (fig 15 11) ~Iost (onshyspieuous and visiblp to the nnhd ey( is a smn1I hllir group extending from the midrib outward N(xt ill impOltnlH( is the brond mHlgillj1 group h(re 411( IlIlilS nrc usutllly S[HlrSp Hlld not I((ognizable wi thou t til( nid of 11 lpns SonwtinHs thesc two hllir groups arc Colllltcted by II narrow bllndof senUcl((j 1111 irs along th( bllSl of the dcwlnp hoshy(CI this hund of hllilS is often wanting Th( top of thc midrib IIllLy show n few soliliu-y hllirs but usually this ((gion is glabrous

rhc inncr dewlap surfn(c nlways hns 11 dlIIS( hlliry eonring (x(ppt in the mnrginnl ZOI1P whit-II mlly hllp f(w(l Iwits than thr out(I SlrshyJuce adjacent to it From the marginal ZOlle inward the hnirs become

24 lECHNICAL BULLErIN 95i u S DEPT OF AGlUCULTURE

o

FIGun~ 13-A Dewlap broud-deltoid undslighUy gUllffcfed n dcwlap nnrroshyliglliute und sleeply sloping C deltoid dewluJl with Iliilrgin sharply folded back sllrface not Kauffercd D nurrow-ligulate dewlup with bl~e line nearh horizontal E flange of ligllie uttenlluted und extending to vcry edge of lIIemiddotllbraIUiccoIIS sheuth margin P flunge of ligule abllts with broad base on triungular extpnsion of sheath lIIargin a uttcll1lutcd f1unge of ligule terminates hefon rcaching sheath lIIarlin

25 ANATOMY AND ~IOHPHOLO(lY OF OIHHl~[

r

f-+~~~shy

~ t

c o t ~ ~

---~

F((il~n~II-BllltI( jOilltlishollil1~ ariatioll ill ItIIlap paUtl1l on til( LIII ~tlllk A Dlllde joint fmlll Ilfar-c(nlml part of talk h()will~ SYlllllIltJild sllMpl sloping 1i~l1llltc dewlaps H IlsYlllnwtrical dewlaps from i(Hf hijdlI IIpllw I(ft dewlap in cOlltinuity with the o(rl~ing sh(llth margin is Iignlal unci slfpl~ liloping the olle to the right is deltoid Illld less sloping ( r11whlpS frolll llhul joint IlbOl( H f) Harrow li~l1latc delap~ from rOllllg(~L blade joint

long IlIHI dlnilp Jln(lting theil fulllsl dlveiopnwl1t ill tltr l1Iidlih it~(lf (fig 15 13) In nmiddot few vnlilj(middots til( bnslll mid db hair are 1101 so long ns the ndjnecnt f1nnge hnils sometimeS not (xt(IHlillg lip to t1IP marshyginnl fringe of the ligUle The f1nnge hiliI hnn II 1(JrirIlPY to (OllshyYcrgr onr the middb mnking the midrib group nppNll (sppiullymnssiYr

771j()7-15-1

26 TECH~ lCAL BCLLETIN 95 i 1 U S DEPT Oli AGHICVWURE

1

8 FJGtml j- (htltI IfI1(middot(1 iell of dewlap showing pUheSGlnce Flange of

ligl1le nhl1tlillg on membranuceous sheath margin visible at left H D(tailed strUetllrc of inner sllrface of blade joint showing pubescence of dewlaps and midrib Ligule partly folded back to expose hlLir grol1ps in back

BCCllllS( 01 its Iplntin uniformity the inner dewlap puhescellce hns little tnxonomic -aim The OutCI dewlnp pubescellce may be us(ful ns a qlllllitntivc charnetel if exceptionnUy well devcioped with the hllir grOlps -isihll to the naked eye The midlib pUHSC(Il((

howlnl (sp((inlly if extending fnl up the INtf is It character of pri shyn1lry impollnnce

27

r

ANAlOJofY AND MORPHOLOGY OF SORUHUM

ANATOMY OF THE STEM

INT~nNoDE

The stem of sorglnlm like tlult of snglleanc is a meristelcl with vascular bundles seattered throughout the fundamental paJenehymaAt the periphery (fig Hi A) the bundles ure smnll an(1 so clos( togethel that they form prltetieally a solid ring The parenchyma eells beshytween the inner bundles are thin-wltHce with small intercellulal SptttCS at their angles Those between thl plIipheJnl bundlcs [ue smnlI thick-walled and lignified Thele is It IHUlOW 1ignified hypodelmil its verticrl (ontinuity occasionaHy blOken by thp iutclpolntioll of parenchyma cells tltiLt abut externnlly on a stOll1lttp Sill(( tile S(lll epidermis of sorghum contains [clativp1y nUlllPlOUS stOll1tll(IS (OI1lshy

pared to sugareune thesf pussage ur(11S nle also rathel llunHlOUS In addition to plastids the l)ilenchyma eells of the lntCInoltips of

many varieties contain stnrch Starch deposits olC first notld in the cclls surrounding tho vnscular bundles and in many vali(tips litis jacket staJeh is the only kind formed III othel Yilietips stalph nlso lucumlliates in the parenchyma bctw((ln the bundlts This ((llmllshyltttion of difluse staleh (fig 16 B) (specinlly in lhe swcttorghlltns nnd the milos can be Ycry mnssive

Fteshly cut cross sections of young StlIlIS lllwc a uniform glftY-WtPl color As the stem matUlcs the cells nClu the centel mny h(l((Jllle lilled with air and take on a cottony-whi te appColallec Tlis PIOCtss of pith formation is often limited to the centel of the stem but it ml~Y cxttnd outward until the entile cross scetion excCpt the pCliphCIlli zone is involved The transformation of palCncbyma into pilh is sometimes followp hy a cOlllplctt c(1l1l1nllmnk-riown that lfw(s lllp cCntlft1 vusclliar hundks suspended in nil with tilt IPlllains of hlOkf1l tiSSllf)

Except in the p(Ji ph(Jnl region thc YILsClIlnl bu ndls of thC in t(rshynode ale lllOmhoid (fig 17 B) 01 nallow onl in (IOSS sl(tion (fig 17 A) Xylem and pltlopm are disposed colllttclally ill Ielation to C11Ch other The PJotoxyl0m consists of annulnI and spilitl clemen Is Since it matures hpforC thC intlIllode elongates the lI11thickpnCt pnlt of the walls of these elenlCnts becomcs elllshCt d l1ling subs(q 1I(JJ t growth Simultaneollsly the small-ctllcd protoxylcm ptllenchynm rccedes forming an ail spuee or Incnna into ddeh plOjeet the l(lllllillS of the secondary wall of the onllulnr 01 spilitl nssels Aboye and lateral to the protoylem are two large metnxylpm y(ss(ls (fig 17) with pitted or reticulat(~-pitted secondary wnlls Iktwcell tw t0 large vessels is a connecting band of smnll pitted trllchcids nnd xyltll1 pnrenchyma The tnngentinl1y Jllltttlled palenchyma cells SlIITOllIH1shying the vessels nre thirk-wnUed and Jignifitd nlHl POSSlSS 1Itieuintc thickenings The pllloem forms all onl mnss of tissue of I(gulnl design composed of Inrge sievc tuhes nnd smull (ompnnioll ((lis The protophlocm is nenrest the outside in mntun bundlcs it is (Iushpc and lignified

The vascular bllllcllts are (ne1oscd hy n llI-mnl((1 slHflth which is most strongly dt(opld Oil thl insirl(middot (fnd tw outsiltit of tht hllllshydlts whtIc it fOllllS trpienl hundlt CILPS At (itt flanks the hNllh is unistlintc 01 bis(liale occilsionally widel Th( slllnth ((lis 111(


FlOeRI~ 10-A CIOSS Hcetioll through the pCliphern1 Icgion of intcrtloclc (hand HCC[iOIl of flesh matcrial) X 40 B Bundlc from ccntral rcgion of illtcrtloclc ~lIn()ullded by pnlcllchynm eclls dcnscly filled with htrch (hand scction of flcsh Illatcrial) X 190

