Studies in the IV. Floral

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82 Acta Phytotax. Geobot. Vol. ^{XXXVII, Nos.} 1-3 ^･

Studies in the IV. ^Floral

floral morphelogy and anatomy of Nymphaeales

anatomy of Nelumbo ^nucijera

Motomi ITo*

ge ue itd ^{: - l v7} gOzaOtk wt ^JZ fig e7 ts-ve

IV.

ixxcDJtEcDpkEIii[IJIia#.E..

^The genus ^IVlelumbo

^{consists of}

^two

^species,

^{the Asian N.}

^{nuctfara and}

^the American N. pentopetala. C}assically, ^this genus belonged to the Nymphaeaceae in

a

broader

sense.

However, the

authors of recent

publications ^have

^{recognized a} monotypic

family, the Nelumbonaceae, because fo11owing

^characters

do

^not

fit

^with

the

other nymphaeaceous

genera;

^{a tricolpate}

pollen ^grain,

^and

^the

^{absence of endosperm and}

perisperm (Li, ^{1955; TAKHTAJAN, 1980;} CRoNptJisT, 1981; TAMuRA, 1982).

^The

^{vascular anatomy of}

IVlelermbo is

characterized

by the large tracheid

and atac-

tostele-like arrangement of vascular

bundles in the peduncle, petiole

^and rhizome(EsAu,

1975; EsAu

and

KosAKAi, ^{1975; MosELEy}

^and

UHL, 1985). These two features

^are

sometimes considered

to be

representative of monocotyledons.

MosELEy

and

UHL

(1985) ^{described the detail fioral}

vasculature mainly

in N. Penttipetala. ^{Here, I intend}

to describe

a

floral

vasculature of

the Asian

^species

^N.

nucij2ira

in detail

compared with

the previous

observations.

Then, the phyiogenetic

relationships

between A]lalumbo

^and

the nymphaeaceous

genera

^will

be discussed.

'

'

Materials

^and

Methods

The flowers

^of

^AJlalermbo

^{nucijlera were collected}

from _plants

cultivated at

Kyoto Uni-

versity.

The

materials were

fixed in FAA, dehydrated ⁱⁿ

^an n-butyl-alcohol series and embedded

in _paraMn following

standard

procedures. Sections

^{were cut at a}

thickness

of

15-20 _"m

^{and stained}

with

^{safranin and}

fast _green. ^{The fioral}

vasculature was re- constructed

from transverse

sections using a

Nikon prejector V-12.

*

Department

of

Botany, Faculty

of

Science, Kyoto University, Kyoto 606, Japan

The Japanese Society for Plant Systematics

The JapaneseSociety for Plant Systematics

September, 1986 Acta Phytotax. Geobot. 83

i'

,

]"ig.

c

1. Vascular bundle

the peduncle ^of IVkelumbo nucijlera.

A: Tran$verse

section ef the

peduncle

(

^×

^{4). B: Central}

^{area of the}

peduncle (

^×

^40),

^{asteroid crystals}

^lining

air canal.

C: peripheral

area of the

peduncle (

^×

^{40). Roman}

^{numerics are responsed to the vascular}

bundle _group,

ex- planation

in

text.

I, laticifer; _ph,

_phloem; xy, xylem.

111

i,･

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84 Acta Phytotax. ^Geobot, VoL ^XXXVII, Nos. 1-3

Observations

Floral

constrttction

In Allelumbo

^{nucijlera each solitary}

^{flower terminates}

^a

]ong _peduncle, ^The

^sepals

and

petals

^{are attached spirally to the elongate} receptacle.

Sepals

^are smaller

than thc petals

^{and are}

^faIlen

^{at anthesis,}

^{In the}

^{upper course of}

^the

receptacle,

the petal

become

^{smaller and narrower.}

^The

^{numerous stamens are attached spirally}

^{to the}

receptacle.

The filaments

^are

filifbrm,

and

the

anthers are extrorse

(Fig. ^{5D). The}

connective

is proniinent. Above the level

^{at which}

^the

^{stamens are attached}

^the

^re-

ceptacle

has the

^nafrowest

diameter. At a higher level, the inverted

^{conical receptacle}

becomes ^{larger in} diameter. ^The gynoecium

^{consists of}

7-18

^carpels.

