ANATOMICAL STUDIES ON THE BARK OF DIFFERENT FOREST TREES OF MADHYA PRADESH

DISSERTATION SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF

MASTER OF PHILOSOPHY IN

BOTANY

BY

MOHAMMAD ALI KHAN

DEPARTMENT OF BOTANY Aligarh Muslim University, Aiigarh

December, 1980

DS286

2 6 AUG 1981'

.to m Momm

CBam^FICATK

mis i t to eortify tbftt th« dl«t«rUtloii m t l t U d

•*^atomloai ttttdiM on th« bark of difforont forott trooi

of Madhya Pradosh**, tubiBittad to tha Aligarfa Kiaslln tkiivartity,

Alisaih, la a bonafida voik oarriad out hy Mr, Mohd. All Khan

^dar my auparvlsloa, in partial fulfilnant of the raquiraaMits

for the award of the dagraa of Haatar of Philosophy in

iiotany.

( Ziattddin ithmad ) Eaadar,

i^apartaant of Botany. iaigaiH Mnalia Ghiirertlty,

Aligaxh.

1 wish to express my high sense of gratitude to

Mr. Ziauddin ahmad, Header, Departi&ent of Botany, for

suggesting the problem and his constant advice, guidance

and eneour^gement generously extended to me during the

course of preparation of td ls manuscript,

Sincere thanks are due to Prof, M, M.H.K. Afrldi,

Head, i)epartinent of iotany, for providing essential

laboratory f a c i l i t i e s ,

I am highly indebted to Or, ii.K.M, Oiouse, Reader,

Department of Jotany, for his valuable suggestions and

enc our ageisent,

Grateful acknowledgeaoits are extended to Prof, dhureef

A. Choghtai, Head, ^aifia College, ahopal, for his help

in the col lect ion of plant material for the study,

I am also thankful to my colleagues and friends for

their kind co-operation and a l l sorts of help whenever

needed*

( Mohaaunad All Khan }

Oeoenbery 1980,

gontm'm

RBVISW OF UT^aAlO&B

0R03S aXRUClTORS OF PHLOEM

SZaUClURiiL DEXAII.S OF SECONDiiaY PHLOEM

aiEVS SLBMiiSIS

ORGAN SbliES

COMPANION CELL

PHLOi^l PARQ^CHSOiA

PHLOEM FIBRES

PHLOEM HAXS

SCLSRBIDS

SEASONAL PRODUCIIw«{ OF SECONDARY PHLOBM

LONQSVITY OF SECONDARY PHLOBM

PERZDBRM AND f»YIlD0M8

MOSPHOLOQY OF PERXDERM AMD RHXnDOMS

c(»fpQN0iis or ms Fmaomn

FLACK OF OBI<&n OF PHffi.LOaflf

DBVSLOPMaiX OF PERID^iM

MBmOOOLOaY

PLiK OF SXODY

BBTBROICJEBi

PAGE

1 - 3

4

5 - 1 1

12-16

16-24

24-30

30-33

33-35

35-39

39-41

41-42

43-44

45-46

47-48

48-50

51f.53

53-56

56-58

59-67

68

69-94

Bark i s a non»teehnieal t«r« appli«tl eosmtonly to a l l

the t lssuas outslda th« vascular eaabiiSB in either a prinary

or secondary state of grow to in an axis , Ihe tern i s

applicable to dieotyledcms and gymnosperms ^ e r e the formation

of periderm takes place any%^ere in the regxcn between tb.9

epidermis and vascular cambium* iiark, therefore, includes

a variety of t i ssues in the cortex, periderm and prisiary

and secondary phloe:., as the case may be. AS such, i t

exhibits caaplex features, because of ttie coifiplexity of

peridennal and phloem t i ssues ,

iiark anatomy of forest trees has been a neglected

aspect of study in the past and i t i s only froc. the l a s t two

decades that the bark study, owing to i t s isDerse econcx^ic

and taxonomic signif icance, i s gaining increasing att«nti(»i

of the anatomists and proving substantially helpful to

foresters , botanists, taxonomists, pharaaoognocists and

forensic experts in various ways.