29 ANATOMYAND MORPHOLOGY OF SORGHUM

r

FIGURI~ 17-11 IArge oval bundle from central region of internode and B large

rhomboid bundle from central region of stem Both X 222

30 TECHNICAL BULLE1IN 95i U S DEPT OF AGIUCULTURE

compactly alJanged with no intercellular spaces between them They am long nnd tnpcring nnd spnringly pitted except nt the poundlnnks Here they nre short nnd rectnngular with circullu pits The sheath cells am in direct contllct with the protophloem but latemlly they am separated from lhe metaphloem by pnrenchyma cells The sheath cells bordering the protophloem am nlways conspicuously huge (fig 17 A) nne thinner wnlled thnll the other sheath cells

The ring of vascullll lisEc just inside the epidermis is composed of Vely smnll and mediB11middotmiddots17ed bundles buried in thick-wnllcd pnJCllshychyma The scierenchYll1atous bundle cap of the phloem pole is small and its cells are usually continuous with the hypodermal selelmiddotenchyma The bundle cap nt the xylem poe is large nnd the sheath nlong the flanges is nlso well developed The xylem of these small bundles is redueed to one or two large metaxylem vessels If two vessels nlC present they ale adjneenL to eaeh othel or sepamted rndiully by smllll-(elllti lignifi(d xylNll plllenehyma or tracheids The phloem is Ielalively well developmiddotd its arpa being proportionnl to Uw size of the IHindle

Among the oulCllllost Imndles nre n few pxtrlIlply smull ones comshyposld of 0111 mdnxylll11 VI5sd nnd some ph loem 1hlY nrc surshyrOllnllpd by It sillgllmiddot-lilY(middotIld lignifiNI shenth This type (If bundle is chnlncterislic of the bilitll intlrnode just ubove the growth ring und is nlso Hwt wilh in lhp lowel Ilodnl (lgion

[he medium-sizld bundlls of the periphernl ring arc not alwnys in eOllluet with lill hypodermnl seielllllhYl11n Ihey lIill([ (hillly from lhp smnllCl type by hning a YIImiddotY Inrge sellleIHmiddothymntolls lnp nt the xylem pol( fhere I1IP nhTnys two mdnxylplli vessp and o((llsiollnlly n Plotoxyl(rn pel11ent

The bUIl(~I(s of (he sl(ond and third eon(pntrie Iing nlso posspss a Inlgp seplmiddotnehyrnntous (np ilL the xylplll polp with 1111 illerensc in the lIumbel of plOtoxylem plellwnts There is it ploglcssivo rcdUdion in thc thiekness of the sheath in bundles farthm awav from the pcriphClY- Tho lIumber of protoxylem epllfl is largm nlld asmnll PIOshytoxylClll InculHt mny be formed Bpginl1ing npproximalely with the fOlllth (ileie 2 mm (pn tll1d fOm the epidellnis the bundles arr like those of the eentll11 intelllod(

[he ontogpny of the stem bundlpfl of sorghum is ill ngrpement with thnt of eoill (512) und SlIgHr(une (1) rhe PIWIIIsors of the bUlldl(s nr( smllll plOenmhinl stlI1l1ds thnt firflt illlIel1S0 in size by diision within tI)( stmnd 10 (onform with the f1l11llI shnpe of the bUlldle thl divisions nle I1t firfl pllmiddotirmiddotlinnl nlld Intllmiddot nt the IInnges Within thlse glOUps of plO(nmhiul11 tiSS1Il dillplI~ntintion prole((ls until nil the (ornpolHnts of the hllndlls hnve b(lPIl formld rhe first sieyc tubrs apprlll al Uw outel murgill-th( Iil-st prOloxylNll pl(lllentsshymnturingsonlPwhnt Iner thnn the firflt si(wl tuhl at the inllrr murgin of the pro(umhial stll1nd Acldilionnl (plls dillmiddot(Imiddotll1liflte 1Intil nil xyl(m and phloem is fOlIllrc Afl thtmiddot (onltiuding elen1lI1ts of the pmiddotjmiddototoxyllm nre clstroyrc dmillg plol1gatioll of tl)( intell1otlp It protoshyxylem In(1Inn ifl forllled

Ihl Imlldlr s1lel1th is initintlc1 during endy plo(amhinl dillllpnshytinlion It fllllIOlllHS (he ellHrgillg hUlldl us n single IIlYr Illit it

31

r

ANATOMY AND lIOHPHOLOGY OF SOHGaUlI

soon becomes several-layered at the xylem and phloem pole Them is no distinction between sheath and vascular tissue PlCClIISOlS in the early ontogeny of the bundle as pointed out by Esau (5) in her detailed study of the ontogeny of the vascular bundlc in corn The sheath is a composite of cells derived from the procambium and adjashycen t parenchyma cells

The procambium cells of the emerging vascular bundle and th(ir immediate derivathes have the appcumllee of a typical cambium from which xylem and phloem cells ale fOlmed This radial seriation of the c(lIs and theil lWJek-shapd appearance in cOss seetion howshyever is not sltffici(nt to argue the existpnce of a lat(ral m(Iistem in sorghum 01 other herbaceous monocots (6)

The epid(lmis of the intclI1otie is formed by a single layer of cells and is very similar in appefilance to that of sugalcane (fig 18) There are three kinds of cells Long cOIk and silien Stomates am rather numerous disposed singly (fig ] 8 G) 01 in veltical files The long cells form four-sided prisms with undulating sili~ified wulls Tlw mean diametel valies bebveen ill and 161l depending on the vUliety There is much variation in the Ipngth of the long Gells within both varietal ulld culm limits But Yllrietul diffClences are quite apparent whln forms with ypry sholt (plls (f N 6] 1) und those with CI- long ((lis Cf K 45(1) ule cOlllpalpd The end wull of the long cell is alwllYs struight neycr sloping as in mltny sugar~~une varieli(s

The cork c(lIs am suberi7ed and relulinl thin-wulltd They um kidney-shaped wilh the gllutest diallwipl at right unglos to the v(rtical axis of the stem Although usually associnted with a silica cell they may oecur alolle (fig 18 D)

The silica cplls are typicully biscuit-shaped with a constIictpd center and oV(Jhanging murginR and wit h the long diameter pUll1l1el with the long axis of the stem Occllsionally they arc broad like tho cork c(Us

Using group structure and distribution of short-cell groups as cril(ria three tpidermal putttII1S ITIUT be lp(ogni7(1 Sill(e there is often much YUliution wit hin (ulm limits t tlY l1Ie of que iOllable importance in tho designution of yurieties 1h(s( puttcrns ale-

Pattern I-Cork and silica cells always in single pairs alternating with long ltplls (fi 18 A)

Pattern 2-Rhort-clt11 groups prevalltnU~ in dOllhllt pairs (fig 18 R) Pattern 3-llany short-cell groups lack silica cells (fig 18 C and D)

It is appulent thut the epidollllis of sorghum is vcry unifollll (omshypared to tlrut of sugal(uill and is of littlo tnxonomic vllue Xot a single vuriet- out of 152 studied showcd Il eomplpoundtp abselHC of Riliea cells whereus in sugareallo mUIlY variltils slrO that ehall1cteristic Them is also only one typo of cork (ell instpad of tho grenl variety of forms o1gtsery((1 in sugar(iil1P Stomates alp Y(IYI11I11HIOUS o((uriing in (onnpded or intcllupt(c Crtienl filps Since tt(ir distrihution is VPIT UllifOllll thpy nre of no uhl( in clnsifi(nlion of vnriplipii AIRo epitPllllal hailS whilc ohRPITed only Iurply in sugalcane are always wunting in the stem epidermis of sorghum

bull bull

33

r


GROWTH RING

A cross section through the growth ring or intercalary zone differs from a similar section thlOugh the intelllode in having a peculiur type of bundle structure and a narrow bundle-free ZOIle inside the epidmmis

The cells of the outer layers of the bundle-free cortex are very small and slightly thickened j those farther inward are large with intershycellular spaces at their angles

Adjoining the cortex is a zone of very large bundles that ale so close together that they appelll like a honeycomb (fig 19 A) in which the walls of the COIllb poundpresent the intmstitial parenchyma that separates tbe bundles The (entlal 01 near-centllll bundlpounds (fig 20 B) are more widely spaced and similar in size to normal internodal bundles

The vas(ular part of the large periphellll bundles (fig 19 A) is very small consisting of a few plotoxylem elements and some phloem SUITotmding this small vascular cOIe is a massive coHpoundnchymatous jaeket which is soft at first but hard and lignified in old stalks