The

^ovary

^is

'

ellipsoidal and the almost

part

^sunk

into

^the receptacle.

The

short styles ^are

terminated

by

^a

dish-shaped

stigma.

The

stylar canal continues

into the locule. The inner

sur-

face

^of

^the

^{stylar canal}

^is

^{covered with}

papilae. ^In

^{each carpc], a single anatropeus}

ovule

is borne

^at

^the

^upper

boundary

of

thc locule

and

lies

^a

little toward the

ventral side.IFIoral

anatomp

^{1. Peduncle to the} base

^of

^the

receptacle-In

the peduncle, ^there

^{are six}

^to

^eight

large

^{air canals arranged}

ⁱⁿ

^{a circle, and small canals are}

located

at

peripheral

^area

Fig.

11,#'I,/,X,･li-l-,Y\.//,i,ag..

maca

^3S-...,.

2,

^Vascular

^bundle

^of

^IVlelumbo

^nrtcijlera

(

^×

²⁰⁾

A: Central

^vascular

^{bundle in}

^the

peduncle

of the receptacle.

C, D: Central

vascular

bundle

succcssively

higher leve], G: A little below level

to the receptacular ^wall around the carpel.

H:

//,,,k

^mge･-

^{ge g} ///G.eq ⁱ i/i,,,llV./ilL/. ⁱ

^X-m

^{, ' ee /}

^me-

^{$ ･ii} eeIR.,

ee

^u-:eeg,ll

^' l,.,er,,,M ^'uaM

. ',K'

, Xi,

^'

lii/W"':-i" S,//inVi,,

^"･m

sEli/･,ima,,l ^,,{.,,/

^･mi

i/i

^'･XliX/g'

CGroup ^{I). B: Central}

^vascular

^bundlc

^{at the}

^base

in

^Lhe rcceptacle, above

B. E-H: Carpel

trace,

^{of carpel.}

Somc

vascular

bundles

are

given

^off

Bottom

of the carpel,

ph, phloeml

^{xy, xylem,}

i

The Japanese Society for Plant Systematics

The JapaneseSociety forPlantSystematics

September,1986 Acta Phytotax, Gcobot. 85

Fig,

A

t'v.'l

S. Transverse

section of' the

flower

of the

Nelumbo

nucij?ra, successively

higher level (

^×

^{6). A:}

Peduncle. B-D:

Bottom ot'thc receptacle.

F-I: Lower half

of the rcceptacle where the sepals and

pctals

^attach.

,J-L: ^{Middlc level}

^{of the receptacle whcre numerous stamens attach.}

^{M-P: Upper} halfof

the rcccptacle

Arrow,

speal or

pctal

^traces;

Arrow head,

cortical vascular

bundlel

c, carpel

trace;

pst pea]t-stamen

^{trace; r, ring vascular}

bundle t'rem

which stamen traces are given off,

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86 Acta Phytotax. Geobot. Vol. XXXVII, Nos. 1-3

caeeew

tw wa rmtva/unmJ,l'

ee"ma,,eessec

1/

The Japanese Society for Plant Systematics

The JapaneseSociety forPlantSystematics

September,1986 Acta Phytotax.Geobot. ⁸⁷

(Figs. ^{IA, 3A, 4A).}

were observed

(Fig.

these

air canals.

Along

^the

inner IB). Throughoutwalls

of

these the peduncleair

canals,

there is

^nomany

asteroid

mterconnectloncrystalsamong

A

^oOe

i, i21g

^e

^{･ lo:}

^e

e

B

%e8 90o

c

sQOoo

Q O

o o

O b

^0eeeb

s

.qQOO`{b

o o

%eebbb

'

1

qqooo

Q O

o o

oeoob eb

D' E F

e eo

e o

o oo

oo e

o o

oo

e

Fig.4. Schcmatic illustratien

of the transverse section of the

fiower

of

IVlrtumbo

nucij}ra.

A: Peduncle.

B: Bettom

ef the receptacle.