Identif ication of plant species i s so far based on

their reproductive structures almost exclusively, ^t times,

i t becomes a t i ck l i sh problem to make a proper identif ication

of plants when toey are devoid of flowers and foliage or lAfB

they have been fe l led and made into logs . Bie only thing that

eaa be helpful at that jmeture i s the knowledge of their voo4

o ^

•ad bArk anatGny. I f aware of the bark features properly,

one e«n face no problem in identliying the Tarioue epeeies

4«8t W easting a glance on their stems. I t i s specially

s ignif icant in forest practices,

farther, there are many barks t^ieh have an immense

significance from therapeutic point of view. Many such barks

in their fragmentary form are mixed with other similar looking

ones and sold in the market in adultrated form, oudi barks

can be eas i ly distinguished frcn^ the adultrants on the basis

of their anatomy.

<*s a consequence, inforciation on an: toxical characters

of tree bark, particularly those of tropical trees i s scanty

and the l iterature elucidating their feasible role in the

identif icat ion comprises of merely a few isolated reports

(Ihorenaar, 1926$ Symington, 1943; wood, If962{ Chattaway, 1953,

1969{ Browne, 1955; ^ihitmore, 196Sa, b, c, 1963; Ksau, 1964|

Begin, 1969 and JTunus, 1976). Ihe possible reasons for the

negllgenee towards ^ i s iaportant aspect of study are at

followst

(a) Structural and deToIc^BMital conqplexity of the system

as eoaqpared to wood.

(b) Oelieate and teehnieally l e s s aeeessible nature of the

various e^Bponents particularly phloem.

(«} hmek of faatidlotts teahnique.

«J

(d) LMk of interest of tti9 coBinorelal world In this tlsso*

tystoR at ootipar«d to xylon, whieh is eoniBsoreially taportant

as tinbsr sineo long* inliils the ralm of barK if detoriBiiiod

largoly by Its eontonts of fibrss, organic substanoes and

arcwiatie compounds such as tannins, latex, drugs and so on,

H. delve of literature reveals that the study an bark

anatomy of tropical trees particularly tiiCMie of Indian forest

trees i s very meagre. In India no such attempt has yet been

made and the present problem pertaining to the forest trees

of .-ladhya iradesh around ^opal, is probably the maiden

attempt to work out the bark anr^toi of a pretty large number

of species in a consolidated manner•

4

Hccordlng to Esau (1965a), the tern: 'bark' la applied

most Gomn-iOrily to a l l t issues outside the vascular oambiun of

the axis , in elttier a primary or seccmdary state of growth.

I t i s also used nore speci f ica l ly to designate the tissue that

accuRulatea on the surface of the plant axis as a result of

phellogen ac t iv i ty , *vS the peridern: develops, i t sepiurates

by means of a nwi-living layer of cork c e l l s , variable amounts

of prisiary and secondary t issues of the axis from tiie subjacent

l iv ing t i s sues . The t issue layers thus separated die . Ihe

term * bark' in i t s restricted o^aning refers to these dead

t issues tOi^ether with tiie layers of periderm. JDius, i f bark

i s used with reference to a l l the t i ssues outside the vascular

cambium, the periderm and the t issues of the axis isolated by

i t may be cooibined imder the designati^m of 'outer bark*.

The technical term for the 'outer bark' i s '^yt idons ' ( De

iary, 1384).

In t^e young stem and roots thus the b&rk constitutes

the epidermis, the cortex, the pericyele, the endodermis and

the different phloie eonponents, while in the older parts of

a plant, the secondary phloea periderm and rhytidone are

involved, partienlerly in dieotyledona and gymnospems vhere

radial growth ooenrs.