The ((ntral stem bundles resemblpound the ordinary type poundxcept that ull yessds ure of the annular 01 spiral types ~0 protoxylem lacuM is prpoundslnt Thpound yasculuT tissur is sUlTotmdNl by a single layrr of thin-wulled lignifipoundd ((middotlls (fig 21 A) This (pIllaYlI forms the il1lHr boundury jf the sclerrn(hymatous jaeket which is Yery massie in til( phloem region but forms only a narrow cr(lsc(nt o(r the xylelll pole (fig 21 R) The ja(klt edls innsting the nlllTOW lignifirrl bundl sheuth tili(krn und lignify progressiwly outward (fig 21 B) until the nanOw sheath us illustrated ill figurl 21 A und the sekr(n(hym atous juekrt oyer the xylem pole and along the flung(middots be(onl( one tissue 1Il( ((middot115 of til( lalgC phloem (ap UTe morC likl (olllnehymu thnn tIl( slllnth ((lis iny(middotsting the bundle as shown by their appenrnn(e and staining ((netion (fig 21 B)

A churactLristie fpatun of the bundlCs of the growth ing is the jueket of stnrch-fillrd cdls surrounding ench bundl( This juek(middott is most promin(nt Ilt til( xylCm lnd of til( bUIlltIl( (fig 20 A) and poorly d(wmiddotlop(d or wanting at the phlo(m pole (fig 21) rhe lurge plripllshyrml bundhos nlso huyC a stnl(h jaeket but it is yery narrow and less conspicuous thun that of the c(ntml bundles

Th( plmiddotculinr stluetme of thc (cntrnl stCm bundles chnlmiddotuctcrizld by tIll lurgc eolhmiddotnehymatous phlo(m eaps und starch jackPts lxt(nltis som( clistnncC l)(yotHI til( growth-ring zonpound as (middotxt(middotmully dplimit(d Also thC rNIrldion of til( (ort(middotx unci th outward mOYJlwnt of the ptliplHrnl bnndhs until tl)( again (stnblish Contuet with th( hnJoshydCrmll I selr(llchymn (fig 22 A) nrc completed only some distallce Ilboc the true growth-ring region

34 TECHNICAL BULLETIN 957 U S DEPT OF AGIUCPLTGRE

1w ntp 1n-A Cross s(ction through pcriphernl region of growth ring (haud ONriort of fr(ih materilL) X 50 8 Cross srcLion through young root hlllld showillg nUlllerous horizontal tmcrs X 100

35 ANATOMY AND )WHPHOLOGY OF SOH(1HlM

r

Flfilln 20 1 ((llml bUlldlt of -(rowth dill with prolllilwnt tlll(lt jali-ll tLrollnd tIl(gt xI(I1I tlel of bllndlpound I hand ((1 ion of frph t1Hl(ritll Ii)) H (reb (lioll Ihroll(h rllt(r n~i)lI of growth riug (hand ~ti)ll of fremiddot11 IIHltltriLiI 2middot1

36 1ECfLX1CAL BLLLETlN n5 u S DEPT OF AGH1(TLITHE

r

bull8

Fwnn 21-1 (Plltml ((111 bundlp of growth rin and Il (Ptllrlllqlltn bundl of growth ring from old stalk (Iland lections of fresh materia1) Both X 12middot1


r

Flanu) 22- 1 (rOH )Petion through peripheral regioll of sf Ill just abo( growth ring (halld (( iOIl of fr(sh matriaIL X 120 H Compound bUlldle of node X 198 C (ross S((tiOIl of stem about 8 lUll below shelth base (hand section of fresh material) X 120

38 JECHNICAL BULLETIN 95 U S DEPT OF AGRlCULTURE

ROOT llANO

Certain anatomical charaei(middotristics of the growth ring reglOnshybundle-free cOIlex large collenchymatous phloem caps and starch ja(kcts-are nlso found unclwnglmiddotd or somewhat modified in the root bUild

1he outer zone of bundles shows the honey(omb appearance charshyacteristie of the growth ling except that the large phloem (aps 01(

strongly lignified Th( outermost bundlls ha-e small phloPIll (aps and lIuge xylelll (aps Yith each Su(cleding laY(I the phlo(m Cups bl(oJlle larger until they gr(atly ex(eed in size the xylem (aps which eYlntually bp(ome reduced to one 01 two laYlrs of (pHs Holizonlal bUlldl(s bmn(hes of the yprtical system Iliay be found at tiH V(ry peliplllry of the Yas(ulnl ZOlHmiddot Otl)(r holizon tal blndl(middots (omposld of sholt pittld clemlnts and lndosing phlo(m tissup arl found seattcIN throughout the (rOss slction (fig ] 9 B) Thp~r eonslitute the tm((s eonn(eting the yas(ular tissu( of the stpm WIth the lOot primordia

The ((ntral 01 neal-(pntll11 bundlPs of the upper root-hand zone are like thospmiddot of til( Oroth Iing The xylpm is (omposNl of a row ot deJllfnts with spirtS lhi(klnings The two large latplUlly pla(pd nsspls are also of the protoxylem type Since tiHse two ((l1sale not (ollll(cted with a xylNn blidge the phloem and the Plotoxylfm palen(iryma form an oval mass of thin-walled tissue into whieh projects the row of proto~ylem (p1s The xyllm poll of the bund(middot is sU[[OI1[1(N by a broad starch ja(klt identi(al with that found in the bundlps of the growth ring In the zone below the root plimoldin the phloem cups are smolIN (fig 23 A) and strongly lifrnifiNI The star(h ja(ket alound the xylm pol( has disapp(ar(d Ill( (pnlral bundlps stillla(k a protoxyl(m la(lllla but til( two nlltllxylem vessels have pitted secondary walls instead of spiral thickenings

NODAL PLATE

Justhplow the insertion of the hoof and extending about 1 cm downshyward is n ZOlH of anastomosing l(af trace bllndl(s til( no(( propPl

hp anatomical pieture of the sttlll (oss sCelion just IgtltgtpoundOIP the bundles entCr the nodnl rpgion from Ixmiddotlow is ill agl((Il1PJll with the anatomy of thr inhll1od( in genclnl The bundles are mono or hmiddotss evenly seattend dN1Sel 11pal til( p(riphery than the ((nl(l with the out(lnlOst bundlrs in (onta(t with tIlt hypodlImal se1el(nehYJllu The central bundles are (middotIongall-oyute with well-deve1oJwd phlo(m and xylem (aps The nwtnxylcm v(sseurols arc pitted and a small protoxylem la(unn is pIpspnt

NUInPIOUS small bmnelHs split off from the bundles as th(y pnt(r the node The di-isions and nnastomosing of tiHse bundl(middots produ(e the vascular mesh that constilutps the nodal plate

The large cpntlal or n(al-llntlal bumllls of the nodal plate haye changed from oyute to broad rhomboid the phloem (aps haye (nshylargcd but the )vlcm caps are now I(eluled to fl singlr-rll II1Y(I 01

have disappeared altogether Othrl buncllrs apPlar illrguinl in shape fl(middotqtwntIy (ompound (fig 22 B) and nnastomosing highll up into separate strands The peripheral bundles have lost conta(t

39 ANATOMY AND MORPHOLOGY OF SOHGHUlI

r

P(nnt 23-A Central bundle from basal region of root band (hand SCCtiOIl of frlsh material) X 132 B Compollnd blllldlcs from peripheral region of node (hand section of fresh IImterilLl) X U3


lIGlIt(- 2--A IJnrge lnternl trace surrounded by small bundles and B part of

horizontally running median trace X 100

41

r


with the hypodermal sclerenchyma (fig 22 G) and receding from the periphery allow for a narrow bundle-free cortex The bundles nlso laye greatly lncreasNI in number nncl frequent annstomosesindicate their intimate relationship to eaeh other The larger bundles in contntt with or immediately centrud to the smnH bundles appear very illltgular )fost of them are compound 01 show signs of annstoshymosing (fig 23 B) In general therp is a gTeat increase in bUll (Ils in the peripheral part of the stem while the center appears mOle parenchymatous

Outstanding mllong the nodal bundles nre (ertnin large obliqu(lly running types thn t r(prtsln t the median nnd lnrge latern traces (fig 24 A) Thesc Inrge truces have nmphivnsalmiddot or sLmiamphiYnsnl structure with the xylcm composcclentircly of narrow clements with spiml secondary thiekeuings The Inrgc median tmce is recognized by its horizontal course (fig 24 B) liS W(lll ns by its structure