C: Lower half

of the receptacle where

petal

^{traces are}

given

^ofll

D:

Mid

of the receptacle wherc many stamen traces are

givcn

^off,

E, F: Upper half

of the receptacle.

e,

Fig.

ptk$"-pt

IIIIS

getev

tuAtae.

dime,fu

^za

P4 h

.E lf･,tkl

^--.Tk･

-･st- - 1-

^,･･

^.. v' {

ec･

5, A: Cortical

vascuiar

bundle (

^×

^{80). B: The}

^{vascular ring which}

^branches

^{the stamen traces}

(

^×

^{4e). C: Transverse}

^{section of the petal.}

^There

^{are two types of} air canal and twe vascular systerns, ^{main systcrn} and auxiliary one

(x40>. ^{D: Tran$verse}

^{seetion of the stamen}

(80).

^a,

vascular

bundle

ofauxiliary system; ac, air canal; m, vascular

bundlc

of main system;

ph, phleem;

st, stamen trace; xy, xylem.

l,･

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88 Acta Phytotax. Geobot. Vol. ^{XXXVII, Nos. 1-3}

'

^The

^vascular

bundles

^{are scattered}

in the peduncle,

^although

^they

^show a certain

regular arrangement and are

grouped ^{into fo11owing four} groups ^by

^their

position.

I). The large

vascular

bundles locating in

the central area of

peduncle (Fig. ^IB-I,

2A):

six to _{eight vascular}

bundles

are arranging

in

a circle.

Each

collateral

bundle

has

some

laticifers between

the xylem ancl

phloem. Each

^xylem

is

composed of usual-

ly three (or

^{rarely one)}

^{large tracheids. The}

^{walls of}

^{the largc tracheid}

^{are often}

^broken

through the expansioh'in its diameter. II). The bundles located in the

septum of

large

air canals

(Fig. ^{IB-II): Each}

^of

^then

^{consists of}

^two

^{collateral vascular}

^bundles,

an outer one and an

inner

^{one, and}

between them

a small

bundle

^{without xylem.}

The

structure of

the

co]lateral

bundles

are

the

same as

that

of

the

central

bundles. The

outgr one

has three large tracheids

and the

inner

one

has

only one.

In the latter, the

xylem and

phloem

^{are reversed}

from

ordinal

direction (Fig. ^{IB-II). III). Small}

^col-

lateral ^bundles

^{scattered around}

^{the large}

^{air canals}

(Fig, ^{IB-III): they}

^{are reversed.}

In

each

bundle,

^there

is

^{only one}

tracheid,

rarely not.

This tracheid is

rather small

comparing with

that

of

the

other

bundles. IV). The

vascular

bundles

scattering at

peripheral

^{area of the}

^peduncle CFig, ^{IC-IV), There}

^are

^{both large}

^{and small}

^bundles

in this

area.

In both, phloem

^and

laticifers

are observed,

The

number of

tracheids

is three in the Iarger

and one

in

smaller

bundles. ^There is

^no

interconnection

^among

any of

the

^vascular

bundles throughout the peduncle. ^This

arrangement

is

almost

similar

to that

of

N. Pentcipetala (MosELEy

^and

^{UHL, l985).}

At ^the basc

^of

the

receptacle, the air canals and

the

vascular strands change

their

shape and

position dramatically. First, the Iarge tracheary

elcments of all vascular

bundles

are replaeed

by

many small

tracheids (Figs. ^{2A-C). The three tracheids}

^as-

./.

sociated with each

Iarge bundle in the pecluncle

^are separated

by the

xylem

parenchyma

in the peduncle. Through in the base

^of

the

receptacle, many small

tracheids

are crowded

together

and are mixed with

few _parenchyma

cells

(Fig. ^{2C). At}

^{a slightly}

higher level

of

the base of the

receptacle,

the

vascular

bundles

^{scattered around}

the peripheral

^area

(Group IV-described IV in

above

text)

and

the

outer

bundles between

large

^{air canals}

(Group ^II)

anastomose,

At the

same

level, the

septa

between the

air canals

disappear

and a ring shaped air canal

is formed. As

a result,

the

receptacle

is

separated

into

^{a central}

_part

^{and a}

peripheral

^ring

^{in the}

^{cross section}

(Figs. ^3B-C,

4B). ^The

anastomosed vas'eular

bundles

^originate

from Group II

and

IV

are rear- ranged ^near

the inner portion

^of

^the peripheral

^ring.