For the purpose of this review phloie eoaponenta wi l l

5

h% taken f irs t follotfed by perid«ri» and rhytidOM and eortax

will not be daalt with at a l l .

aaosti t>rHUCfUiiii: oi pnLom

thloem Is the principal food conducting tissue and

canstitutea the vascular system of the plant body in oombina-

tloa. with xylea. It develops as a peripheral vascular tissue

and forr s a part of the bark in stes-.s and roots of the vascular

plants, Jepending cm the origin and time of appearance in

relatiOTi to development of the plant or the organ, the phloem

can he classified into two major groupsi

i) Priaary phloera

11) Secondary phloem.

^) PriiarY BfalfftK

Ihe phlowD that develops from procambium in the priniary

plant body i s termed as 'pr^&ry phloem.' Ihe priotary phloem

initiates during the embryonic stage of the plant and i t s

differentiation completes after the formation of prlaftry body.

I t is distinguished into the f irst formed protophlocm and

the later formed metaphlow elements. Ihey are different in

their positiOQ, structure and developmsat.

\}

a) ProtophloiBt

The portion of tho phlow tSiat difforoatiatoft f l r t t i t

tormed as 'Protophloem* by Hustow (1B72),

ihe protophloem, together with protoxylem, ewistltutos

the vascular t issue of the young elongating parts of a plant

and ccK^talns s ieve elements possessing the usual specialized

characterist ics , that I s , highly vacuolate, enucleate, proto­

plasts and walls bearing sieve areas, Ihere i s soiree doubt

regarding the niorpholOs»lc nature of the protophloem elenwnts

in the gyrcnosperms and aince no sieve areais have been recog­

nized in tnem; they are referred to as precursory phloem c e l l s

(i^isau, 1950{ ^rlth, 1958). In anglosperms s ieve elet&ents

have been observed in ^ e protophloem of roots, stems and

leaves in woody and herbaceous species (£sau, 1939, 1950).

iliese s ieve elenents possess sieve-tube members but, in many

plants, they lack well-developed sieve plates , s ieve areas and

eoiapanion ce l l s (Bsau, 3966a; lahn, 1967). Xhe sieve-tube

a«Bbers are long and narrow, and s ieve areas can not be

distinguished eas i ly . Die walls are somewhat thiek and e e l l

contents stain s l i gh t ly . The s ieve tubes of the protophloem

function only for a short duration, and soon get erushed or

obliterated ( Ohouse « ! l i . , 1972) • In many dieotgrledons

the parenthyaa of the protophloem remains after the obliteration

of the • ! • • • • tube neoibers, and differentiates into fibres

(BIyth, ]958| lieger, 1B97). Ihe prot<^loeB in anglospermt

(H«ev«, 1942t Crafts, 1943a, b{ i^gard, 1944} Esau, 1943a, b,

1945 and Starling, 1946) dlffarantiates In aoropatal mannar

and praeadas protoxylam dlffarantlatlon (ChauTaaud, 1900;

Chang, 1935; Ksau, 1938a and Ohousa fil AI** 1372). Ttim detai ls

of the developmant of proto and metaphloam was studied by

fichnelder (1945) in peach and by i isau (1943) in the grapevine.

Xhe part of the prijQary phloem vdileh develops after the

formation of protophloem ani raatures after the growth in length

of the organ and the adjacent t i s sues , has been nannsd

' MetaphloeKi* by Van Tiei^em (1887), I t forms the main food

conducting part in soiae herbaceous dicotyledons, most Bono-

cotyledons and many vascular plants v^ich do not show secondary

grow^ and i t remains active in pteridophyta and long-living

monoootyledois, such as the Palmae, for several years after

the ful l development of their primary bodies (i^sau, 1965a)*

On the ottier hand, in woody and herbaeeoiw species , showing

caabial secondary growth, i t soon beeones inactive after the

formatioQ of secondary eonduetlng elements. Hetaphloem sieve

elemwdts may be either partly crushed or ooa^letely obliterated

in these plants (Ssau, 1965a).

