Morphology and anatomy of Polygonum viviparum

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MORPHOLOGY AND ANATOMY 01 POLYGONUM VIYIPARUM by JOHN JEROME SPERRY

A THESIS P r e s e n t e d t o th e F a c u l t y o f The G ra d u a te C o lle g e i n t h e U n i v e r s i t y o f N e b ra sk a I n P a r t i a l f u l f i l l m e n t o f R e q u ire m e n ts F o r th e D eg ree o f D o c to r o f P h ilo s o p h y D e p artm en t o f B o tan y

L in c o ln , N e b ra sk a J u l y , 1942

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UMI Number: DP13951

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MOBPHOLOOT AHD AHATCMT OF POLTOOHUM VIYIPAHOM*

Polygonum T lrip a ru a L, i s of in te r e s t because i t re p ro ­ duces T a g sta tiv a ly by swans of b u lb ils appearing on tb s f l o r a l a x is (fig* 14) .

T his in v e s tig a tio n i s aenearnad

p rim arily w ith datarm ination of tb s n a tu rs of tb s flow er, tb s b u lb il, tb s peduncle, and tb s rhizome.

Tbs s tru o tu ra

of tb a le a v e s and r o o ts i s a lso oonaidarad.

Tbs anatomy of

tb s aaadling i s not d aso rtb ad , f o r ssa d lin g s vara not found i n n atu rs and eould n et ba produead from tb a aebsnas th a t vara e e lle s te d l a t a in tb s growing season* M odification of tba infXorasoansa f o r

to

duotion oeours in many groups of Aagiospenaa*

g e ta tlv e re p ro polygonum

v lv lp a ru a i s tb s only sp ae ia s in tb s Pelygonaoaaa known to raprodues in t b i a manner*

Tbara ars se v e ra l o tb sr sp aeias

i n tb a f a n il y , however, tb a t M ultiply by o tb ar v e g e ta tiv e methods*

According to sa b ro a ta r (1908) polygonum b i s t o r t s

sands out m a ll side branebss from tb s fla s h y , ereokad r h i sons*

At tb a ends of tb asa branebss nav p la n ts o r i s s .

Aaeordiag to & g l (1907-1931) Polygonum a i t s baa e rssp in g atama tb a t produoa ro o ts and shoots a t tb a nodes,

B ailey

and B ailey (1941) s ta te tb a t asexual rep red n etien earn ba brought about by h o r tic u ltu r a l n sth ed s in A trephaxis, Ooeoolo b is , Muahlanbaokia, and Rheum rhapontlcum .

Tba foregoing

^ A t i f i l ablm ^diig& aai i i ndda f4#' abahsmib A ssistance hid' suggestions by Hr* X* H* t a l k e r , under whose su p erv isio n t h i s in v e s tig a tio n was conducted, Tba ra ssa ro b was sup* p o rted In p a rt by a g ra n t from tb s American A ssociation f a r tb a Advancement o f d sien ee , to whoa a p p ra s la tlo n i s ax* p re sse d ,

400948

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8

snaaplaa do not c o n s tltu t* a ooaplst* H a t of a l l polygoaaosous p la n ts

loam, to roprodncs T s g s ta tir s ly .

Polygeana YiTlnarua l a a w idely d is trib u te d alplno sp aelo s.

Htogi (190?«1931) a ta ta a tb a t tb a p la n t occurs gan-

o r a lly througbout tbo bigb aountalna and a r o tle reg io n s of Jttropo and A sia, in clu ding tbo Pyrenees, tbo A lps, tbo JUras, tbo nountalns of soandanaria, Hortb Xngland, amd Scotland*

H a m and Clements (1988) ro p o rt tb a t tbo p la n t

i o ono of tb o eblof spool*s of tbo ounmor aopoot o f tbo p otren tundra o f Hortb imerloa*

sm all (1895) rooorda spool*

mono f r m tbo o u m lto of tb o ib l to Mountains* T his spooleo i s found l a abundanoo in tbo T le la lty o f Windy Point on tbo Gog R ailroad on tbo soutboaot slops of Plkos Peak i n T a ilo r County, Colorado*

I t grows fro n an ap*

proximate a ltltu d o o f 11,000 to 18,000 foot* tbo lower alp ln o and uppor oabalpino sense*

Tblo inoludoo Iro n twenty*

f I t s to ono bundrtd twenty p la n ts ooro found p ar square notor*

Tbo g ro a to r number ooourrod i n tbooo a re a s tb a t bad

boon d is tu rb s ! by anim als o r by tbo dSTolopoont of experi* n o n ta l gardoas* MATXRIALS AMD MXTBOOS Tbo n o to r ia l uosd In tb lo study was oollootod in JUns, JUly, August, and o a rly ssptsnbor In tbo aron o f tbo Windy P oint s ta tio n on Plkos Poak In Colorado*

Moot o f tbo mater*

l a l was fix e d In to rm lln ~ a e e tle « a le e b e l so lu tio n , and tbo ro aaln d o r in Kawashin*s and G ra f's so lu tio n s*

Wbon tbo

Ofmpnont vas a r a ila b le , a i r woo renewed fro n tbo tio a u o s

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bafora f ix a tio n .

Tba m o to rlal v e i aaetlonad a t tbioknaaaaa

Taryiag from 8 to £8 a io ro a a . hand oaotiona vara mada.

Whararar p ra e tie a b la f r a a*

Gross o b aarratio n a vara aada v i t b

tb a a id of a b ln a s a la r d la a a a tln g mleroaoepa. D iffa ra n tia l sta in in g van ap p llad according to tb a kind o f tia a u a .

Tba prooaduraa uaad vara B tldsabain*s iro n -

haamatoxylin and Oranga 0 (C haabarlain, 1998)* lacto -p h an o l Otenaval, 1998)} ta n n ia aoid and iro n eb lo rld a (F o a ta r, 1984)} Flaming*a t r i p l a (Qhambarlaia, 1988)} aad (tenant*a quadrupla (fobanasn, 1940)*

in a d d itio n to tb a abata

aatbada a daabla a ta in of a a fra n in 0 aad f a s t groan and a a fra n in alona vara as^loyad.

Tba aaotiana vara moantad i n

Oanada balaam and l a Iso b at y l a a th a o ry la ta . tm aaaaaaafnl a f f o r ta vara aada to proaota normal g ra v tb o f tb a rblssm a and b u lb ils in tb a graanhonaa and la b o ra to ry a t L in o eln , Habraafca.

Thaaa vara p lan tad i n o rd in ary sand,

q u arts aand* loam* aad n ativ a tnndra s o i l .

Daring a pariod

o f a ig b t aan tb a tb a rbisomaa prodnoad a nnslman of tb ra a la a v a s , a f t a r vbiab g ra v tb vaa ratardad*

B o lb ila , ondar

a i a i l a r c o n d itio n s, s ta r ta d g rav tb v i tb ln a pariod o f ta n days.

Tbay grav f o r a abort pariod b at diod a f ta r prodaeing

a man mum of tb ra a is a v a s . F u tila attam pta v ara aada to induoa p r o lif o r a tio n by a o ttin g tb a rbisom aa, tb a b o lb ila , tb a laaYoa, and tb a psd n n sla.

Attampta vara alao aada to g araln ata aabanaa

takan from tb a ao a t mature flo v a ra a v a ila h la i n l a t a Avgust and a a rly ssptaO bsr.