^{The inner}

^vascular

^bundles be-

tween the large

^{air canals}

(Group ^II)

^and

^the

^vascular

^{bundles inside the large}

^air

canal

(Group ^III)

^move

^to

^central

part

^{and arrange at}

^its peripheral

^region

(Figs. ^2C,

3B). ^These bundles

^{are usually collateral and reversed as}

in ^the _peduncle. In

some cases,

these bundle become

concentric.

The

vascular

bundles in the

central

part

^of

peduncle (Group ^I)

^{establish a central system}

^{in the}

receptacle.

The

central air canal

becomes

^{smaller and}

is gradually

^replaced

by _{pith parenchyma} (Figs. ^{2C, 3B).}

NII-ElectronicL.i

The Japanese Society for Plant Systematics

The Japanese Society for

September, 89 At

^a

higher level,

some of

the

neighboring coliateral vascular

bundlcs in _peripheral

ring unite

together

and

form

^a concentric vascular

bundle. In the

same manner,

the

vascular

bundles in the periphery

^{ofthe central area also}

become

concentric

(Fig. ^5A).

At

^a

higher level, the peripheral

^{ring and central area}

become

^{continuous again;}

the

ring-shaped air canal

is

successively

divided by parenchyma

^and

gradually disappears

(Figs. ^{3D-F). The}

^concentric

^bundles

^of

^{both the} peripheral

^{ring and central area}

spread

into the

cortex of receptacle and

form

^{a cortical vascular system}

(Fig. ^5A).

2. ^Lower half

of

the

receptacle･

At the level

where

the first

^sepal

traces

are es-

tablished, the

number of vascular strands

in the

central vascular system

is

about

20,

although

the boundary

of each vascular strand

is

not always clear

(Fig. ^{2D). In}

^some

cases,

two to four bundles have

^{a single common}

_protoxylem

although metaxylem and

phloem

^{of each}

^bundle is discrete. On the

other

hand, two

or three

protoxylems

^are

sometimes observed

in

one

bundle. The

number of

the

vascular

bundles in

central

stele

increases by the

radial

division

of each vascular strand

in the

upper course of the

receptacle

(Figs. ^{3D-H, 4C).}

^{Two types}

^{ef air canals are observed}

^{in the}

^sepals.

One is larger,

^oval

in

^cross

section and arranged

in

one

line

aleng

the

middle of

the

sepal.

The

^other

is

smaller

and almost circles

in

cross section and

distributed

^near

both

^{the adaxial and abaxial}

surfaces

(Fig. ^{5C). Two}

^{vascular systems are observed}

^{in the}

^{sepal, a main system and}

auxiliary one.

The

^{main vascular system} consists of several

large

vascular

bundles located between large

air canals

in the

middle.

The

structure of

these

vascular

bundle

resembles

that

of

peduncle. Each bundle has

one

to three large tracheid. The

other or auxiliary system consists ofsmaller

bundles distributed

^near

the both

^surfaces.

Each bundle is

associated with one

large tracheid

or

is

without xylem

(Fig. ^{5C). All}

^{of the}

vascular

bundle in the

sepal are collateral

(when

^xylem

^{is present). The}

^{structure and}

arrangement of vascular

bundles in

the

petals

^{are similar}

^{to those} in

sepals.

Both ^the

^{central and cortical vascular systems supply}

^the

^vascular

^{bundle tQ}

^each

ofsepals and

petals. ^{The trace} branched from

^{central system}

becomes

^{the main system}

and the traces

from

cortical system

become

^{auxiliary system.}

A _trace for

main system

is branched

^via

bifurcation

^or

trifurcation

of central vascular strands which remain

for

a rather

long distance in ^the

centrai stele.

Then, the trace

runs obliquely

in the

cortex.

This trace is divided into five to

nine

parts

^at

^{the base}

^{of each}

petal.

In ^the petal, ^the

^sequence

^{that the trace for}

^{main and auxiliary systems}

is

^similar

to that

of

the

sepal;

traces

supplying auxiliary system of abaxial side, main system, and adaxial auxiliary

one (Fig. ^{6). The}

^{vascular suppiy of}

^the petals

^resembles

^that

^of

^the

sepals.