Ihe s ieve elements of metaphloem are longer and wider

with more prominent sieve areas than of the protophloem. In

dieotyledons, the metaphloem postetses eompanion e e l l s and

o 0

t^Xo^n parttnehyoa iihil« in noiioeot]rl«(loQ8| the « ! • • • tub«t

and eoE^anlon o«Il8 often form strands containing no phIo«B

paronohyma ea l l s anong them, aXthou^ such esXis taay iMi pros«nt

on thtt periphery of the strands (Cheadle and Uhl, 1948).

Presence of fibres in the primary phloem has also been

reported in iicotyledonous plants by i lames and Mac Daniels

(1947) but according to Ssau (1950) they are absent from ^ e

metaphloera of pri. ary phloem and i f at a l l present in the

prii ary phloem, they arise in the protophloera as reported by

ichneider (1945) in peach and Ssau (194S) in grapevine, Kven

in those cases ^ e r e fibres develop later in the secondary

phloesj of the ssune plant, generally, a l l pri-aary phloens fibres

are longer than the secondary ones.

Ihe various c e l l types of phloem idiich are derived from

the outer derivatives of the vascular eambinm are termed as

'secondary phlo«i*, Ihey generally include four d is t inct

eoiif>oiients, i , e « , s ieve eleiMnts ( s ieve c e l l s or sieve-tube

aenbers associated with albuoinotis e e l l s and companion e e l l s

respeet lvely) , phloem parenchyaa, phloem fibres and rays«

Usually, the s ieve eleaents of seooodary phloem are shorter

in s i s e thm the primary ones. Secondary phloem consists of

well-defined sieve areas and s ieve plates arranged in radial

;i

f l l « s , a f«ature which dlstinguish«a It from tha prlnary part

of thm i^loeiB (Esau, 1965a). Iha secondary phloem Is made up

of two main systems, the axial or Tertieal system derived froa

fusiform I n i t i a l s of the cambium and the ray or transverse

system derived fror?> ray i n i t i a l s respectively. In case of

ccmifers the principal components of the axial system are the

s ieve c e l l s , phloem paroichyma including albuminous c e l l s ,

and phloeis f ibres, ihe s ieve areas are located to the radial

walls of the s ieve c e l l s (#tbbe and Crafts, 1939$ otrasburger,

1391), and phloem parenchyma c<»isists of starch, res ins ,

tannins and crystals (Jrivastava, 1963)• Xhe ray or transverse

systeci in conifers i s usually composed of ray parenchyma ce l l s

only, but socetimes albuminous c e l l s may also be present. In

conifers the rays are i:»>3tly uniseriate and homogeneous.

Ihe sec<Midary phloem in dicotyledons shows a broader

diversity of patterns of c e l l arrangement and more variations

in the phloem coiq>oa<mts than the seecmdary phloem in oMiifers.