Scntte]ed nmong the larger bundles nre numerous smnH truces some running Ylrtieally others horizontnlly or obliquely Th(y nre eomshyposlc1 of short-pitted xylem dements enelosing some phloem in selllishynmphivnsal or Int(lfll[(lTnngpment 111(ls( small tru(fS nnnstomose very frequently nnd chnnge their course from the Y(lticnl to the horishyzontnl nnd back ngal11

Just below the insertion of the l(af the p(ripheral bundles recnter the cortex in grcatly altercd form and cxtend thence upward into the lenf

The course of the vnscu]al bundles and the relationship of the leaf truces has been desclibNI in detail for corn (5 12 13) nnd sugar(nne (1) Sorghum shows no cltyiatioll from the normal pnttell1 1shythough the nwcLian tlfice (fig 24 B) nppoaches dose to the ceuter of the stem the lnrge lattrul truc(s (fig 24 A) btnci only a few millishymeters inward fi behador thnt is mort in ngrNmfnt with Shnrmnns conclusion (12) thnn with Esaus (5) who innstigated the ](aI-tmce relationship in corn

ANATOMY OF THE LEAF

SHEATII BASE

The shell th base is composed of (latinIy meristema6c tissuc from which the sheath e]ongntts It is comparable to the growth ring or intercalary zone of the stem

A (ross seetion through the basnl part of this 1(gion shows the ascu]al bundl(s clistrihutNl morp 01 ](ss evcn] OWl the entip cross section (fig 25 B) Tht smnlllst hundhs I1rpIl(xt to the olltpr (Ipishydermis the ]argpst nrC llpnr-((ntlfil som(what (Ios( to the inner epidermis All bllndlls hnn enormolls phloem (nrs hilt the ynsclIlnr cor( is yery smnll consisting only of a f(w spirll 1 (I(lllpnis The l~rg(l bundllt~s (fig 25 C) nr( similnr in strueiurp to those of til( powth lmg

In the upppr pnrt of th( shenth joint the bundlfs nn more (onshyc(ntrnted in 01lt outer periph(rnl rfgion of tlle (-ross sP([ion The ]nrge bundles POS5(SS enormous phlo(IJll nps Ihnt nl( Iignifi(ld wholly or in part (fig 25 C) TIle YnsCulnr (orp is surlOlll1d(d by fl IlnlTOW lignified jacket as in the growth ring (fig 21 A) The two melnxylpl11

42 TEClINICAL BULLETIN 95i U S DEPT OF AGRICULTURE

Fwung 25-A (104 sec[ion throllgh periph~ml J(gion of sheath base showing hairs (hand sectiQn of fresh Illltteriai) X fi4 If Cross secLion through basal region of sheath base (cross section of fresh material) X 34 C Large bundle from sheath base (hlmd section of fresh nmterial) X 110

43

r


vessels are pitted and connected with one another by a narrow xylem bridge (fill 25 0) A protoxylem lacuna is wanting The phloem is especiafiy well developed and more massive than in the bundles of the growth ring

The dense matting of hairs (fig 25 A) that delimits the sheath base externally is of Stlbepidermal origin The basal part of the hair is strongly lignified the free part is of cellulose

LEAF SHEATH

The leaf sheath is traY(lsed by vascular bundles running parallel to one another They are join(d at intervals by cross-connecting vein lets similar to those described by Moreland and Flint (9) for sugarcane

A cross section of the sheath shows bundles of different ranks In the narrow part of the sll(ath small and Ituge bundl(s alternate but in the broad central region an intermediate size is interpolated (fig 26 A)

The smallest bundles lie close to the epidermis Since they are formed late in ontogeny they lack protoxylem Their sclerenchymashytous sheath is narrow at the xylem pole massive in the phloem region and always confluent with the hypodellllal scl(renchyma

The medium-IarOe bundles poss(ss protoxylem but often lack a lacuna The bundYe cap of the phlo(m pole is (itlwr in contact with the hypodermal seler(nchyma or separated from it by several rows of parenchyma cells (fig 26 A)

The largest bundles (fig 26 B) r(semble in structure the central bundles of the internode except that the phloem of the sheath bundl(s is more massiye The seier(nchymatolls bundle cap at the phloem (nd is in contact with the hypodmmal sclerenchyma except where the bundle occupi(middots a near-cpntml position Radial sheets of chlorenshychyma about fiYe cells wide (fig 26 A) connect the YVIm pole of the large bundles with small groups of thin-walled hypouClJllal sclerenmiddot chyma of the lopr epidermis rrhe area between the radial sheets of chlorenchyma is filltmiddotd with large empty (ells rrhe tissue appeals white and spongy in old sheaths but there is rarely actual cell break-down as in the sheaths of sugarcane

The transverse bundle connections are in the nature of branches that run obliquely or at rigllt angles between the large yertical veins and effect a union between them RtlUctulLlly and ontogeneticully they are similar to the cross-connCctillg veilliets of the lamina consistshying of seveJllI rows of shod-urtieulated xylplll clements of the reticulateshypitted type rhe bundles ale formed late in ontogeny tllpir initials connected witll the cambiumlike region of the vertical bundle system

The outer epidermis of the sheath in the region between the veins eonsists of regular long cells altpmating with gIOUpS of shOlmiddott cells The epidermis overlying the veins is com posed entirely of cork and silica cells llanking tlle v(in arelt and separated from it by two IOS of Ipgulal intercostal epid(rmal cells arc single vertical files of stomates Topograpliieully the vein arpa appCars depressed (fig 26 A) the intercostul area sliglltly mispd The innCl epidermis is composed ~olely of long thin-wallecl cells

44 TELHXltAL BlLLETIX lIii t ~ DI~PT or (H1((LITHE

111(1 Ilul hlndp I JlllldllI-(middotjIlPd likP till 1Iltllh but til( 110shy(OIlIlP(tillg pjoltmiddot1- un lIot 0 prolilill111 II III hc 11(11(11

TII(Ipound tlrl (0 tY]1ps of ((1I1111 1111 III lip illltlil roUlld OIlIS ill glOllJH of 7 to J) a[IllIlillill wItll larp otll 1lll1d[( rfi 21 11 TltllllI hlllHllps i(lpnPlltillg [lip plillipill middotin or [h Iaf occupy III( (1IIi1t c1lpllt or tl1( llttll (10- S(([1011 (fig 27 tI) til( smull Ioulld 11)( is llIlilldd(ld ill pnl(Il(IIYllIa ill (10( plOxill1ity to t[lp [O(l lpidplllIisIIi 27 1 l

Ilp bu( hUlldh ill (IUllllln[Jy ilililal to tllO( of (lip 1(lnr slllnt II I~H(II bUlIdl1 i jnltlclaquo1 by fllllll1(I [igllifipd -11(middotnlh hi(11 I in ((lilshy

t Il1uit wit h t hl hypod(llIUll wl(]lJlCYIllIl ilt lh( plllollIl poll lillt j lpnla(pd flOill IIt( xylem poll b n ingl( Inylt1 of larg( thill-itll(d plnll(h1Ht ((11 111( n(lJ1nl part of (h( 1gt1lll1l1(middot ii Jikp lllnl dpshy(rilJ(( [or lhp 1l(lilth

Ill( lllalllJulldl( or til( l(Hf IJlndp (fig 27 H) 111( l01lnd 01 Iihth 0111 Snwll nod -orJl(llwt 1Iff1 Illllldl( IIltplllal( wit It (ll( fll(tltpj

TIH Inlgltl t~IH ili 27 bull 11 (((11) IlOI (PIlIlnl 0Pfloil( it but (lpnlltpcl by fi( 01 ix InII or PIIIPIl(liYIIlH (pll- j n Illnll glOllP or middotwllJ(middotncImll ill (olllnet Iilh Illp upper ( pi( Illli Thc n(ulll pnll or til( bUlldlp cOIlit or n rpw pill(d Xylplll ((1 nlld n IOIIJl or ph[O(l) ((lIs (fig 27 B) S1IIlOU11lt illg (iI11 IHllallp j it jn(( or 11l1l

XAT()~[Y AX) ()IPllOLlllY UF Omilll[

r

amp

--ic l((dIU i bull bull 1 (10-- (tiOIl of har 1trIp ltawl ((middottill f fmiddot-ll IIml(middotdd

bull Ij() H (Iq _I jilt IIr har ClIiaCfd lImit tllll 1II~1I HIj- I ( Pari illl (10 111 IoU I troll I Lmiddot luidrilJ blJ ii diIIilIlI ilJl IIf 11111111middot haud Pl1 illll r frl-11 malmiddotIhl li[j