In the

_upper

(inner) petals, ^{however, the trace from the}

^{central system}

becomes

cencentric

in the

cortex,

This

^{concentric strand}

bifurcates,

and one of

them

enters

a

petal

^and

^the

^{other remains}

in the

cortex.

This

remaining strand continues

to

run

upwards

in the

cortex and

later

^{supplies a stamen at a}

higher level (Figs. ^{3I, 6).}

Plant Systematics

1986 Acta Phytotax. GeobDt.

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90 Acta Phytotax. Geobot. vol. xxxvll, Nos. 1-3

C

P

ama

SE

.

'

t'1

. co ce

Fig. 6. Schematic illustration

ofthe

floral

vasculature ofIVketumbo nucij2ira.

Stamens

and cortical vascular

^{system on right}

^half

^{are omitted.}

^Thick

^solid

^line,

^{central vascular systcrn; thin sold}

^line,

^cortical

^{vascular system;}

^C,

^carpel;

^{P, petal; SE,}

^sepal;

ST,

s-tamen.

.

^At

^a

^{level higher than that}

^of

^{the insertion}

^ofthe upperrnost

petal,

^{a vascular strand}

which

is branched

off

from

each of

petal ^traces

^spreads

in

^a

tangenital direction

and makes a

lateral interconnection

^with

^the

neighboring strands.

As

^{a result of}

this

^anas-

tomose,

a ring-shaped vascular

plexus ^is

established.

Numerous lower

^stamen

traces

are

fbrmed

successively

from this

structure

(Figs. 3J, ^{4D, 5B). A}

^{single vascular}

^trace

enters each stamen.

MosELEy

and

UHL (1985) ^have

^observed

^that

^{a stamen receives the}

traces derived from

cortical system ^as well as

from

^{main system.}

^However, in the

present

^{study, such}

^traces

^{were not observed.}

It

may

be

^related

^{to the}

^{size of observed}

flower. ^The

^vascular

bundle in ^the

^stamen

^is

^collatera!

(Fig. ^{5D). At}

^a

^level

^slightly

above

the departure

^of

^the

^stamen

^{traces, the}

ring-shaped vascular

plexus divides into

many small concentric vascular strands

(Figs. ^{!U-K). These}

^{remaining strands supply}

the trace to the

^{upper stamens}

by

^successive

divisions

^and

they gradually become

^smaller

and

finally disappear in the

upper

half

of

the

receptacle.

.

3. Upper half of the

receptacle-Just

below the

^narrowest

part

^of

^the

receptacle,

the

strands of

the

central system number about

20. This

number

decreases through

fusion

^and

becomes

^about

7-18

^strands

in

^{accordance with}

^the

^{carpel number}

(Figs.

3K-M, 4E). At

a

higher ^level,

^about

half

^of

these

strands move outward and are ar-

ranged

in two

or

three

whorls

(Figs. ^{3N, 4F). Each}

^strand

^becomes

^{concentric at a}

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The JapaneseSociety for Plant Systematics

September, ^{1986 Acta Phytotax. Geobot. 91}

higher level (Figs. ^{2E, F),}

^{and supplies a single carpel.}

^The

^{vascular strands of}

^the

cortical system are scattered

throughout the

receptacle.

At this level

^{many air canals}

appear again and

gradually ^they become larger in

accordance with

the diameter

^ef

the

receptacle.

The parenchymatous

^{tissue of the receptacle}

is partitioned by those

air

canals and

the

resulting

parenchymatous ^tissue becomes two to

six cells

layer's

thick

in

^{a cross section}

(Figs. ^{2F, 30). The}

^{concentric strand radially}

^{divides into}

^six

^tQ

eight, and

the

alternate

three

or

four

^{strands run outwardly and run up}

^{in the}

^receptacle

around

the

carpel

(Fig. ^{2G). The}

^{rest of}

^{the bundles divides into}

^{numerous ones}

just

below the bottom

^of

the

carpel

(Figs. ^{2H,3P). The bundles}

^{enter each carpel and}

fbllow the

ovary wall where

the

strand varies'in size.