atoried, iaternediate and non->storied arrangements of c e l l s

are seen in the secondary phloem. Xhis arrangement of ce l l s

can be determined by the nature of cambium, i . e . , i^ether i t i t

storied or non-storied and also by the extent of elongation of

the various elMtientt of the vert ical systwn during the

differontiation of c o l l s . Ihe rays may be uniseriate, b iser iate ,

• t t l t i ser iate and heterogeneous. Ihe heterogeneous rays art

eonpottd of partneh/oa e t l l t . Otnerally the phloem appeart to

io

•hov re lat ive ly more developmental disturbance In the arrange­

ment of Its eonponenta than that of conifers vrtildi I t Inherits

from the eanbluB because of the uniformity In the c e l l slxe*

In dicotyledons the components of tiie axial system are th9

sieve-elements (either sieve c e l l s or sieve-tube nembers, the

l a t t er usually associated with con^anlon c e l l s ) , axial phloem

parenchyma and fibres idille the ray system Is gwierally made up

of only parenchyma c e l l s . In both the systems the sc lere lds ,

l a t l c l f e r s , the various Idloblasts v l ^ specialized contents

(iiilsau, 1965a) and the secretory elements of schlzogenous and

lyslgenous origin,may be se©n#

Ihe variation in secondary phloem Is exhibited !^ the

fact that in herbaceous and sore woody dicotyledons the

secondary phloem Is ncm-storled and sieve-tube members are

elongated, having mostly compound sieve plates on the long

tapering end walls (iiftglft, JliiillUfift, IlZiZ&i&f MftQ&ilSUt ffrBf»

i m a , and ziavDhus. In contrast to th i s , in some advanced

dicotyledons iisSLf iftlfatritlit ijauOaUS.* iMttL* igiSlliaU* and

]2Ii!lt) the phloen i s storied and sieve-tube members are

shortened, possessing s l ight ly inclined or transverse end wal ls ,

ttiually with simile s ieve plates , fiie sieve-tube members in

some genera of sub-family Pomoideae (Hoeaeeae) exhibit primitive

features and show an i^proach towards the s ieve e e l l s of

conifers (Evert, 1960, 1903a).

Herbaeeoos dieotyledons possessing seeondary growth nay

Xi .

haw stooQdiiry phloem, showing close resemblance to those of

the woody Species (HfiaSiiaa. JSSJX&iUi)* Gaourbita ase the

exeeptlcns anong herbaoeotis dicotyledons where secondary

phloem may not be distinguished easily from the primary c»e

except in having larger ce l l s .

D«p«ndlng <m the structure and fimcticm of sleve«tul>e

elwients, the secondary phloem can be divided into the following

two types}>

i) Canductlng phlcwfo

11) ;«Qiiocaniuotiag phloem.

^e phloeia i s sa i i to be difiereatiatei Into conducting

phloec; yihec the sieve elesants become enucleate and develop

the other associated specialized characteristics, such as

well defined sieve areas with oallose, the conducting strands

between ttie cel ls ; presence of nacreous walls in sieve-elementst

appearance of fibres.

Ihe anount of yearly increment of active phloem produced

in one season varies with plant to plant and seasonal eonditloat,

Ihe eoddueting region of phloen is generally eoosiderably

narrower than the corresponding inorenent of xylen. Moreovert

in the deeidaoiis species of dicotyledons a given inerensnt of

phloea eoBMonly funetlons in eondoetion for a single season,

in the evergreen diootyledons and tlie eonifers for two seasons

probably (Orillos and Saitb, 19S9| Huber, 1939)• Ihe anoont of

the aetlve phloen in a l l tesyerate trees nMSnros a fraction of

niillla«t«r except In n i i a where It i s about I oi l l iaeter

(Uoldhelde, 1951). Ihe width of eoaduetlng zone of phloem

•arlet in tropical trees in and around Aligax^ from 0,25 urn

in Anona »anaao»a (^mad fti l i . , 1977) to 1.500 - 2,400 BB in

f9lyftjLffi4ft IqBKU?!!* (3jouae nt AIM W76) in around idigarh

and from 0.2 mm in igLML& (Holdheido, 1951) to a, IB ma in

msXSi msml9» i*>hm&d al AL*« 3^77) In regions of tropics

(Table 1). I t is evident that the active phloem constitutes

cmly a small part of bark as a v^ole.

l i ) M9ii"g9aattQtAng ptilwi»

Ihe part of the phloem in i^ich sieve-elements have

ceased to function is often referred to as non>condueting

phloem. Ihe former widely used term, non-conducting phloem,

i s ambiguous since the phloon in n^ich the sieve-elements have

beccme non-conducting, coooiaaly possesses parenchyma ce l l s

that continue to store starch, tannins and o^er substances

until the tissue is separated from living parts of the phloem

toy the a c t i v i ^ of phellogen and beoones dead (Esau, 1965a).