46 TECHNICAL BULLETIN 95i U S DEPT OF AGRICULTURE

thick-walled chlorenchyma cells containing nllmClOllS large plastids All bundles are connected at frequent intervals with one another

by naiTOW branches that run diagonally (fig 27 A) or at right angles to each othm Structurally they are the same as the cross-connecting veinlels of the sheath

The mesophyll of the leaf consists of relatively compact chlorenshychyma with p1astids that are smaller and more numpoundrous than those of the chloIophyllacpoundolis sheath surrounding the small bundles TheIe is no well-devdope( palisade layeI

The cells of the epidermis of the blade are similar to those of the stem except that they me thinner walled and vary in the percentage distribution of the various elements The cell patlpoundtns of the 10wlI and the upper epidlrmis show a periodically rpound(UITlnt dlsign that is 1(shylated to the type of (pound11 underlying tho lpidolmis Except for the added presence of bullifonll cells (fig 28 A) tll(rl is littll difterence in structure between tIll UppOl and the 10wlI llaf lpiclrmis

The epidermal cllls that oYldie the vascular bundlls form several longitudinal IOWS their number dlpending on til( size of the veins The cells are long and narrow (fig 28 B) and may lie lnel to end in a series but usually they abut on a cOIk-silica glOUP The centm of the zone is oc(upied by rows of long cells altell1nting with rows made up of cork-silica groups only Flanking tho vlin zone on eithpr side are s(weml rows of lpi(krmal (ells compos((1 of long cdls that lin pnd to end 01 of long cells nltell1ating with cork-silica groups 01 cork (plls only Conspicuous are the vlItienl fillS of stomates oc(urring in single or double rows Thl long (llIs hehn~ln the IOWS of siomatos contain small two-Cpoundllld apPllssld hairs that tnke the placl of the cork-silica group O((nsionnlly there ale found short spines and twoshycllled hairs occurring bltweln the long cpllf flanking the veins TlIlY all of the type dlscribed for tho epiciPlmis of Sllgal(ane (3) The bullifolm 01 motor cells OCCtll in single 01 double files (fig 27 B and fig 28 A) and are not npoundally so plominlnt as thORpound in sugarcane

The midrib of the blade appears crlSClnt- or half-moon shap((1 in cross slction The lower sUlface is studded with asculal bundlls (fig 27 C) while the UpPll is undpoundr1aid with a thick band of hyposhydlrmal sclllenchyma The mass of tissue bltween the bundles and the sc1lrenehynul is pallnchymatous

The parlnchyma of tl1l midrib of young Ilavpounds is rich in sap Vilwld from aboyl thl tissue has a watpoundI-soakld applamncp As thl leaf matures the parcnchyma I)((omes filled with ail giving the lntile midrib a whitish appeanU1CI that contrllsts strongly with the grlen (0101 of thl lamina In YariEtirs with color((1 midrihs a pigshymlnt is found in thl parpoundnehyma U1l(lCrlying tht UpPPl epidllmis

The hundlls of the midrib ale like those of the shrath IJalge and small bundles alternate (fig 27 0) and all are (onnlctcd with the 1011 epidlrmis by hypod(rmal sclerenchyma The pCrcpoundntngp disshytribution of the large bundlpounds how(yer is much gnat(r than in the lamina wle the large bundles OC(lU on1y at gllnl intllvals

Tlw 10wlI lpicilrmis of the midrib is similar to that of the lamina eX(lpt that the longitudinal files of (olk-silica groups and thl IllUIOW long (ells aIpound moll l1UmplOUS This is to be poundxp(eipd sinee all yascu1al bundles of th( midrib abut on hypodpImal sclpoundJ(n(hyma


r

FIGURE 28-A SlIrfacemiddot vicw of IIpper epicermif of blade B surface view

of lower epidermis Both X 170

48 TECHNICAL BtLLETIN 957 U S DEPT OF AGRICULTCRE

FlGnl~ 29--1 Cross petioli throu~h d(lIiap (hand ctiOIl of frp~h lIIaf(riai) X GO n Enlar~((1 d( from lIIar~illai dowil1Jl rlgion howing dttailcd itrlldurc of iarp hllndk X 2middot10

The upper lpidtrmis of till midrib is mndp liP of IUlge long cells nltlrnating with cork ((115 Thel( 11( 110 sili(il ellls lnt nO 5tomntcs In (lImiddottnin varilmiddotli(s mnny (pi((ormnl tells ()spNinlly in thc tentlnl Iegioll hnn glOwn out into InJgp ulliedllllnl hnirs These IHlirs Ue of epidelmnl origin nncl do not 1)((on1( lignifi(middotd

1)1[1

Ihc nnntomieul pidul( of lIw (IpwhlP I(S(1I11gt1(5 that of the Inmina but till l1sculnl bundks IWHr l eiowr IPintionsilip to those of tlH shCflth base

r

ANATOMY AND MORPHOLOGY OF SORGI[ln-I 49

A cross s(ction through th( dewlap (fig 29 In s1l0s n su(((ssion of Y(ry large und VlIT smnH bundks embedded in par(ll(hymn S(vernl nry slllllLl bundles nlternntc wilh 01H lillg( bundle The Intter occupi(s the centt[ of the cross s(ction whilt lhe smnll bund(s have an oir-central position closer to the lower than to thc upper epid(rmis

TI( Inrge buncllts art sUlTound(d by a nntTOW jocht of thin-walltd lignified cells (fig 29 B) Tlw phlo(m is eo Plwd wi th (olltllchyma thnt exttnds to the locr epiclPrmis (fig 29 L1) A similnr ~roup of collenchymn is found nt till xylem tnd of the bundh but it IS not in continuity with the shtuth (Plls i)ping sltpulOted from them by one 01 two luyers of palenehymn The xylem is eomposed lntinmiddotly of eltllwnts with spiral secondtllY thiekenings A Plotoxylem lacunn is wallting

The small bundks of the de-lnp contain a f(w nnlTOW protoxyl(m elements ilnd some pblocmiddotm ThlY nlc jn(kPlld with a hrOltd Inytr of colknchYllla thnt is especially prominent 011 the phloem end of the bUlldle

Both cpickrmal sUlfnces ar( ClothNI with hnils likP thos( found on tlw shealh joint Ill( (plls of llil lpitiell1Iis nre ilTlguhu and lliillshywallld stomates arc present

ANATO~lY OF TIlE ROOT

LAHGE LATEltALS

A (ross scdion through a Inrgc latcnll root shows a sipholloRtlle with a lentrnl pith nnd brond eorl(x (fig 30 AJ The eortlX is eOIll shyposld of 12 layers of ((lis Ill( IWriph(middotrul litytr fOIl11S 1111 rxod(Imis that hns plOmillcnt waIL thiekenillgs involving lhl ouLCL tang(nlill wnll Sorghum difrcls in this llsl)(ct from Rugnl(1IW whose fooL exoticImis luts U-shapNI thiehnings invohing the illlHr tnngPlllinl and mtiial walls ExtPlnally tlw (xodlrrnis ngtuts til( usual t-IW of lhin-wnlkd root ppirilrrnis ill whieh mallY of the ((middotlIs ItOYt (ongnlpd to form root hailS 1hl bulk of tlH cort(middotx is eompos(d of Inrg(middot l(gular ((Is wi lit sq ullrish in t(JCllLuliu spn(Cs 1)(tnPIl tltpm Till ((lis of the two most (pntrnd lnYlrs dos(st to the enclodermis nle bliek-ShnllCc IInllOW ancl Y(IT Ilgular

lhr (cUs of tbl (middotndod(middotrmis are (ry uniform and there a I cmiddot no inlel(Cllulal spa((s bCt-(cn them At mallllity till ((lis han th(shyU-shnved typc of thielning in whilt-h tlH inlwr tnngltnlinl wnll nnt purt of the ntijfl(pnt mdinl mILs urI r(infol(pd (fig 30 C) The tt1nglnlial wnIL thi(kpnings aI( topped with Rilin knobs of irl(gulnl contour (fig 31 Ii l Silica (x(I(Lions similar to lIJ(S( bUl mol( I(middotgulnr in shape tn( hl(n dts(ribld by thl f1uthOI 101 UI( (l1dodermis of sugar(lllle (J l pl 23 D)