Tbaat flo v a ra bad not y at f a ll a n from

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4

tlit in flo rescen ce*

No germ ination v a t obtained* G1KXRAL MORPHOLOGY

Polygonum vlvlparm a I t p eren n ial by te e n s of rhizom es. The rhisome* a r t found from e a s -b a lf t o t a t inch b e lt* th e t o l l su rfao e. shapes*

They a r t always te a to rte d ta d a r t of varying

The o o a to rtio a t a r t probably d at to t i n m echanical

o b stru c tio n s imposed upon th e growing p o in t by th e o e ra a t, by t b t te x tu re o f tb a t o i l , aad by com petition o f f t r t d by adjacent p lan ts*

Rhizomes o f p l a a t t growing i a d t a t t awards

o f aadgaa any bo a t a r ly v o r t i t a l l y s tra ig h t*

They a r t f r o a

4 to 10 am* v id t aad from 6 to 18 n * long*

Tbt ro o t ayotaa

I t d t a t t aad f ib r o a t aad extends l a t e t b t t o l l f a r about tb ra a l a t b t t l a a l l d lre a tle a s *

from tb ra a to fiv e o r a tt to

la a a a o la tt la a t a a a r t produatd from t b t rhizome t a t b s ta te s* Tbt lo a v tt a r t a l t o r a a t t aad i a too ranks*

Tbt iatam o d aa

a r t about 2 ma. l a length* Oat to tb ro o flo w tr s ta lk s a r t p r td a ttd by a p lan t t a t b season*

Attached t o t b t p td u a a lt a r t from tb raa to f l v t

o v ate, la n c e o la te , o r l in e a r le a v e s (fig * 29) a rra a g td a l ­ te r n a te ly l a te e ra n k s.

AH lo av es a r t so arranged tb a t

t b s l r a p ite s a r t a t about tb s same le v e l ( f ig . 2 9 ).

Tbt

la a v a s of t b t pedunola o fte n bare t b t g re a te r p o rtio n o f t b s l r p e tio le s m odified f o r elaap in g (fig s* 29,24)* vbereas tb s lo av es o f t b t rb lz o n t bate only a s t a l l b a sa l p o rtio n th u s modified* Tbs in f lo re so ta o t o o a ta ia s f lo v e rs o r b u lb ils o r a to n b la a tle a o f t b t tee*

There s to a t t o be no o rd e rly arrange*

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meat of tii* f l o w s 1m r e la ti o n to b v lb lli*

The b u lb il* may

bo ia to rsp e ra e d throughout th e in flo resc en ce but u s u a lly occupy tb s proximal p o rtio n .

Counts of b u lb ils sad flow er*

a s th ey appoar on tb s in f lo ro sooneo show tb s t tb sro nay bo on a given a x is as many a s SO flow ers and no b u lb ils o r tbo ra rsrso .

A ll ia te rg ra d e * of tb sso co n d itio n s e x ist*

Tbs fla v o r s aro p e d ie e le d , aad tb o b u lb ils aro a tta c h e d by abort s t a l k s .

Gao o r tv s flo v o rs 9 o cc asio n ally th re e , o r

ono b u lb il, appoar a t a nods. tended by s im ila r b ra c ts ,

B u lb ils aad f la v o r s aro sub*

o o o asio aally tbo b ra c ts subtoad*

la g b u lb ils a t tb s bass of tb s apiks develop o b lo ro p b y ll. Tbs f i r s t b u lb ils m y roach m atu rity a s o a rly a s tbo l a t t a r p a r t of Jfeas.

T his i s p rio r to a a tb o a is ( f i g . i d ) .

Tbo fla v o rin g period extends fro n o arly JUly to add b ar.

9 spt.cn-

Vhilo b u lb ils aro u s u a lly dissem inated by mid s u m s r,

tb s y may bo formed throughout tb s gresriag season.

Numerous

b u lb ils ro e o n tly o s ta b iisb s d i a tb o s o i l aad l a v ario u s sta g e s of dovolopm at voro observed from o a rly Ib ly to o a rly September.

Tbs in flo re sc e n c e s develop a t d if f e r e n t r a te s

under varying environm ental c o n d itio n s.

Tbo flo v e rs are

white when th e y f i r s t bloom, but l a t e r become flooded v i t b a re d d ish e e lo r. One specimen of Polygonum viviparum was found tb a t pro­ duced a slen d er rkU ens tb a t grew to a d ista n c e of about 16 cm, from th e p aren t p la n t aad a t i t s t i p produced b u lb ils . T his c o n d itio n th e w r ite r found acumen i n Polygonum b i s t o r t s

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6

TBS fXOVBt Bach flow er l a o t i l t p i l by an eneheathlng te n e t ( f i g , 1} which l a two to fo u r e e l l a i n th ic k n e ss a t i t a base and th in n e r above* th i n o u tlo le*

Tba o u te r la y e r of a a lla l a covered w ith a O ccasionally i n t e r e e ll u l a r a i r apacea are

p resen t In I t a neaophyll*

The aheath l a devoid of v a sc u la r

supply o r any m echanical tls e u e .

Zt nay be adnata to th e

p e d ic e l f o r a abort d istan ce l a th e b ae el region* The p ed icel ranges from 9 to 4 an* In len g th aad la about 300 a le ro n a in diameter*

There l a a eenepleoetts rid g e

(fig * 11) a t th e d l a ta l end Ju st below th e veeeptaele* Between t h i s ridge (fig * 19p) aad th e re c e p ta c le (fig * 19R) th e re l e a band of abaelete.cn tis s u e abent 36 a le re a e wide c o n a la tla g o f th re e t o fiv e la y e ra of e e lla ( f i g s . H A , 19a )*

Theac e e ll a have denae protoplasm and la rg e nuclei*

The bend i a In te rru p te d only by th e paaaage of th e v ae ealar bundles whoae nylon l a t h i s la y e r l a reduced to traehelde* The o o rten o f th e p ed ice l e o n te e ta o f two o r th re e la y e ra o f parenchynatone e e l l a , aone of which tahe a v ery deep atain*

According to Johansen (1940) th e eeeurrenee of auch

deeply a ta ln la g c e l l s In d ic a te a th e prceenee of phlobephene which l a a re a u lta n t of th e o x id atio n of tannine*

so lc re d e r

(1906) has s ta te d th a t tan n in e eeemenly occur in polygonae* sous p laate*

The e e l l a t e r a l bundles In tha b asal p o rtio n

a rc earrennded by a aelerenehymatena aheath, which In the upper p o rtio n re a a ln e parenehymatoua.

The p ith l e amall*

dlatinguiC hable endodermie end p e ri cycle do net eaclct*

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A

The re c e p ta c le i t concert ta d pa mm* g ra d u a lly l a t a th e ta p a la of th e perigyneua fla v o r ( f i g . U ) .

fo r th is

raaaoa th e flo v e r i s o fte n re fe rr e d to aa hypogynoua*

Tba

ex ten t of tbo sinuous epiderm al c a l l a which cower tb a ta p a la aad s e t tb a ra c a p ta o la may ae rre t a d a l i a i t tb a two ra g io a a . Tba v ascu lar bundles i a tb a re e e p ta e le converge t a f a r * a cy lin d er ( f i g s . 8-0,11) from which tb a divergence of trao a a occurs*

Tba v asc u la r tlaau a l a eaapoaad la rg e ly o f

tra e b a ld a aad conducting paraaabyma.

Tba parenchyma outaida

o f tb a v a sc u la r c y lin d e r l a lo e e c ly arranged* w ith numerous i n t e r c e ll u l a r apaaea.

Many of tb a a o lla c o n tain druses aad

tannin* Tba ta p a la ara q u incuncialiy arranged (fig* 9)* a re d is tin c tly p atalo ld*

They

T heir aaaopbyll la poorly d o tal*

oped, c o n sistin g o f bat few la y a ra o f apengy parenchymatous ao lla*

Tba apidarm ia i a eowarad w ith a th ic k a a tlo lo aad i a

devoid of trichom as*

There a ra only a fow stom ata sur­

rounded by o a la rla a a guard c e llo .

Bach te p a l i a aappliod by

oaa median tra c e v l t b tv a l a t a r a l a branching from It* A c i r c le of se c re to ry tla a u a lin o a tb a reo ep tao u lar tup a t tb a baaa of tb a atamans*

T his tlaaua* although form­

in g a continuous ring* conai a t s of d l a ta l p o rtio n s tb a t ara d i a t i a a t n ec ta ry lo baa (flga* 1111,18) •

Tba number of lo b es

v aries* but th a ra a ra alv ay a tb ra a larg o ones which aro ad­ n ata to tb a tb ra a ia n a r atmaana (flga* 11*18)*

Thara ara

m aaller ir r e g u la r lobaa to which tb a o u te r atamena may or may not bo attached*

Tba t o t a l number of lo b aa l a not

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c o n sta n t, but f i t * are u s u a lly p resen t (fig* 1 8 ). The n a e ta ris e (fig* 18} a ra composed of a mass o f u n i­ form a o lla , r i c h i a p ro to p la m aad w ith largo nuclei*

They

aro surrounded by a sia g lo la y e r o f s u tla ls e d o p id o rsal e o lla which aro non~aeeretery*

Widely sc a tte re d sto n ata

aro found oa the epiderm la ( f ig , i f ) ,

th e stomata appear

to bo oororod w ith out in aad probably do not fuaotioa* Baaed upea a n a to a ie a l oridonee, tbo roloaao o f n o o tar begins l a tbo baaal p o rtio n of tbo n o etary .