The largest

^one

has

^a

few big

tracheid,

while

the

smallest one consists ofonly

phloem. Up to the

stigma

these

strands run

parallel

^and

^have

^no

interconnection. Most

of

the larger

ones supply an ovule.

At

the top of the ovary all of the ovular supplies converge on

the

ventral side and simul-

taneously

enter

the ovule. In the funicle, five bundles

^are

observed. The

vascular strands remaining

in the

carpel wall move up a

little distance in the

short style.

Discussions Structure qf

^vascular

bundte

In the peduncle,

^vascular

bundles

are characterized

by having

a

few Iarge tra-

cheids

in

xylem.

In the

receptacle

the

vascular

'bundle

exhibits a

different feature;

having

many slender

tracheids in

xylem

(Figs. ^{2C, D).}

^The genus ^IVktumbo

^resembles

^the Nymphaeaceae in the fo11owing

anatomical

fea-

tures

ofthe

peduncle;

^the

presence

^of

large

air canals, scattered vascular

bundles (except

Brasenia

^and

Cabomba)

^and

the presence

^of

laticifers. On the

other

hands, ATlalermbo

diflbrs from thesc genera ^{in the}

^{absence of}

protoxylem ^{lacuna, Protoxylem lacunae}

have been

observed

in

all

genera

^of

Nymphaeaceae (MosELEy, ^{1961, 1965; SaHNEmER,}

1976; ITo, 1983, I986). The function

of

these Iacunae is

related

to the transportation

of water to

fioral

organs

(ITo, ^{1983). In}

^the

^peduncle

^of

^IVblumbo,

^{there are}

^{large tra-}

cheary elements and

their function

^seems

to be

^{same as}

the protoxylem lacuna in the

nymphaeaceous

plants.

"FZoral vasculature

In the flowers

of

IVlrlumbo the

receptacular vasculature consists of

two

systems, central

(ce ^{in Fig. 6)}

^{and cortical system}

(co ^{in Fig. 6)} (MosELEy

^and

^{UHL, 1985).}

Traces

of

the

main vascular system

to the

sepals and

petals (m ^{in Fig. 6)}

^{are supplied}

from ^{the central}

^systems.

In the present ^study it is

^confirmed

that

central and cortical vascular systems are rather

independent

each other.

The

cortical vascular system con-

tributes to the

adaxial and _{abaxial vascular system}

(auxiliary

^{system, a}

^{in Fig. 6)}

^of

^the

sepals and

petals.

^The

^{auxiliary vascular system ofa}

petal

^{and!or sepal}

is

also

found in Victoria, Eunyale

The Japanese Society for Plant Systematics

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The JapaneseSociety forPlant Systematics

92 Acta Phytotax. Geobot. Vol. XXXVll, Nos.1-3

(MosELEy, ^{1958; SaHNEmER, 1976; ITo,} unpulbished data) and

some species

of dym- Phaea (vAN ^{HEEL, 1977). In both Victoria}

^and

^Eittyage,

^vascular

^bundles

^{of auxiliary}

systern are supplied

from peripheral

^vascular

bundles

which are

located in

^the

_peri- pheral

^regions

ⁱⁿ

^their receptacles

(ScHNEiDER, ^{1976). These bundles}

^can

^{be designated}

corticai

bundles,

although

they

contribute only

to

sepals and

petals. P'ictoria

and

Euryale

have large fiowers

and

their

auxiliary vascular

bunclles

^seems

to have

^a

_parallel func-

tion to the

^cortical

bundles in IVlelumbe.

The presence

^{of a cortical vascular system}

^{in the flower has} been

reported

in

several

families

as

listed in ^Table 4

of

MosELEy

and

UHL (1985). These ^{plants have}

^rather

large

receptacle.

As pointed

^out

^by

^{some authors}

(CARLguisT, ^{1968s MosELEy}

^and

UHL, 1985), large fiowers

need to

have

^{many vascular}

bundles for their physiological

Table1. Diameter

ofreceptacle and presence or absence of cortical vascular system

in Nymphaeales

and

Berberidaceae.