AS such, the phloem parenehyna remains functional while the

phloem is non-conducting. Iherefore, i t i s more understanding

to use the term non-conducting Instead of ncn-functional and

so on* file sieve elements represent various ways of the

inactive state, fiie sieve areas are either covered by a BMS

of substance known as eallose,*definitive eallose* leading to

lABLE 1

iieotti of conducting JBhlMm

His, iiPKClBi NO.

! • ACer

2 . M&LS. IBATMlPS

3 . i«<mft ffQVlM9»ft

4 . JslUlA

S. a) g.astU t%»%ViX9k

b) £• grandls

e) £.• lavaniea

o) £• Il2dJHlA

^) £• AiftMA

6. Dalonix XigJA

8« iACUI.

9* y«gflni* l i a o n i ^

10. FMxlnna

11« IHf^iilf

!£ • Hmnmitmrm^ iitdift>

13* WIlflltBi ftlaUfi

yiiPIH IN (ami)

O.a-0 .3

2 . 1 6

0 .25

0 . 2 - 0 . 3

0 .72

0 .63

0 . 7 1

0 . 9 3

1.48

1.41

0 .22-0 .675

0 . 6 1

0 . 2 - 0 . 3

1.27

0 . 2

0 . 4 - 0 . 7

1.02

0 .900 .1 .500

am KH£2iCE£>

Holdheide (1951}

«hiBad sSi &L* (1977)

- d o ­

ll oldholde (1951)

/jhriad si ML* C 1977)

- d o -

- d o -

^.««. - d o -

a>do—

-do -

ahou8« and Hashni (1976)

^B»d t i A]L* < 1^77)

Holdh«ld« (1951)

Ahnad l i AL. (1977)

ZlBBwrMnn (1961)

HoIdh«id« (1951)

^^^ t i AL* (1977)

QhoiiB« and Haihni (1976)

T4dk£ 1 (Contd.)

SL. NO,

SPECIES DKPXH IN (mm)

RBTBRBNCBS

14* li2Clli. alM 15. .flii}9ph9rm fgrr\isto9W

16. P9iyal<;t U I9ii«4f9l^a 17. PODUlUl

19. Cggreus

21. laaartoaaa Aaaica

23. itratoftlia idma

24. iziim. 2 5 . ^lavnhna m^MyitianA

0.45 ^mad e t a l . (1977)

O.aoOo 1.650 C»iou8e and Hashmi (1976)

1.500-2.400 -do -

0 . 8 - 1 . 0 doldheido (3551)

0 .750-1 .300 ::toou8e and Hashmi (1976)

0 . 2 - 0 . 3 :ioldholde (1951)

0 . 8 - 1 . 0 - d o -

1.25 Ahmad et^ a l . (1977)

0 . 2 Zlmmeroann (1961)

1.200-1.950 Ohous* and Hashmi (1976)

0 . 4 - 0 , 7 Hoiah«ida (1961)

1*36 Ahmad aX A I * (1976)

u

diteardlng of their function, Ihis i s folIOMd bx eo«pI«t«

ditappoaranee of oalloaa fron tha pores of the sieve areas

after rendering the tubes non-eondueting. Ihe eon tents of the

s iere-eleoents may be entirely disorganised, or they may

disappear and the ultimate c e l l s are f i l l e d with gases. This

state should not be mistaken with the teisiporary suspensicn of

act iv i ty or partial suspension by the depositicm of proTis i^al

cal lose during winter in Iect<»ia and some monocotyledons

(Jane /.evski, 1881)• Ihe identif ication of the non-ftinetioning

state of the sieve tubes i s particularly certain i f the s ieve-

elements are more or l e s s collapsed or crushed. In dicotyledcma

the coi&panion c e l l s and sone of the parenchyma c e l l s , while in

conifers, the albuminous c e l l s cease to function and evwa

collapse (Bsau, 1965a).