The Yaslulal ring is 1(lnlilly 111l1lOW (fig 30 A) and is limited on the iURicir by a s(l(rclHhymntolls ju(rt thnt follos lIw undulnling (Ol1tOllr of the inwardly proj((ling llwlnxylcm Y(ssc lhlmll11h(J of protoxylrlll points is -alinlJlr dllwntiing on lhe si7l of til( root and thc number of mcolnxvl(m (hmrllt~ Thlr( 01( (ommonh~ thr(( protoxylem Rtrnnds and nil lqwd llumbrl of phlo(111 groups fo (u(h n1Ptaxyl(m VliSll Bolh p1oloxy(m and phloem ahut till pNieydc The laUer consists of n single layer of cells that become thick-walled

50 rECHXICAL BULLETIN 95i U S DEPT OF AORICCLlTRE

FIGURE 30-01 (ross section Ihro1ph IlLTC IUINal rool (hand pelion of fresh llIateria) X 33 13 Cross scciOIl throuph hltltn joot (hand S((tiOIl of frcsh lIla((rial) X 19 C Partial elllarl(lllent of I to how dttlis in structurc of endod(lmi lnd fi~cllar tissue (n Endod(rmis 1IX metlLshyxylem (sHel p1 protoxYllI 8 lti(gt( tlllgt( j p perieyc(gt sk si iell kno) X j(i7

51 ANATO~[Y AND MOHPHOLOGY OF SOHGHGM

r

FG1IU 31-1 (ross ectioll thlOlIgh nlsclIlar tisuc of large Jutpral roof CT

Endodermis with silica knobs 718 protophloem sir( tube s large UI(middottnshyphloem sieye tubr r compllnion cpll ]i pnlieyel(gt X 1000 n Longiludilllli seetioll through baRnl region of root tip iihowing apical JIl(middotrisltmiddotlIl X 425

52 TECHXICALBCLLETI~ 05i U S DEPT OF AGRICCLTCRE

fairly clrly in ontogln) In I()(nli~((l regions Ihcl)(lilyee is two cells tluck

The phlolm glOups nle V(IT uniform (OI1SiStillg of fh( 01 six bull el(ments (fig 31 A) lhlle is nmiddot single pIolophlo(m sieve tllho bull (fif 3211 nnd fig 31 A) Jll1nked by lm (ompnniol1 (Ills of similnl si~1 (fig 3] A) Cenllllcl to this lIind of ((lis is 11 vely lnlge silve luhe with a hOle of 20~ to 22~ und above it ure one 01 two sieve tll bps of smnll(I si~I

The (ells SUlTOlllHling the xykm und phloem thieken und lignify with ag( so thnt in old roots the enlill vns(ullll Iing is composed of thiek-wnllNI lignified tissllc in whieh only the phloem (ells rcmain thin-wnlled and nonlignifilll

The pilh is (omposNI of very reglllul tis~ue tlw (ells arc roundish wilh illtel(dlullll spn((s b(teell th(m The wnlls of the pith (ells of ollaquoll Iools Ihif]Wll nnd lignifi(aliol1 which is nt first spotty overshytakes most of the tissue

III the denloplll(lll of the vns(ulnl tissue the IllIge metuxylem elellHnts urI initinl(d befol( th( PlotoXyl(lll (fig 31 B und 32 It) but th( Inllt11l11111ll( enrliCI ThC plotophlo(lll iCve tub(ls (fig 32 B) 111nl1I( nhNHI of lhe PlOtoxyltll1 in ngl((ll1(llt Iilh tissue difrerentiashytion ill oihel plnnts

IkTTHESS HOOTS

The stludulC of thl lllHl(lgTOUIHI pn1 I of the buliless roots is i(Cnli(nl wilh thnl of lh( InlgC Intlmls rh( nCrinl S((tiOll ho((l is difi(rrnt QllHntilntiC difiP((IH(S eonsist in n glent(r root dinmshy((PI H Inlg(( ntllllJrr of nHlnxyltlll Y(gts(1 nml eon(omitllntlYI1 g((ln(PI lllllnlHr of protoxylrlll points nlld ph1c)(l11 groups (fig 30 R) nnd n Igtroult(1( (ort(x ThC groups of phlo(lll (onnin 11 In 11(11 l1UIl1Jel of lis Ihn 11 tllC IlllW In terHls nnd Ul( 111111 ngrn1(n I of (ho pltlorlll ((Is is lCss symmrtrienl Qunlilftlivply t1( nNinl pnrt of lhe hlltshylrss 1001 difi(rs fro1l1 thr 11lld(middotrground pnrt hy hnving It strollg rpidrllnis in which thr OlltPl lnllgltlItinl nl lind 111(1 rlldinl walls IlIe rein fOI(ld 11( NHlod(rlllis wltilr similnr lu(ks siliell k1l0bs Thple is no distin(t CxoclImis ilh lnng(lltinly Ihi(k(nCc 1115 but a ll1odifi((( ptlipllPlUl (OICX eonsisling of tIll(( or foUl layers of smull th ick-wnlkd (ells

Ill( filiform IniNllls of lh 5Pond nnltl third 01((1 nre whitish Illld of approxilllil icily (((11ft d innwll

Th( IInHlol1lhnl slrlltillr is simpll ln(h loollrt consisls of n hlond (ortlx (fig 33 1) sl11Tollnding n miniutllr( sill( ith th( u1111 type of tissue fllTnllgfl11(nt lhi stnl(lll1C difiels chidly from lhlll of the Inlgl 1001 in lacking nn (xO((Imi and poss(ssing only on( 01 two n1(t Ilxykm v(ss(ls

TIt( (ntllll (orC is ex(((diugly smllll (fig 33 13) hning a bore sOni(what smnlllr thnn lhnt of n sing( n1(lnx~Itm (s(1 of n Inlge Inlplnl Thl Cl1(odllmnl ((15 nrl of t11( tYPl d(s(riigtNI fOl lhl Inrgc lOOl (X(ppt thnt tllP siliea knobs nTP oftln wnnlillg nt IWR ill(onshySpi(llOllS Thpr( is n 111lilnY(rlc IWri(T(I( (onllisting of ntPlllnlillg $(tOlS of ttick-wnlled nnd t1lin-wllllrc ((lis nIP prliey(lic ((lis opposite the phlocm groups UlC usually lhill-nlled (fig 33 B) The

53 AN AIO1[1 AND 1WHPHOLOO1 OF 80HGI-IllI

r

FIGlIn 32~- 1 irallYltrH ((liOll of root lip l(lkclI JjOIl frolll t hlt apillll cdypshy

t rogltll III Clisllt III r tOOl (ltp (ltgioll 111 IlHtaxylCJIl Y(~(I illititLl~ X I-I) H TIlIlf(Slt ((1 ion t III-ol1h root ip tllkpll IjOIl from tho api(lit calyptlOg(1l llI(rist(nI (1 I~lldod(rlllis ps pIotophlo(1ll icYO lullj mI lHolfixylclll(lt L X 7-1-1

54 lECHNICAL BULLETIN 957 U S DEPT OF AGRICULTURE

A shy

bull

~ 8 yen~ ~

FIGum 33-1 Cross Hcction of Il1null laf(ml rootictX 18middot1 n Cross ection of central cylinder grently enlarged cn EIlc1o((rlllisi IX llletlLxylem vesseli p pericycle px protoxylell1 8 sieve tube X 799

smallest roots with only a single (elllllli melnxylcm v(Siel usually have foul Plotoxylcm points and an equalnumbel of phloem groups In steles with two metaxylem Ycss(ls the numbCl is slightly Inlgc1

55 ANAlO~IY AND MORPHOLOGY OF SORGHUM

The cortex of many rootlets eventually breaks down and becomes

r disorrranized Such rootlets have a brownish exterior Their funcshytional capacity as organs of conduction is plobably unimpaiIed howshyever as the vascular tissue protected by the thick-walled endodennis appears normal (fig 33 B)

(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED

AUTSCHWAmm E 1925 ANAIO~I1 OF THI VIWETNIIVE OIWANS OF SUGARCANE Jour

Agr Res 30 197-221 Hlus

1930 A COMPARNrIVE STUDY OF IHE SIIM IWIIJIollUIIS O~ (ERTAIN SUGARshyCANE VAlUEriES Jour Agr Res 41 853-865 illus

1940 MORPHOLOGY O THE YICWrAlIVE OllGANS OJo SUGARCANE Jour Agr Res 60 503-549 ilus

1942 A CO~IPAnATIVB NALYSTS OF TilE VEGETNIIVI-l CIIAUACIEUISTICS OF SO~II VAltIANrs OF SA((HAlIUM SlONIANEU~l U S Dept Agr Tech Bul 811 55 pp illus