Species ^Diameter

ofreceptacle-(mm)

Cortical bundle ^Reference Nelumbonaceae

IVlelumbe

nucgera

Nymphaeaceae

NpmPhaea

tetragoua

Eiiny7aleforox

Victoria

amozonica

Barclt!ya

motleii

IVbePharsubintagerrmtmi

Brasenaischreberii Clabomba

eareliniewa

Berberidaceae

IVlandia demestica Beriberis thunbergii Mlahoniajoponica Ranzaniaj'oponica EPimedium grantdePerum

VLinceuveria hexandea Plagierhagma dubia

JofZ,rsonia ^dipfp,Ua

Achllystripnjlla CLiulopfp,ltum

thalictroides

Leontice teontopeltalum (!),mnospermiumaibertii Bengardia

ahcysagonum

Dipdylleia grayi DisosmaPteiantha

Pb`kiplyll"mPeltatttm

P. hexandra

15. 0 4. 5ILO31.

2*4.

1 5.53.

22.

1 L O*

Q 8*' l. O*

l. 3*

O. 8*

O. 6*

O. 9*

1. 6*

O. 4*O.

5*O, 9*

1. 0*O.

9*

1. 0*3.

9*3.

3*2.

8*

resentp

'absentauxirally

auxirally

hbsentabsentabsentabsent

abseritabsentabsentpresentabsentabsentabsentabsentabsentabsentabsentabsentabsent

(proscnt)

presentpresentpresent ITo, l984 ITo,

unpublished

SCHNEIDER, 1976 ITo,

unpublished

ibid.ITo, 1986a ITo, 1985a

TERABAyAsm, 1983c TERABi\yAsHi, 1978 ibld.TERABAyAsHl,

1977 TERABAyAsHi, 1979 ibid.TERABAyAsHI,

1981 ibid.ibid.TERAByAsHi,

1983c

ibid.ibid.ibid.TERABAyAsHI,

l983b ibid.ibid.ibid.

*

Measured from figures in

original

papers.

The Japanese Society for Plant Systematics

The JapaneseSociety for Plant Systematics

September, 1986 Acta Phytotax. Geobot. 93

'

need, and

the presence

^{of cortical vascular system seems}

^to be

^related

to the

size of re- ccptacle,

The

relationships

between

^receptacle size and

presence

^{or absence of}

^the

cortical

bundle in two groups, Berberidaceae

and

Nymphaeales

are summarized

in ^Table

1. There is

a

positive

correlation

between them. Nevertheless, the presence

^{or absence}

ofcortical

bundles is also

^related

to phylogenetical

^relations,

for

^{example, this nature}

is

constant

in

^each

Tribe in Berberidaceae (Table ^{1). For further} discussion,

comprehen-

sive survey of

the flowers

^which

have

cortical vascular system

is

need.

^The

^stamen

^traces

^are

derived

^not

from the

central vascular system

directly, but from the

vascular strands _which

branched from the

upper

petal ^{traces. These traces} branched the

successive stamen

traces (Figs. 3j-L, ^6). Suchavascular

supply

to

^stamens

is

also reported

in

some

families (vAN ^{HEEL, 1966).}

^rl"his

^type

^{of stamen supply re-}

presented ⁱⁿ

^these

families is _generally interpreted

as an

increment

of

the

stamen num-

ber. In IVlelumbe,

^as

pointed

^out

by MesELEy

and

UHL (1985), ^the

^{stamen vascular}

supply also suggests an evolutionary

trend

of

increment

of stamen number.

The

^receptacle vasculature

in ATlalumbo diffl]rs from the

nymphaeaceous

genera ⁱⁿ

the fbllewing features: 1)

absence of

girdling bundle

^at

the base

^of

the

receptacle,

2) presence

^{of a}

^typical

^eustele

in the

receptacle and

3) _presence

of a cortical vascular system.

^{In the}

nymphzeaceous

genera, ^there is

a complicated vasculature at

the base

of receptacle which _{was referred}

to

a receptacular

plexus, As discussed in

my

previous paper, ^the presence

^{ef the}

girdling bundle is

a method

to increase the

^number ofvascular strands

in the

receptacle

(ITo, ^{1984). In IVblumbe, the}

^{central system}

^{in the peduncle}

consists ef six

to

eight vascular strands and at

the

middle

Ievel

of the receptacle

they increase to

about

20 ⁱⁿ

^number

^{by the} successive dtvision

of

the lewer

vascular strands.