Tbo e o lla i a

t b i a re g io n , la e lu d la g tbo epiderm is, aro f i r a t t o loao t h a i r doaao oontonta*

Thoao e o lla brook down, fo rs ia g a

lyaigoaoua c a v ity ( f i g . I f ) ,

T bia breaking down proooaa

oo atiaao a progrooaiwoly from tba b aaal to tbo upper p o rtio n o f th e noetary*

Tba nootar l a th a a re le a se d l a t e tbo eup-

l i k e groove a t tb a baee of tbo ovary* Tba aadrooelua o o aalata n o z sa lly o f o lg b t f e r t i l e sta* none arranged l a two w herla,

Tba th re e la s e r at erneas aro

p a r tly adnata to tb a la rg o noetary lo b e s (fig s* i s , 18) aad aro opposite tb a in n er ta p a la ( f i g , 9,T4,T8) aad tbo in n e r and o u te r te p a l ( f i g , 9,T8$,

Tbo f iv e e a te r stamens s a y bo

atta c h e d to tbo w a l l e r n o etary lo b e s o r d iro o tly to tbo re e e p ta e le (fig e * ie ,ll£ } *

They aro a lte r n a te w ith tb a

t a p a l a .( f i g . 9 ). Tba a n th e rs o f the th ro e in n er stamens aro e x tre m e , whereas tb a a n th e rs o f th e fiv e o u te r stamens aro in ty e n s s , AH of tb o an th em aro v e r s a t i l e .

The w all of th e an th er

c o n s is ts of th ro e la y e r s o f s e l l s , th e epiderm is, tbo

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endotheciua, aad th e ta p e tu a ,

Tba l a t e r a l v a i l s a t tb a e p i­

dermal c s l l a ara th ick en ed , and tb a s e l l s are f i l l e d v l t b tannin*

The tap e turn and endotheoium d is in te g ra te a t tb a

tim e o f dehiscence.

Tba p o lle n g ra in s are spheroidal and

range from 98 to 48 miorons i n diameter*

in g en eral th ey

f i t tb a h asie form deseribed by lodaboase (1995) f o r tb s Polygonaceae.

They are trio o lp a te v l t b no eoneplenene

in te ro o lp e r th ic k e n in g s. The gynoeelum l e paraoarpous (Terminology o f V ileon aad Jfest, 1999) v l t b a s ty le c o n sistin g of tb re e n e a rly separate s ty le d la ( f i g . 19) • (fig s* 9,18)*

Tbs ovary I s s s s s ile and sb arp ly angled

Tbo ovary v a i l c o n s is ts of a h ig h ly m odified

e a te r epiderm is, a eonepleaeae hy p ed erals, a fev la y e r s o f parenchymatous s e l l s , and a marked in n er epiderm is.

Tbs

o u te r epiderm al s e l l s are p rism atic v l t b t b s l r r a d ia l v e r t i ­ c a l axes elo n g ated .

The parenchyma i s most ex tensive a t tb s

base and in th e an g les o f tb s ovary.

Iben th e ovary I s

mature an exten siv e ca v ity i s sehisolyslgenously formed betveen the hypode m i s and th e in n er epiderm is which rem ains about tbo o v u le.

The ovary i s supplied by tb re e d o rs a l,

c o l l a t e r a l v a sc u la r bundles ( f i g s . f ,8 ,9 ,1 1 ). Tbs s ty le d la a re filam entous and o f r e la tiv e ly uniform diam eter.

Kaoh i s supplied by a s in g le , weakly developed,

c o lla te r a l bundle tb a t i s lo c a te d on tb s ab a x ia l side of the styledlum ( f i g . 15) •

The xylem elem ents o f th e bundle

are rin g ed and s p ir a l v e s s e ls ,

asperated from th e epiderm is

by two o r th re e la y e r s of parenchyma i s a compact fa s e ie le

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10

o f tra n s m ittin g tis s u e (fig s* 1 0 ,1 5 ).

The c o lls o f till*

tis s u e a r t thin * w alled , g re a tly elongated, have dense pro* toplaam , and e x h ib it a d ea rth of vacuole a.

The e o lla aro

r e la ti v e l y n a i l in tra n s e c tio n , b at aro not n e c e s s a rily n a l l o r than tho adjacent parenchymatous o o lle fro n which th e y aro d lffo ro n tla to d by th e proaonoo of t h e i r donao eon* to u t (flg a* 1 0 ,1 5 ). o f th e stylodium .

T his fa a a lo lo ra d la to a a t both aada In th e baaal p o rtio n tho tra n sm ittin g

tla a a o o f a l l tho a ty lo d la of tho a ty lo u n ite and extend fls g o rllk n in to tho loeulo over tho n lo ro p y le, and In some ease a almost oono In o ontaet w ith tho Inner integum ents and tho n u o ellu a.

Thus above tho apex o f tho ovule l a le*

oatod a oolld canopy of g lan d u lar tra n sm ittin g tla a a o . Tho stig m atie head i a e a p lta to (fig* 10) and l a s lig h tl y lo n g er th an broad.

Tho e o lla o f tho o u ter la y e r o f tho

atlg m atle head aro rounded b at not p a p ille e e . Tho orthoptropoua ©rule ( f i g . 11) occupies n eat of tho space of tho looulo aad appears to bo almost s e s s i l e .

Al­

though th e fu n lo le appears to bo much shortened, i t la apparent t h a t i t extends to tho baao o f tho ovary.

I t la

adnato w ith the tla a a o of th e ovary v a i l In tho lo v e r one* th i r d .

I t l a separated fro n tho ovary v a i l by a tannin*

bearing parenchymatous sheath.

The o u te r integument i n the

o h a la sa l reg io n l a th re e e e U e i n th ic k n e s s.

Those gradn*

a l l y ta p e r to f o r a an integument tv e e o lla v id e .

The Inner

integument l a th ro e c o l l s In th ic k n e ss a t th e e h a la z a l end and fo u r to six e o lla vide a t tho n lero p y le .where I t l a

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

tru n c a te .

The n u c e llu s l a fla sk -sh a p e d .

Tha baaal p o rtio n

i a lim ite d by a la y e r of heavy-walled e a ll e .

Tba ta p e rin g

upper p o rtio n protrude a between tb a in n e r integum ents, ex­ tending beyond tb e a toward tb a tra n sm ittin g tia a u e ( f ig . 1 1 ).

A s im ila r eo n d itio n i a found in Polygenes p s r a ic a r la .

according to Socages (1919) •

The embryo eae i n i t a d e f in i-

t i r e a ta te eo n tain a tb a n o rsa l e ig b t m ielel ( f i g . U ) . Ho n a tu re aead ware obtained during tb a p eriod tb a t e o lle e tio n a ware aada*

However, tb a s a t a r i a l e o lle e te d in

e a r ly September oontained embryos wbiab bad reached about tb a 1 0 -e s ll s ta g e , ovary and ovule*

sev eral ebaages bad oeourred i a tb a Moat of tb a parenchymatous e e ll a o f tb a

ovary bad d is in te g ra te d , leav in g only tb a bypoderaia, tba in n e r epiderm is, aad tba sbeatb around tba v ase u la r tis s u e below tb a ovule*

Only tba o u te r integument p e rsiste d *

There

i a foxaad a t t b i a t i n e a scanty supply of endosperm. Tba V ascular Anatomy of tb £ n e w e r Two v a sc u la r bundles e n te r tb a p ed ice l from tb a ra o h is (fig * n r ) .

One of tba as im mediately s p l i t s in to two ( f ig .