ESAt K 1943 ONTOmNY OF THE VASCULAR BUNDLE IN ZEA ~IAYS Hilgardia 15

327-356 illus

1943 ORIGIN AND DEVELOlMENr OF IRIMAR1 VASCULAR TISSUES IN SEED PLANrs Bot Rev 9 125-206 illus

JEsWnlT 1 1916 DES(lImJVING D)RSOORrEN VAN JI~I SVIfEnRnl~ ~m~sn HIJshy

DUAGE ~IOII1I0LOGH VAN HJolI SUIKEIIHUI Arch Suikeshyrindus Nederland ludic Medeel 24 [3591-429 ilJus

M ILIEU E C 1916 COMPARATITIl SrUDY OF rim nOOT SYSTI~lIS A)o1D IEA F AREAS OF

COliN AND THE SORGIlO~IS Jour Agr Res 6 311-332 ill us ilolullAND C F nnel FLINl JJ H

19middot12 rim DI~VrrOIIENr Or YS( (TIn (ONNI~(TIONS IN THI- LEA SHBATH OF SUGAII(AN~l ArneI Jour Bot 29 3IiO-3(J2

QUINn) J R nnd KARPlm R E 1945 THE INIHJIIITA)o1(l OF THRlm Gmms TnNr INFLtT~lXCE Tum OF

FLOHAL lIIIN1ION A)o1D ~1AIUHllY DATE IN MIIO Amer Soc Agron Jour 37 916-03(J illus

REZNIK lL A 1034 ETUDFJ ANAIOIIQIIE DE JAllANrULB DB SOnGHO Re Gen ele

Bot 46 [385]-119 illus SnAn~IAN B C

1942 D~VFJLOP~IJo]NTAl ANAIOIY ot Ill SHOOT OF ZEA MAYS L Ann BoL [London] 6~ [245]-282 illus

S1RASlItT nmm E 1891 UBJolU IH)o1 HAlT UND nUJ VEltRWIl1INGFJX DrJR TEITUNnSlIAHNEN

IN DIJN IfJANZEN In Strasburger F HisLologische Beitrage v 3 1000 ppbullbull illus T(nn

YINALL H N SlEIIHJ)o1S 1 C nnd lLlllIN 1 H 19313 lDlNTIFI(ATIO)o1 I1ISTOny )01() 1l1S1middotIUIllTIOX OF (O~UION SORGHUM

VAIUEJJoS 1middot H Dept AI-(r Tech Bill iOU 99 pp illus WEA vmt J E

1926 nOOT IHnl(OI~IBNl OF~HrD (HOIS 291 pp illus New York and Toncloll

UScGOVEFNMENT PRUHING onlCpoundt 1948

lor 51110 by thl ~llllPlll1lpIHIfl1t ltJf J)O(lllJl(lltll r i (1O(lII1rt11nt Printing omcc middotashillgton 5 D C - Pll~l 20 C(llts

t

---- - shy

Technical Bulletin No 957 bull JUlie 1948

Anatomy and Morphology of the Vegetative Organs of Sorghum Vulgare 1

By ERNSI ARTtlCHWAOBII Blnior 1)la1l1 (watolllist DillillOIl of Sligar Plant hlllcsshyligations Bureau of PllInt Indllstry Soils and Agricultural Engineering Agricultural Research Administrationl

CONTENTS Pugc Page

Prcvious investigations________ _ 1 Anatomy of the stem _________ _ shy-Internode _________________ _ Matcril~ls and methodsbull______ _ 3 27Gross morphology ____________ _ Growlh ring_______________ _ 3 33Root blLlleL ________________ _ Morphology of thc sttlk _____ bull __ 8 38

Internode __________ _____ _ NodlLl plate____ _ ______ _ 8 38Nodc _______________ _____ _ bull 1 Anatomy of the le~f _ _ ____ _ 41Growth ring_____________ _ j 1 ShelLth base ___ ____ ___ _ 41Root band_______________ _ Leaf sheath_______ ______ _11 43Bud ____________________ _ 12 lelLf blade____ -- ________ _ 4418 Dewlap _________ bull _________ _MorphologyHli the IcaL ______ _ 48LcGObllLdel ________ bull ______ _ I) Anatomy of the root__________ _ 40 ~

Ieiltshllnt)_______________ _ 20 Large laterals______________ _ - 4) 21 Butttmiddotpss root Hbullbullbullbull __ bull ___ _ 52- Bl~joint---- ----------shy______ Sl II e ~ ~ _ 21 Small Inteml _ 52

= ~lcGwlap -- - - - 23 LiterntlIrc eiLeei __ 55

~ J ~ gt 0 PREVIOUS INVESTIGATIONS SO~hU1i~Svlglmm llljate PP[s) is a plltut of It1I(i(m lillrugc nmiddotll lt Istublislwcr as all pcollomie CTOP (Yl1I ill Uw lHdi(st dilizH tiOllS The litelLtllll dpuling wiLhits history gtographi( disltib1ltioll alld 3 botanieal d(scription of vlLri(middotti(s is (xtellsiv( It hHs (1((IIUy b(tll

Tcviewcd in (ttuii by VillHll Stlpll(ns nnd ~lllrtill (14)1 TlHir tleatise gives the uotnlIienl ci(seliption of muny sOIghlllll vnlittits an(l a key for stpnmtillg th(m

in til( dps(Iiption of sllgnrenllc varitties minut1 mOlphologicul and eV(It anntomical (hn[lwt(rs UI( us(d hilt in sorghum taxonomy rcmiddotiiancc hns bN1l plltlNI chidly on thl SlructulC of illnOnS(PII(P spikehmiddott and sllmiddotd Tow SUIP sOlghulll shows a Iplativ pnueity of such ehnlll(ttrs ns Conn til( basis for suglu(Il1middot (lnssificlttioll but somc of the (hnruettls nle of glNt distin(tll(Ss unci of I(ILI vuluc in the des(ription of variCti(s and ill the annlysis of a hyblid populatioll

Th(I( is no litelutul-( dluling sppcifillIlty with thc millut(middot mOIpholshyogy nnd anatomy of SOlghll1ll pxcppt for a papPI 011 thl dlv(lopn1lntlli allntomy allCl homologi(middots in til( young R(middot((lling (J 1) but its stluClllr( I Rubmitlecl for Jlublication Oct 11 1)4fi

2 Credit is duo )Ii~s ROH~ Mary Schipman for the prppl1f11lion of till drawings 3 Italic nlllllbCls in parentheses refer to Literature Cited p ~5

78605iO-48-1 1





Morphological





























GROSS iIOHPJlOLOGY









t

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J 1________________1 17 23 16 27 231 28 10 26 18 20 16 23 14 16 -_____ ______ t1 10________________1 191 22 20 2(i f 23 I 26 I 21 27 21 20 16 24 18 17 ______ ______ ~ H_________________ 21 22 15 2(i 22 21 21 30 20 19 12 23 20 18 328 28 Y 8_______________bull 23 23 18 281 23 28

1

22 30 19 19 15 22 19 19 5 17 7 ________________1 23 24 12 25 23 27 24 30 20 19 17 23 22 19 3 14 ~



I 1 I


fill

r




I Starch 1


-----------------------------middotI------~------I------




475 584




NODE


GltOWTfI IIING


HOOT IIANll






lIun











r










I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

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23

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DEWLAP








o



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f-+~~~shy

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771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy





Morphological





























GROSS iIOHPJlOLOGY









t

I r I

c















INTEnNol)lo~









r

~~~

~ lt7 ~ I~ tti~~~~~r

HB J~l Ifr






J 1________________1 17 23 16 27 231 28 10 26 18 20 16 23 14 16 -_____ ______ t1 10________________1 191 22 20 2(i f 23 I 26 I 21 27 21 20 16 24 18 17 ______ ______ ~ H_________________ 21 22 15 2(i 22 21 21 30 20 19 12 23 20 18 328 28 Y 8_______________bull 23 23 18 281 23 28

1

22 30 19 19 15 22 19 19 5 17 7 ________________1 23 24 12 25 23 27 24 30 20 19 17 23 22 19 3 14 ~



I 1 I


fill

r




I Starch 1


-----------------------------middotI------~------I------




475 584




NODE


GltOWTfI IIING


HOOT IIANll






lIun











r










I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

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---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy







GROSS iIOHPJlOLOGY









t

I r I

c















INTEnNol)lo~









r