The pattern

^whereby

^the

^{number ofvascular strands}

^{increment is}

^not

^the

^same

^between IVlrlumbo

and

the

nymphaeaceous

genera. ^The

^{absence of such}

bundles in ^IVlalumbo

suggests

that it toek

another evelutionary

path.

^In

^{addition to}

^them, IVlalumbo has

^{many characters which}

difller from the

nym-

phaeaceous genera; ^tricolpate pollen (UENo

^and

^{KiTAGuaHi, 1961),}

^{seed without}

peri-

sperm and endosperm

(Ll, ¹⁹⁵⁵⁾

^clearly

^{bladed first leaf} (HAiNs

^and

^{LyE, 1975; ITo,}

1982),

and

prcsence

^{of vessels}

(KosAKAi

^{et al.,}

^1970). Cytological

^and serological studies also reveal

the distinctiveness

of

IVielumbe (SiMoN, ^{1971a, b; OKADA}

^and

^TAMuRA

1981), The _present

^{study shows}

that the diflerence in the

morphology of the vascular

Table 2. Anatemical

characters

in

the

Nymphaeales

Character IVlelu.Bras.CLtbo.MPh.Alymp'Eury.vact.Clera.

Protoxylem lac-una Vascular _plexux

Cortical (c)

^{or auxilary}

(a)

vascular system

c

++ ++ ++ +

+-ora

++a ++a +

The Japanese Society for Plant Systematics

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94 Acta Phytotax. ^Geobot. Vol. XXXVll, Nos. ^1-3

bundles

^and

in the fioral

vasculature

between IVlelumba

^and

^the

nymphaeaceous

genera

is

^also clear.

These differences

are sumTnarized

in Table 2. ^'

The following

special

features

of

floral

vasculature

fbvnd in IVketumbo,

such as

pre-

sence of

cortical

^vascular

system

^and

its

contribution

to

sepals and

petals,

^common

vascular supply

to

stamens which run up cortical region, are also

found in Magnoliaceae,

altheugh vasculature

in gynoecial

^region

is diflerent

^{each other}

(SKipwoRTH

^and

^PmupsoN,

1966; UEDA, l986). This fact

may suggest a

phylegenetical･relationship between

^them.

Further

studies of many aspects of

them

are need

to

clarify

this problem. ^'

Acknowledgment

^I

^would

^{like to}

^{express my cordial}

^{thanks to} Pro £ Kunio IwATsuKi

^of

^{Tokyo Uni-}

versity _and

Profl Shoichi KAwANo ^{for their}

^{valuable advice.}

^{My thanks}

^{are also}

^due

to Pro £ Nobuyuki FuKuoicA

of

Shoei Junior ^{College, Dr. Susumu} TERABAryAsHi

^of

^Tsu-

mura

Laboratory,

and

Dr. Kunihiko UEDA

of

University

of

Osaka ^Prefecture fbr

^critical-

ly

reading

the

manuscript and

Dr. Fredrich H. UTEaH fbr linguistical

check.

'

Sumniary

'

Floral

^{morphology of}

IVlelumbo

nucijiira

is

observed chiefly

from ^the

^anatomical

point

of view.

Vascular bundles in the peduncle ^have large tracheids. They

are classified

into five types. In the

receptacle morphology of

the

central vascular

bundles

changes

to

typical collateral

bundles

^which

have

^{xylem with many slender}

^tracheids.

The flora}

vasculature

in

^{the receptacle consists of}

^two

^{systems, central and}

cortical systems.

The

main vascuiar system of

the perianth is

supplied

by

a single

trace from the central

vascular system.

These traces depart

spirally.

Traces to

stamens are supplied

from the

cortical vascular system.

Usually

^{a single cortical strand sup-}

'

plies

^{several stamens}

^by

^the

^division

^{of these strands.}

The

comparative

diflerences in floral

vasculature

to

nymphaeaceous

genera

^are

fo11ewing

respects:

1)

absence of

girdling bu.ndles

at

the base

of

the

receptacle,

2)

existence of

typical

eustele

in the teceptacle, 3)

^{existence of a cortical vascular system.}

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