1 ) , and tb a second a le e d iv id e s , forming fo u r bundles by tbs tim e tb a a b sc issio n la y e r i s reaebed ( f ig . Sa ).

in tb a re *

eep taele about midway between tb a a b sc issio n la y e r and tb a base of tb a ovary tb a bundles u n ite by tb a form ation of tra n sv e rse stra n d s c o n sistin g m ostly of tra e b e ld s and con­ ducting parenchyma to form a v a sc u la r c e ll a r ( f ig s . 3 ,1 1 ). l a t b i a reg io n th e re i s a r e la tiv e ly larg e p i t h containing parenchyma and eclcrenabyma.

400948

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Tba tra o a a to t b t ta p a la a r t f i r a t t a a ria a (fig* 8, T l-5) • nannar*

Tbt a t do not a r i a t aim oltantoualy but In a s p ir a l Tbt t r a t t a may ba nux&barad in o rd tr of t h a i r o rig in

a t i a in d ie a ta d by f i g a r t a 3 , 9 T1-T5.

Tbt tra o a a to tba

o u tar ta p a la a r i a t f lr a t* tboaa to tba la a a r and o u ta r ta p a l aaoond, to tb a in n ar ta p a la la a t*

At a a llg b t diatanoa

abort tb a p o in t of o rig in of ta p a l t;;o « 5t tba tra o a a to tb a o u ttr atamona a ra fozntd (flga* 4*08} 11 l a f t )• a lto a r i a t a p ir a lly .

Tboaa

Apparantly tb a tra o a a batvaan ta p a l

tra o a a 1 aad 3 aad batvaan ta p a l tra o a a 3 and 4 appoar f l r a t*

Tba o tb a r tra o a a to tb a o a ta r atsaana a r la a batvaan

tb a tra o a a to ta p a la 1 and 4 , ta p a la 2 and 5 , and ta p a la 3 and 5* Tbo in n a r otaaana a r i a t a la o a t a im lta n a o u s ly b at not a t tb o aaao la v a l (flga* 8 t JSj 118)*

Tba tra o a to tb a

in n a r a t t a i n tb a t a r ia a a i a a p o a ltle n near ta p a l 3 appaara f l r a t*

Tbo tra o a a to tb a o tb a r tv o in n a r a to a n a a r la a

oppeaita tb a tra o a a to ta p a la 4 and 3 (fig* 3,1s)* Oaoaalonal v a a tig la l tra o a a hava boon obaorrad a t a la v a l batvaan tboaa o f tb a Innar and o u ta r ataaana* Tba d o raal tra o a a of tba ovary a r la a in tb a ra eap tao la a abort diatanoa balev tb a ovary (flg a* 3D* 11)*

Up to

t b i a t i a a tba v aao u lar o y lln d ar p rag raaalv aly daaraaaaa l a d la n a ta r v l t b a eonaa quant d ia l nut io n o f tb a p ltb a ra a (flga* 3 ,4 ,5 * 6 ).

Tba doraal tra o a a a rla a approxlm ataly

oppoaita ta p a l tra o a 1« ta p a l tra o a £ , and an o n tar atanan tra o a (flga* 6 ,9 ).

Bafora a gap l a forsaad, tvo a a a ll tra o a a

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appear l a t e r a l to each doraal tra c e ( f ig . 6V)•

These do not

pass in to th e w all of th e ovary hut pass upward and fuse w ith th e vaaoular supply to th e ovule (fig* 7)•

The d o ra al

tr a c e s pass d ir e s tly in to th e stylodia* The vaaoular supply to the ovule c o n s is ts o f a o iro le of s o lid vaaoular tis s u e whose a re a i s much deer eased (fig s* 8 ,9 ,1 1 ).

oust below th e heavy la y e r o f eh a la z a l

c e l l s t h i s s o lid tr a c e d iv id e s in to th re e , whieh are oppo­ s i t e th e d o rsa l traces*

These th re e p ass outward and up­

ward around th e b asal p o rtio n of th e ovule, a t which tin e l a t e r a l tra c h e ld s diverge to fo n a a cup (fig* 11) * A n a i l amount of v a sc u la r tis s u e p asses from t h i s in to th e ehalaza f o r a sh o rt distance* TBS BULBIL The b u lb il o rig in a te s as a d e riv a tiv e frost th e a p le a l maristem o f th e peduncle*

I t b eg in s a s a p a p illa te protub­

erance in th e e a rly autunn*

By e a r ly Jhne th e flow er s ta lk

i s about 3 cm* in le n g th , and th e b u lb ils are re a d ily d is ­ tin g u ish ab le*

The mature b u lb il (fig * £9) I s a sm all, p i r i ­

form s tr u c tu r e , ranging from 9 to 6 am* along th e c e n tr a l a x is and from one-h alf to 9 mm* a t th e w idest point of th e b asal portion*

Sice v a r ia tio n s a re ap p aren tly due to d i f f e r ­

en t environmental eonditlons* The b u lb il

c o n s is ts of a sh o rt stem , en larg ed a t tho

base (fig s* 9 8 ,2 9 ,9 0 ,3 9 ), to which i s attach ed two ao alell k e leav e s (fig* 88L).

The oerea of the o u te r com pletely

covers the e n tir e in n e r one a s w ell a s the 3 o r 4 embryonic

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leaves*

They are a lte r n a te aad arranged i a tv o ranks*

The

oerea o f tho tv o b aaal loaves sorvo p rim a rily a s a p ro te c ­ tiv e sheath (figs* 86,36), A sh allo v depreseion i s lo c a te d a t th e bass of t h i s atom marking tb a p o in t of severance fro n tho parent plant* P rotruding s lig h tly from t h i s d epression are remnants of th e b u lb il stalk *

T h is s ta lk c o n s is ts of the tvo c e n tr a l

stra n d s o f v asc u la r tis s u e (fig* 33) aad tvo o r th re e la y e r s of parenchymatous s e l l s betveen th e vaaoular bundles aad th e epidermis*

l a th e lovexmost p o rtio n of th e s ta lk

an abac le s io n reg io n

00 ours*

T his c o n s is ts o f se v e ra l

la y e rs of elongated* th in -v a ile d c e lls * The stem of th e b u lb il i s covered v l t h an epiderm is vhleh co n tain s no conspicuous in c lu sio n s (fig* 86) * p e r i­ derm form ation ta k e s place i a th e hypedermls* o f tvo to fo u r lay ers* u su ally th re e (fig* 86)* fo u r la y e rs tvo of them are phellem .

I t c o n s is ts When of

The phellsxa e e l l s

become f i l l e d v l t h tan n in vhloh i s a t f i r s t homogeneous (fig* 86) and l a t e r granular*

The v a i l s become more dense

vhieh i s In d ic a tiv e of th e presence of suberln*

In th e

b a sa l p o rtio n o f th e b u lb il vhere the periderm a t t a i n s i t s g re a te s t development th e epiderm is becomes atro p h ied and begins to slough off*

The s o r t l e a l and s te la r re g io n s of

th e b u lb il stem c o n s ist m ostly of storage parenchyma i n t e r ­ spersed v lth a i r spaces, many o f vhieh are extensive* A ll of th e parenchymatous e e l l s contain inclusions* The most common i s s ta re h (fig* 86) a s determined by the

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iodine t e a t ,

T his i s composed of o ra l g ra in s v h ieh are

marked v l t h conspicuous oonosntrle s t r l a t Ions (fig* 25) and, a t m a tu rity , densely f i l l th e s e ll*

The la r g e s t sta re h

g ra in s a re about 10 ailerons long*

S o attered throughout the

starch -co n tain in g parenchyma a re ta n n ln lfa re a s e e U s (fig* 26, h eav ily stip p led )*

The l a t t e r e e l l s may he sin g le o r

in groups of too to te n and fre q u e n tly extend i n lo n g itu d In a l revs*

C olls eo n talnlng d ru ses aro of l e s s frequent

occurrenoe and a re found n ear the ra s e u la r tis s u e s and In the s o r t e r a t th e le a f nodee*

Tho vaaoular handles a re tho

only h ls to lo g le a l s tru c tu re s th a t serve to d e li n l t th e ex­ te n t o f th e e o r ti e a l and s t a l e r re g io n s, an endodermis o r a p s rie y e le not being d istin g u ish ab le* The v aao u lar e y lln d e r i s a d isse e te d slp h o n o stele, tho bundles o f vhieh are e o lla to ra l*

in a d d itio n to tho u su al

vaaoular elem ents of xylem and phloem th o bundles are s u r­ rounded by sev e ral la y e rs of elongated parenehymatous e e ll s v l t h dense protoplam alo c o n te n ts.