~~~

~ lt7 ~ I~ tti~~~~~r

HB J~l Ifr






J 1________________1 17 23 16 27 231 28 10 26 18 20 16 23 14 16 -_____ ______ t1 10________________1 191 22 20 2(i f 23 I 26 I 21 27 21 20 16 24 18 17 ______ ______ ~ H_________________ 21 22 15 2(i 22 21 21 30 20 19 12 23 20 18 328 28 Y 8_______________bull 23 23 18 281 23 28

1

22 30 19 19 15 22 19 19 5 17 7 ________________1 23 24 12 25 23 27 24 30 20 19 17 23 22 19 3 14 ~



I 1 I


fill

r




I Starch 1


-----------------------------middotI------~------I------




475 584




NODE


GltOWTfI IIING


HOOT IIANll






lIun











r










I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy







t

I r I

c















INTEnNol)lo~









r

~~~

~ lt7 ~ I~ tti~~~~~r

HB J~l Ifr






J 1________________1 17 23 16 27 231 28 10 26 18 20 16 23 14 16 -_____ ______ t1 10________________1 191 22 20 2(i f 23 I 26 I 21 27 21 20 16 24 18 17 ______ ______ ~ H_________________ 21 22 15 2(i 22 21 21 30 20 19 12 23 20 18 328 28 Y 8_______________bull 23 23 18 281 23 28

1

22 30 19 19 15 22 19 19 5 17 7 ________________1 23 24 12 25 23 27 24 30 20 19 17 23 22 19 3 14 ~



I 1 I


fill

r




I Starch 1


-----------------------------middotI------~------I------




475 584




NODE


GltOWTfI IIING


HOOT IIANll






lIun











r










I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy





t

I r I

c















INTEnNol)lo~









r

~~~

~ lt7 ~ I~ tti~~~~~r

HB J~l Ifr






J 1________________1 17 23 16 27 231 28 10 26 18 20 16 23 14 16 -_____ ______ t1 10________________1 191 22 20 2(i f 23 I 26 I 21 27 21 20 16 24 18 17 ______ ______ ~ H_________________ 21 22 15 2(i 22 21 21 30 20 19 12 23 20 18 328 28 Y 8_______________bull 23 23 18 281 23 28

1

22 30 19 19 15 22 19 19 5 17 7 ________________1 23 24 12 25 23 27 24 30 20 19 17 23 22 19 3 14 ~



I 1 I


fill

r




I Starch 1


-----------------------------middotI------~------I------




475 584




NODE


GltOWTfI IIING


HOOT IIANll






lIun











r










I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy













INTEnNol)lo~









r

~~~

~ lt7 ~ I~ tti~~~~~r

HB J~l Ifr






J 1________________1 17 23 16 27 231 28 10 26 18 20 16 23 14 16 -_____ ______ t1 10________________1 191 22 20 2(i f 23 I 26 I 21 27 21 20 16 24 18 17 ______ ______ ~ H_________________ 21 22 15 2(i 22 21 21 30 20 19 12 23 20 18 328 28 Y 8_______________bull 23 23 18 281 23 28

1

22 30 19 19 15 22 19 19 5 17 7 ________________1 23 24 12 25 23 27 24 30 20 19 17 23 22 19 3 14 ~



I 1 I


fill

r




I Starch 1


-----------------------------middotI------~------I------




475 584




NODE


GltOWTfI IIING


HOOT IIANll






lIun











r










I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy









INTEnNol)lo~









r

~~~

~ lt7 ~ I~ tti~~~~~r

HB J~l Ifr






J 1________________1 17 23 16 27 231 28 10 26 18 20 16 23 14 16 -_____ ______ t1 10________________1 191 22 20 2(i f 23 I 26 I 21 27 21 20 16 24 18 17 ______ ______ ~ H_________________ 21 22 15 2(i 22 21 21 30 20 19 12 23 20 18 328 28 Y 8_______________bull 23 23 18 281 23 28

1

22 30 19 19 15 22 19 19 5 17 7 ________________1 23 24 12 25 23 27 24 30 20 19 17 23 22 19 3 14 ~



I 1 I


fill

r




I Starch 1


-----------------------------middotI------~------I------




475 584




NODE


GltOWTfI IIING


HOOT IIANll






lIun











r










I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy





INTEnNol)lo~









r

~~~

~ lt7 ~ I~ tti~~~~~r

HB J~l Ifr






J 1________________1 17 23 16 27 231 28 10 26 18 20 16 23 14 16 -_____ ______ t1 10________________1 191 22 20 2(i f 23 I 26 I 21 27 21 20 16 24 18 17 ______ ______ ~ H_________________ 21 22 15 2(i 22 21 21 30 20 19 12 23 20 18 328 28 Y 8_______________bull 23 23 18 281 23 28

1

22 30 19 19 15 22 19 19 5 17 7 ________________1 23 24 12 25 23 27 24 30 20 19 17 23 22 19 3 14 ~



I 1 I


fill

r




I Starch 1


-----------------------------middotI------~------I------




475 584




NODE


GltOWTfI IIING


HOOT IIANll






lIun











r










I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


r

~~~

~ lt7 ~ I~ tti~~~~~r

HB J~l Ifr






J 1________________1 17 23 16 27 231 28 10 26 18 20 16 23 14 16 -_____ ______ t1 10________________1 191 22 20 2(i f 23 I 26 I 21 27 21 20 16 24 18 17 ______ ______ ~ H_________________ 21 22 15 2(i 22 21 21 30 20 19 12 23 20 18 328 28 Y 8_______________bull 23 23 18 281 23 28

1

22 30 19 19 15 22 19 19 5 17 7 ________________1 23 24 12 25 23 27 24 30 20 19 17 23 22 19 3 14 ~



I 1 I


fill

r




I Starch 1


-----------------------------middotI------~------I------




475 584




NODE


GltOWTfI IIING


HOOT IIANll






lIun











r










I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy





J 1________________1 17 23 16 27 231 28 10 26 18 20 16 23 14 16 -_____ ______ t1 10________________1 191 22 20 2(i f 23 I 26 I 21 27 21 20 16 24 18 17 ______ ______ ~ H_________________ 21 22 15 2(i 22 21 21 30 20 19 12 23 20 18 328 28 Y 8_______________bull 23 23 18 281 23 28

1

22 30 19 19 15 22 19 19 5 17 7 ________________1 23 24 12 25 23 27 24 30 20 19 17 23 22 19 3 14 ~



I 1 I


fill

r




I Starch 1


-----------------------------middotI------~------I------




475 584




NODE


GltOWTfI IIING


HOOT IIANll






lIun











r










I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy

r




I Starch 1


-----------------------------middotI------~------I------




475 584




NODE


GltOWTfI IIING


HOOT IIANll






lIun











r










I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy





lIun











r










I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


r










I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy







I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy




I ~J l)~


i Ir


1



Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


Groups on sides

Groups on wing

middotmiddot16

middotmiddot26


17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy

17




































19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy










19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy

19

r



LEAF BLADE












L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy





L~F SIIETII


21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy

21

r








JlLAlm JOINT






(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


(shy I shyt~

0


23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy

23

r




DEWLAP








o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


o



r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


r

f-+~~~shy

~ t

c o t ~ ~

---~



771j()7-15-1


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


1





27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy

27

r


ANATOMY OF THE STEM

INT~nNoDE










r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy




r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


r











31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy









31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy

31

r











bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy

bull bull

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy

33

r


GROWTH RING











r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy




r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


r



r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


r

bull8



r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


r



ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


ROOT llANO





NODAL PLATE







r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


r





41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy




41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy

41

r







ANATOMY OF THE LEAF

SHEATII BASE






43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy



43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy

43

r




LEAF SHEATH
















r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy









r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


r

amp














r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy












r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


r






1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy




1)1[1


r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy

r







LAHGE LATEltALS







r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy




r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


r









IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy







IkTTHESS HOOTS






r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


r




A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy


A shy

bull

~ 8 yen~ ~






(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy




(1)

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)

(10)

(11)

(12)

(13)

(14)

(15)

LITERATURE CITED







327-356 illus



















t

---- - shy

t

---- - shy

Documents

Anatomy and Morphology of the Vegetative Organs of Sorghum ... · Technical Bulletin No. 957 • JUlie 1948 .Anatomy and Morphology of the V'egetative . Organs of Sorghum Vulgare