There a re no selerenehy-

aato u s e e l l s around th e vaaoular bundles or In any o th e r p o rtio n of th e stem o f the b u lb il* At the apex of the stem of th e b u lb il th e re I s a m a i l , c o n io a l, m u ltl- e e llu la r growing point*

A c r i t i c a l study o f

t h i s term inus has net been mads, but m orphological zonatlon of th e a p ic a l n e r ls te n earn be eeen* regions*

There are tvo d is tin c t

The o u te r reg io n c o n s is ts of a single la y e r o f

e e l l s vhieh divide only in an a n tic lin a l p lan e, aad i s e v i­ d en tly th e p recu rso r o f th e spiderede*

T h is germ la y e r vould

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16

be considered a t th e deraatogen of B uaatein (1666) o r th e tu n ic s o f Schmidt (1984)*

The in n a r g am la y e r s a m of

homogeneous s tru c tu re and divide in a n t i c l i n a l , p e r l e l l n a l , and tra n sv e rse plana a,

These e a l l s give r la a to tb a i n t e r -

n a l tla a u a a of tba a tm aad leaves*

They can n o t, however,

ba d irid a d on tb a b a s is of appaaranoa aloaa In to tb a p a ri* blem and pierone o f b&nsteln (1863) but r a t bar sa y ba dla~ tin g u iab ad aa tba corpus o f Schmidt (1924) . d if f e r e n tia tio n l a abort*

Tba re g io n of

Parenchymatous a a ll a containing

a tarcb and ta n n in are f l r a t observed a t a diatanoa of about 78 microns p o s te rio r to tba growing p o in t, and tb a fe ra a tlo n o f v a sc u la r tla a u a from p rev aseu lar tla a u a a t about t i n same lev el*

Tba a i r apaoaa f l r a t oeeur about

800 o lerona from tbo apex* Tba o u ta r la a r a a o f tba b u lb il a t f l r a t bare sm all bin da a (flga* £61, 29,80) and ao n alat m ostly o f o o raal t i n ­ eas*

Tbo p e tio le a are abort and tb a laminae a ra but

s lig h tly developed (fig* 8 2 ).

In tb a lamina of tb a f l r a t

l e a f th e re la a m a l l amount of p a lisa d e and spongy para a obyma* L a te r tba blade may expand to a g re a te r o r la a a ex tan t (fig* 36)*

saob successive le a f develops to a

g re a te r ex tan t u n t i l tba normal le a f l a attain ed * Loaf development i a continuous w ith tb a exception of a p ariod of enforced dormancy tb a t tb a b u lb il may undergo* Tba f l r a t evidence of a le a f l a tba presence of a s lig h t rid g e (fig * 88) around tb a v e g e ta tiv e cone (fig* 52a ) , consequent growth o f tb a a e r ls ta a a tlo rid g e l a uniform

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Tba

u n t i l th e growing p o in t of tbo atom ha a boon surpassed. S h o rtly a f t a r t b i a tb a ra i a an in araaaa i n tb a r a ta o f development on ana aide o f tb a r id g e .

Tbia a a a a la ra tia n

marks tba baginnlng of tb a form ation o f tba p a tio la aad lam ina ( f i g s .331,341) •

Tbia growth i a divergent f r a * tb a

remainder of tb a r id g a .

At t b i a time tb a a p io a l m e riste a

i a com pletely oowarad w ith tb a exception of tb a a a a ll o r i fio a i n tba aabryonie ooraa ( f ig . 35 OB) •

Due to tba to n -

daaoy fo r a more ra p id growth on tb a aida of lam ina forma* ti o n , tba o r if ic e o fta n eoaaa t o bo lo a a ta d a t an anglo ra tb a r than s tra ig h t ab o rt tba growing point o f tb a atom ( f i g . 34 OB).

Tba ambryonio ooraa continue a i t a growth

p rim a rily by a n t i e l i n a l and tra n sv e rse d iria io n a .

I t ia

evident tb a t few p e r le lln a l d iria io n a ooour baaanaa tb a mature ooraa l a tb a aaaa th ick n ess i n any given region aa tb a aabryonie o o raa.

About tba time tb a t tb a prim ordla o f

tb a lam ina aad ooraa bare baooma d if f e r e n tia te d , tb a vega* ta tiv a p o in t forma another rid g a . When tb a lamina l a about primordium 400 alero n a lo n g , an other la a fAhaa formed an tba opposite aide o f tba aada ( f i g . 3 4 ). Thua i t can ba aeon tb a t tb a ra l a a rhythmic a c ti v i t y o oeurriag b a re .

Tba sequence i n aunmary i a aa follow at

(a ) Tba form ation of a protuberance surrounding tb a a p ic a l marl atom | (b) tb a aurpaaaing o f tba meriatam by tb ia rid g a ; (a) tb a d if f e r e n tia tio n o f tba lo a f and p a tio la p rim ordla, follow ed by (d) form ation of an a d d itio n a l rid g a of mer i ­ ot am in tba a p ic a l growing p o in t, ag ain tb a aneompaaaing

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18

o f the growing p o in t and tba i n i t i a t i o n of a lam ina aad p a tio la primordlnm on tb a ©ppoaita aida o f tb a axis* A d re n titio u s ro o t a i n p ro g re a e ite atagaa o f development may ba found in tb a o o rtlo a l rag io n of tb a b u lb il. prim ordia (flga*

2 0 , 2 ?)

Tba ro o t

a ra formad i n tb a p e rie y e lle ragion

ad jacen t to tba p ro ta a e u la r tla a u a .

ITa n a lly tb ra a ro o t a

(flg a* 27,30,51) a r la a a t aaab ra a o u la r sodas o cc asio n ally tb a ra a ra b at ono or two*

Tba ro o t a a ra always ad jaeen t to

tba tra n s re rs e ra a o u la r atr&nda, and a t aaab noda tb a ro o ts a r la a i n about tb a seas r s l a t l r s p o s itio n (fig* 51}*

Tba

ro o ts divide tba s lrs u n fs rs n e s o f tb a atom in to tb ra a n a a rly equal a ro s , b at wbsn only two ro o ts o rig in a te a t a noda tb a a r t s ara unequal, tba p o s itio n o f tb a two being tb a asms a s tb ay would ba I f tb ra a ware p re se n t.

Tba ro o t f i r s t pasaas

upward through tb a e o rte x , and a t t b i a time a la rg e ro o t oap l a formed*

By tb a tin s i t baa r e ashed tb a periderm ,

th e proximal p o rtio n h a s besoms d if f e r e n tia te d in to tba prim ary tis s u e a o f tba root*

Tba t i p ru p tu re s tb a p a rld e m

and paaaaa out in to tb a a x il of tb a ooraa a t tb a le a f node* Tba ro o t may p a n e tra te tba ooraa (flga* 29,30) o r p ass out between tb a ooraa and tb a stem* Vaaoular Anatomy of $!» B u lb il Two tra o a a e n te r tba b u lb il from tba ra o b ls (fig* H J).

21

Tboaa d lta rg e a t ones and p a ss outward and upward

(fig* 35) • These ara tba prim ary traoaa*

These ta o e u la r

tra o a a soon branch and anastomose forming a c y lin d r ic a l c o lla r (figs* 31 VN,35).

In t h i s tb a t a s s e ls c h a r a c te r is tic

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o f tlit bundlea aro replaced by many abort h o riz o n ta l tra e h e ld s , and th a ra t a aa laoraaaa l a tb a amount of con­ ducting parenchyma,

Tbia ra g io n t a tb a ra a o u la r noda ( f i g .

51 TO) aa d latin g u lah ed from tb a rag io n of a tta e ta u n t of tb a la a r e e , tb a lo a f noda ( f ig . 31 UT). Sight tra o a a paaa from tb a f l r a t ra a o u la r node, too of vhioh ara a co n tin u atio n o f tb a prim ary tra o a a tb a t a a ta ra d tba b u lb il*

ir o n an a-h alf to tv o -th ird a tb a diatanoa bo-

tra a n tb a f t r a t and tb a aaoend ra a o u la r noda a, tb a ra paaaaa from ono o f tb a prim ary ra a o u la r atran d a a larg o tr a o a . Thla paaaaa in to tbo p a tio la r rag io n o f tb a f i r at l a a f (fig* 31 LN) * F requently tba la rg a ra a o u la r tra o a on tb a oppoaita aida o f tb a b u lb il o p lita and anaatomoaaa v l tb an ad jacen t tra o a ( f ig . 3 1 ).

Tvo of tb a romaining tam o a vhieh

a ra appraxlm ataly a q u id ia ta n t from tb a prim ary tra o a a paaa in to tb a o a ra a l ra g io n o f tb a f l r a t l a a f noda*

Tba aaeond

ra a o u la r noda i a a abort diatanoa abora tb a f l r a t lo a f noda, tb a la a f noda a i n aro ry eaaa balag from tv o -th ird a to tb r a a fo u rtb a tba diatanoa batvaan tb a ra a o u la r nodoa. Tba form ation of tb a o o lla r a t tb a aaeond ra a o u la r node and a t aaob auoooaalro one l a tb a aama aa a t tb a f l r a t . Tba number o f lo a f tra e e a paaalng from tb a aaoend ra a o u la r noda may ba tb ra a or fo u r, r a r e ly f i r e .

Tba branch from

tb a primary tra o a paaaaa in to tb a p a tio la r re g io n of tbo l o a f , th e o th e r tra e e a paaalng in to tb o o e re a l re g lo n a . O coaaionally one o r tvo m anlier tra o a a paaa in to tba pat* i o l a r reg io n of tho l e a f .

I t l a a t t h l a aaeond ra a o u la r

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20

node th a t tint f i r s t a d v e n titio u s ro o ts o rig in a te f f ig s .

2f,

31 B ).

Prom th s th i r d v ascu lar node tr a c e s pass in to the t h ir d le a f of th e b u lb il,

Thera seems to ha no r u la a s to th e

number of nodes through which a tra c e p a sse s before e n te rin g a lo a f .

Three tr a c e s , a prim ary tra c e aad too l a t e r a l

tr a c e s , always pass in to th e p e tlo la r re g io n , the l a t e r a l tr a c e s being approxim ately e q u id is ta n t from th e prim ary tr a c e .

The ooreal tr a c e s range in cumber from t i e to many.

T heir p o s itio n has no re g u la r re la tio n s h ip to th e p e tlo la r tr a c e s .

However, in th e f i r s t le a v e s th e re a re u su a lly t i e

la rg e o o re al tra c e s i n a l a t e r a l , median p o s itio n .

These

soon fo rk ( f i g . 87 L ). There e x is ts a m u l l amount of ln te ra o d a l s p l i t t i n g and anastomosing o f t r a c e s .

A v a sc u la r tra c e may s p l i t to

surround a m u ll Is la n d o f parenchymatous c e l l s which i s two or th re e c e l l s in th ic k n e ss and may extend f o r a die* tanee o f about 200 m icrons.

In d iv id u al tr a c e s may s p l i t a

sh o rt d istan ce below th e point o f form ation o f th e v a sc u la r node.

Thus o ccasio n ally in a sin g le c ro ss se c tio n a l view ,

the v a sc u la r c o lla r may appear s lig h tly in te rru p te d , whereas i t i s probably continuous but I r r e g u la r i n so f a r a s a s tr a i g h t , h o riz o n ta l p o s itio n i s concerned. THE FXSXm&S The peduncle c o n s is ts of a heavy-walled epiderm is, a narrow c o rte x , a s i phono s te le of c o l l a t e r a l bundles, and a la rg e p i t h .

The outerm ost la y e rs of c e l l s of the c o rte x

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21

are of collenchyma.

There i s a conspicuous endodenais

whose c e l l s are somewhat enlarged and co n tain s ta rc h .

The

p crio y ele c o n s is ts o f several la y e rs o f elongated s e le re n chyaatoua c e lls *

The v asc u la r handles are th e most highly

developed of any found In the plan t*

The la rg e r ones con­

t a i n , in a d d itio n to th e primary t is s u e s , a w ell organised cambium and a meall amount o f secondary xylcm and phloem* The l e s s mature handles co n sist p rim a rily of phloem w ith a few stra n d s o f protoxylem .

in th e I n te r nodal reg io n the

handles branch and anastomose fre e ly *

There a re many uuoh-

elongated ta n n la ife ro u a c e l l s i n the p ith and th e cortex* The druse-co n tain in g c o lic w hile s c a tte re d through th s stem a re most numerous In th e nodal region*

At the nodes the

v a sc u la r handles anastomose, forming a c y lin d ric a l c o l l a r , numerous bundles of varying number pass in to th e leaf*

The

gaps are sh o rt and are closed by l a t e r a l tra c e s from a d ja ­ cen t bundles. In th e ra o h ls th e re i s a m anlier amount of colleeehyma and selerenehyma w ith an in crease in th e amount of d ru seco n tain in g and ta n n in ife re u s parenchyma*

The eadodermls

rem ains conspicuous but i s broken a t the numerous nodes* The course of th e v a sc u la r system has been follow ed to th e f i f t h node.

About 250 microns below the f i r s t node

two adjacent tr a c e s anastomose and continue to th e nodal region*

Biro a branch anastomoses w ith an adjacent tra e e

(fig * 21)*

At a p o in t about 50 to 100 microns above t h i s a

sm all tra c e p asses from th e o r ig in a l tra e e in to th e b ra c t

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

(fig * S I B)«

At ft d istan ce o f 10 to GO microns above th s

p o in t o f d eparture of th e b ract tra c e th e larg o remaining tra o a branches In to to e approxim ately egual ones*

7b*aa

supply a flow er (fig * 21 y) o r a b u lb il (fig* s i BO) * D ire c tly above oaeb nod* u su a lly an adjacent tra o * s p l i t s to f i l l th s gap (fig *

21 ),

so a r tb* apex of the a x is tb * r* la

a d *pl*tlon In tb* number o f trao*a* Then two flow ers appear a t a nod* th e procedure la s lig h tly m odified.

In a rid g e which l a formed below the

p e d ic e l each of th e tra* * a th a t ar* formed above tb* b ra c t tra c e div id e in to two (fig*

21

y y ).

jsach p ed icel th u e r e -

celve a two v aa eu lar tra e e a , th e earn* a s a t a node where bat one p ed icel l a present* THS mny.nifK The anatomy of th e rhinom* l a e s s e n tia lly th e same a s th a t described f o r th e stem of th e b u lb il.

As th e rhizome

in c re a se s In diam eter th ere I s an In crease l a th e number of v asc u la r bundles*

The xylem elem ents c o n s is t almost en­

t i r e l y of traeh eld a*

The v a sc u la r bundles are surrounded

by a parenchim atous sheath* in th e rhizom e.

There a re no f ib e r s to be found

A ll o f th e parenchymatous c e l l s co n tain

conspicuous amounts of e it h e r s ta rc h o r tannin*

Svea those

in th e xylem and phloem reg io n s c o n ta in stareh*

The p e r i­

derm In c re a se s somewhat In thickness*

As many a s th re e

la y e r s of cork c e l l s have bean observed*

l a th e o ld e r r h i ­

zomes th e o u ter one o r two la y e rs of periderm are sloughed o ff.

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n

Roots axis* l a th s rhizcma in th s b u lb il*

asm

manner ms l a th s

At th s nodes branching and anastomosing oeour l a a

Banner comparable to t h a t o f th s b u l b i l .

B ow ver, th s

c y lin d e r i s acre In te rru p te d hseauss of th s passags of a g ra n to r number of bundles to sash appendage. THI LSAT Ths hlada o f th s n atu re lo a f c o n s is ts o f th s epiderm is, oas to

tour la p s r s o f p a llsa d s c a l l s , th ra a o f fo o r la y e rs

o f spongy meaophyil s o i l s , sad th s T asen lar bundles*

Ths

epiderm al s o i l s , w ith th s o x ssp tlo a of thosa over th s aid * r l h , era sinuous*

Ths appsr epiderm is i s covered w ith a

th ic k la y a r of rldgad c u ti c le .

Oa th s lower a p ld a ra is

th a ra i s lo s s o u tic la aad a ra la tlT a ahaadaass of s la g ls* s s l l s d , alongatad trichom as, found h s ra .

a few g lan d u lar h a ir s a r s a ls o

Ths sto a a ta a rs s lig h tly sunkaa aad ara aora

am orous oa th s ab a x la l su rfa c e . c a l.

Ths guard s s U s a rs ty p i­

Ths oh lo ro p h y ll-bearlng parenchyma i s not continuous

aoross th s s a t i r e lo a f hlada hat i s in te rru p te d by th s aid * r i b aad by t i e r s o f parenchim atous s e l l s th a t l i e b etssen th s la r g e r v aa su la r bundles aad th e epiderm is. The Taset&lar bundles r e tic u la te l a th e lamina surrounding s n a il i s l e t s o f parenchyma* l ik e i n stru ctu re *

(fig * £4)

The m idrib i s stem­

Several e e te p h ls le bundles are arranged

i n a ty p ic a l slphoaostele*

As branches pass from th s m idrib

to th s blade th e re i s branching aad anastomosing*

l a th s

blade th e re I s but l i t t l e aslerenshyaatoua t i s s u e , but th s bundles ara surrounded by a sheath o f ta n a in ifs re u s paren*

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

84

chyme* T1m s tru c tu re o f tb s p e tio le i s th e seme o« th a t of th s m idrib v i t b th e ex cep tio n th a t th e re i s considerable M lfN A O h p U it

Tbo s tru c tu re of th s ooroao I s s im ila r to th a t of tb s lo a f w ith sev e ral exceptions*

In tb s b a sa l reg io n th e y aro

r a th s r th ic k , bat so ar tb s margins tb s 7 aro rodaood to tbroo o r f e a r s o i l s in thickness* in g number pass in to tb s ooroao*

T asealar tr a s o s

at T arr­

A ll o f tboso in tb s b asa l

p o rtio n aro p a r ti a l l y surrounded by solerenohyma*

S trands

of seloronshyaa which aro net ad jacen t to tb s r a ocular bandies aro a ls o present*

Tbs number o f tra e o s to tb s

ooroao in o ro aso s in oaeb saooossiTo lo a f from tb s f i r s t lo a f o f tb s b u lb il to tb s mature loaf* TBS BOOT A ll of tb s ro o ts observed bate asso cia te d v i t b them eo to tro p io mycorrbisa*

Tbs anatomy of tb s r o o t, p a rtic u ­

l a r l y v i t b re fere n ce to tb s m o d ificatio n by m yeerrbisa, has boon describ ed by B»coalman (1000) * Supplementary t o Bisoelman*s o b serv atio n s, i t i s to be noted t b a t no ro o t h a ir s a re formed on tb s young ro o ts a s th e y corns from tb s b u lb il*

In th e specimens observed t b s c o r ti c a l c e l l s o f tb s

o ld e r ro o ts do not lo se t h e i r id e n tity a s com pletely a s in ­ d ic a te d by Hssselnan*

Tbs s to le o f tb s ro o ts produced by

tb s b u lb il i s t r i a r c h , vhereas tb a t of tb s r o o ts produced from tb s mature rhlsem s i s peat arch*

Due to th e o f f s e ts of

th e m ycorrbisa tb s c o r tic a l e e l l s aro destroyed aad tb s storage of s ta rc h ta k e s place i n tb s p e ri cycle a n d e n d o d e rm is .

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

25

BXSOJBSXOg

5aX5iX8 aro probably tho c h ie f M ana o f reproduction l a Polygonum Tivjparum . a le s s o r e x te n t.

Reproduction by ooods mar eeeur to

fho f a il u r e to fin d n a tu re aooda map bo dao

to th s fo o t th a t i t mas not poaalblo f o r the observations to bo continued a f t or th e f i r s t week of September. However, th s r ig o r s of v ia to r l a th s o lp lao tu ad re map oaaoo a pro* a a to ro term in atio n of th s dorelopneat o f th s sood aad map ho rssp o n slb ls f o r a high r a t s o f m o rta lity of seedlings I f thop occur.

fho b u lb ils aro prodaood oarXlor th aa tho

flow ers so bosoms ostabXishad l a th s sams soasoa l a vhloh thop aro developed.

B u lb il p ro d a etlo a I s probabXp a e t a

rssuXt of o o a d ltlo a s th a t aro unfavorable f o r sood produc­ tio n u n le ss i t i s th o sh o rtn sss of th o soasoa.

Both b u lb ils

aad fla v o rs ooear oa tho asms p lo a t aad l a about oqual abuadaaoo.

Thsrs i s ao srld sn o s t h a t asoauaX ro p red u etlo a

ooeurs from d s d lf f s r s n tla ts d t i s s u e .

fho b u lb il i s slmplp

a atom branch th a t bosoms s ssTorsd from tho p a rs a t p la a t aad grovs d lr s o tly in to tho rhiaomo o f tho a d u lt. fho aaatom leal srld sao s shows t h a t tho flow sr i s a m odifiod atom, a s la d lo a to d l a g en e ral f o r flo v o rs bp semes and ttaoBanlels (1926), and th a t th s b u lb il i s a ls o a modi* f l s d atom.

Bus to th s dsaorlbod d lffo ro ao o s l a tho nataro

aad number of tho appendages, aad tho v asc u la r system s, i t i s evident th a t eo sv cralea from oas stru o tu ro to tho othor h as not taken plaoo.

50 attem pt

has boon mads to determine

tho fo o te r s vhloh laflu oaoo th o r o la tiv o development of th s

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two stru ctu res* The anatom ical fe a tu re * of th o b u lb il and th e rhizome aro sim ilar*

Tb* eh i* f d iffe re n c e between tb * too at rue*

tu ro a i e size* which l a only * a a tt « r o f age aloe* tb* b u lb i l by continued growth become* transform ed in to tb* rhizome o f tb * ad u lt plant*

Tb* rhizca* baa a g re a te r

storage a r* a t a g re a te r number of c o ll a te r a l v a sc u la r bundles, and a broader periderm .

The form ation of a vaseu*

l a r c o lla r a t tb * node* a* w ell aa tb* o rig in of ro o t* i a tb* same In tb* too*

Tb* f i r at ro o t* f r a a tb* b u lb il a re

t r i a r e b 9 whereaa tb* ro o t* from tb* r h i none a re p«atareb* Tb* q p lra l and rin g ed w eaaela o f t i n b u lb il are rep laeed by tr a e b tld a a a i t develop* in to a rhiscne* Tb* peduncle* aa a temporary a e r i a l stem* ahova a g re a te r number of m o d ification*, and i t d i f f e r s markedly in aeveral r e p e a t * from tb* rhizome and b u lb il*

S ir* th ere

i * m ob meobanieal t i s s u e , whiob l a absent in tb* b u lb il and rbisoa**

Only i n tb* p*dunel* i s tb*r* found pronouneed

cambium and aeeondary nylem*

Tb* only place In tb* peduncle

tb a t sta rc h storage occurs l a l a th e endodermls, whereaa i n tb* rb ise a e and b u lb il i t l a continuous throughout*

The

tan n in e * U s in tb* b u lb il aad rb ise a e are not elongated aa th e y a re in th e peduncle* Tb* presence of multjQLaeunar l e a f nodes* according t o d ln a o tt and B ailey (1914} 1* an advanced condition*

The

m u lti^ a c u n a r supply a t tb * le a f node* can be in te rp re te d by tb * h y p o th esis tb a t tb * oorea* represent* aa s ta te d by

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S in n o tt end B ailey , a rev of fuaed s t i p u le s ,

i n th e B u lb il

tb s supply to tb s f i r s t tv o l e a s t s i s tri^ la c u n a r*

T h is,

aoeording to S innott (1914), i s probably a p rim itiv e condi­ tio n f o r th s o rder Polygonal* s .

The number of tra e e s pass*

*

ing to a le a f range from th re e to many.

Thus on t h i s p la n t

p rim itiv e and advanced e o n d itio n s e x i s t .

T his probably ean

be e o rre la te d w ith th e f a s t s th a t th e oereae

th e n atu re

le a v e s re p re se n t numerous fused s tip u le s aad th a t a g re a te r number of tra e e s pass to th e enlarged laminae of th e mature le a v e s .

Arber (1999) s ta te s i n re sp e ct to th e fla v o r th a t

th e v ascu lar system oorresponda in e x te n t to th e organ vhloh i t su p p lie s .

Applying t h i s statem ent to e th e r ogams

i t i s to be expeeted th a t a re d u ctio n o r in crease in number o f fu sed s tip u le s or i n th e siz e o f th e le a f blade vould be d ir e c tly e o rre la te d v l t h th e number of le a f tr e s e s a t a node. The c o n trib u tio n o ffe re d by th e study of th e f l o r a l anatomy o f t h i s sp ec ie s le n d s emphasis to th e statem ent by laubengayer (199?) th a t " th e Polygonaeeae re p re se n ts a v ery homogeneous and n a tu ra l fam ily a s i s ahsva by th e uniform anatom ical s tru c tu re in th e f lo s e r " .

The f l o r a l anatomy of

Polygonum v lv ln aru n i s s im ila r to th a t o f th e sev eral Species of Polygonum described by him.

The study o f f l o r a l

morphology v lth ln sev e ral o th er fa m ilie s co n trib u ted ad d i­ tio n a l evidence of th e re la tio n s h ip s v i th in a given taxo­ nomic group (Davaon, 193Cf Lawrence, 1937 j Thomson, 1942). The conclusion reached by L&ubengayer (199?) th a t the

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vhorled co n d itio n i s a fundamental one o f th e fam ily Folygon&oeae la not su b sta n tia te d l a ao f a r aa the o rig in o f te p a ls l a eoneerned*

Aa has been d escrib ed , th e te p a ls l a

PoXyeoaott TlYtparum a r l a t l a a s p ir a l manner*

Tim ataaaaa

l a th e same species do ao t a ria a sim ultaneously w ith in e ith e r the o u te r o f la a o r w horl, th u s p resen tin g evidence th a t tho p rim itiv e c o n d itio n was s p i r a l .

The r e s u l t s ob­

ta in e d l a t h i s study would agree aaaa eXoaaXy w ith th ese of Bauer (1922) ohoaa s tu d ie s waro ontogenetic* X&ubengayer (193?) reaches tho aoaaXaaloa th a t perigyny l a u n iv e rs a l l a tho Polygenaeeae.

Saunders (1939) a ta ta a

th a t tho flo w ers l a t h l a fam ily aro hypogynous.

I t l a c r l*

dent th a t Polygonum riv ip a ra n l a perlgyaous# Laubengayer (1937) fooad l a Ooeooloha aad B ranalohla t h a t th o ovule l a supplied hy a la rg o o o n tral mass sur­ rounded hy a t h l a ahoath o f e e lls *

Ba I n te r p r e ts t h l a a a a

■eentrel bundle raparaaoatlag a fa a lo a of tho v a a tr a la aad tho ovular tr a c e s , aad suggests t h a t a a u ltl- o v u la r oondlt l o a o&oo ex isted * Polygomta vlvlnarum*

fho aaaa evidence l a p resen t l a fho ahoath around tho v a sc u la r aapply

to th e ovule l a stro n g ly marked In t h l a apooloa*

T herefore,

l a Polygonum vlvlparem . aa l a o th er opeeloa of tho Pelygonaeoae (Xanbengayer, 1937), th o ovule s ta lk l a r e a lly a fu n lo u lu a r a th e r than a p la c e n ta , aad tho p la e e n ta tle a l a f r e e c e n tra l r a th e r th a n p a r ie ta l aa I t la o rd in a r ily de­ sc rib e d .

Saunders (1939) elate a that sectaries are of te a present

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botwooa th s stamens l a Polygonum* constant fe a tu re o f tho group*

They aro ap p a ren tly a

fho morphology and anatomy

o f tho n e c ta rife ro u s dloo has boon described*

There has

boon aoao d iscu ssio n ao to tho source o f th o Aloe*

flo h o r

(1928) in to rp ro to d tho n o etary In tho v lllo v ao a perianth* Bo Candolle (1825 J* Vpdlor ilflfiO), aad Moero (1956) eoneluded th a t tho aootariforom o dioo l a th o Phaseoleae of tho Legumlnoseae ropro sent o a oot of undeveloped stamens.

la

Polygonum riv ip a ru n thoro aro u s u a lly fiv e lohoo present* and thop aro attaeh o d to a p a rt o r a l l of tho stamens* Thla vould suggest th a t th e dloo re p re s e n ts a whorl of a o d lflo d ataaoao la to ra o d la to botwooa tho la a o r aad tho o ato r whorlo o f ataaoao*

Oeoaolonal v e s ti g i a l trao o o aro

found botwooa tho trao o o loading to th o la a o r and o a to r stamens* b a t thooo aro not of s u f f le le a tly constant ooeurroaoo to w arrant tho ooaolaaloa t h a t thop aro ro a aaato o f tho vaoeular supply to aa la to ra o d la to whorl of otoaoao* SOMUHY Polygonum vivlparum prodaooo aa e re e t a x le f r o a whloh aap bo p r e lo a d flow oro o r b u lb ils o r both*

Tho b u lb ils

develop e a rly , follow ed bp tho floworo* Tho flow er i s pertgpnousf w ith eu iao u n elally arraagod tep als* two whorlo o f stamens* a paraoarpous gyaoeelum* aad la rg e aootarp lo b es whloh aap bo a o d lflo d steaoas* Tho trao o o to tho to p o la a r is e s p ira lly *

Those to tho

o th e r organs aro aore o r lo o s sp ira l* Tho vaooular supply to the ovary o o n s ls ts o f th re e

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d o rsa l t r n t i which p a s t l a t e th e s ty lo d la .

That of th e

ovule l a In te rp re te d aa the r e s a lt o f th e fu sio n o f the v e n tra l tra e e e w ith the o v u lar trac e* The b u lb il l a a m odified stem h earlag f e a r o r f i r e la are a.

The oereae of the f i r a t leave a aerre aa a p ro te c ­

tiv e cover.

The ate* o f the h e lh tl l e covered v lth a

periderm aad c o n tain s v teh feed sto rag e tissu e * l a the h a lh lla th e ro o t a a r le e l a th e perioycH o region a e a r the a p ic a l s e r ia te * . Leaves develop fro * a a e r ie tan a t i c rid g e which ear* rounds th e v e g e ta tiv e p o ia t • The groviag p o ia t l a in te rp re te d aa c o n sistin g of tv e germ la y e r s , a sin g le -la y e re d tu n ic a aad a se v e ra l-la y e re d corpus. The v a e e a la r system o f th e h o lh ll l a c h a ra c te rise d by th e form ation o f a c y lin d ric a l c e l l a r a t the v a sc u la r nodes* From t h i s varying numbers of tra c e s a rise *

The tr a c e s In ­

crease i n number i n accordance v lth th e com plexity o f th e organ supplied* The peduncle l a a m odified a e r i a l a te * .

I t d if f e r s

fro * th e h o lh ll and rhisome i n th a t i t c o n tain s much s c le r enehyma, cambium, secondary xylsm , and a narked endodermis. The v a a e n la r anatomy o f th e b a s a l p o rtio n of th e ra e h is has been described*

One tra c e su p p lies th e b r a c t,

two a b u lb il o r a flo w er.

The normal tv e tr a c e s may fo rk

to supply two flowers* The rhizome l e markedly s im ila r to the b u lb i l, d iffe rin g

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*1

c h ie fly l a i l u , and l a th e n atu re of th e z y le a elem ents whloh l a th o r h is e a i a r t tra c h e ld e . fho Xoairoo aro m ostly laaeooXatO aad r e tic u l a t e ly re in e d .

Tho ▼aeeolar system of tho p o tlo lo aad m idrib la

a elphenoatele*

fho ooroao aro membranaceous aad o o a ta la

aaeh aolerenehyaa. fho r a a e u la r huadXo of tho ro o t l a e ith e r t r i e r eh o r p entaroh.

S tarch l a atorod l a tho perieyeXe