The character i s t i c s of the inactive phloem as a whole

vary in different plants. In certain dieotyledcms such as

fa4rl9atBtilr9B (Cheadle and Ksau, 1964), lUJj i , EfiBMlM «nd

Jijglans the shape of non-funotioiiing sieve-tube ehanget s l i gh t ly ,

as reported by Bsau (1965a). wftiile in others, l ike AristoloahlAy

Mii lBU ( ^ a u , 1965a), sose Oalbergia species (Ohouse and Yunus,

1975), soBM species of Hyrtaceae (Ohouse Ai A ! * , 1976b), sooe

Yerbenaeeae (Khan t i AL», 1978) and sone deciduous and ever­

green speeles of tropical plants (Ohouse and Hashai, 1976), the

• l«ve*«l«ients and their associated c e l l s collapse cosipletely.

^B XUiift vlalfegm the noa-eondueting sleve->tubes beeoM f i l l e t

ID

vith tyloses Ilk* proliferations from Tortieal parenebyma e o l l t

(Ksau, 1948}• In conifers, having fibres in the funetionlesf

phloem, the old s ieve c e l l s get crushed between the fibres and

the enlarging phloem parenchyma ce l l s (iibbe and Crafts, 1939).

Ibe non-conducting phloem usually undergoes an intensive

sc l er i f i ca t ion , especial ly by the developiaent of fibres or

sc ler ids from axial and ray parenchyma c e l l s ((Siouse and Hashmi,

1976; ithmad 9% al«f 1977), Hie intrusive growth Va&t modifies

the spatial relation araong the c e l l s , may precede sc ler i f icat ion,

The old phloesi a lso accumulates orgastic substances nalnly

crystals and phenolic compounds. Crystals are found In the

conducting phloeni and their number frequently increases with

the sclerif icaticm of parenchyma c e l l s (l-isau, 1965a} • Ihe

distribution and the types of the crystals are enough character­

i s t i c s to be useful in the cos^arative studies of phloem

(Holdheide, 1951) • Qam of the phenomena which affects the

appearance of t^e non-ccmducting phloem i s the di latation of

parenehynatotts c e l l s , by which the phloem i s adjusted to the

inerease In oirouMferenee of the axis resulting from secondaxy

growth (Esau, I9«6a)*

Ihe anoiBt of noo-eondueting phloem depends on the maimer

of phellogen fomation v i s . , the plant nay have a broad sane

of inactive phloem, i f the phellogen i s superficial , and i t not

replaced by deeply-lylag phellogen for many years as in Priy^f

u

(Sehn«ider, 1045). On tha other hand, th« inactive phloam i s

in tmall aDotmt, i f tha phallogan is fontad yaar aftar yaar

in daepar layars as in n t i a and L- MM^MSL ( ^^au, 1948 and

Khan, 1977).

oidve-elements are the principal food ccmducting elamenta

of the phloem, Hartig (1337) was in a position to differentiate

between sieve->tubes and s i e v e - c e l l s , and tho la t ter tera la

applied to the tubular structure subdivided into Individual

units Dy sonavdiat oblique end walls , Cheadle and t^itford

(1941) called the conducting elements of the phloem as »Sieve

elements* but tfaay separated the loss specialized elements

having sieve areas on overlapped end walls as the 's ieve c e l l s '

frc»i! the 'sieve*tube members' \rfiich are distinguished by having

the highly specialized sieve*areas grouped together to form a

s ieve-plate at the ends. Turner, ^ e two types of s ieve

elements, the s ieve c e l l s and the sieve-tube members differ In

the degree of differentiation of their s ieve areas and in the

distribution of tiiese areas on the walls .

Sieve c e l l s are reported In gymnosperms and vascular

oryptegams in vtildti they consist of l e s s developed sieve areas

without BMirkable differentiation froa one another and having

no s ieve plate l ike struetare on their end walls . Ihey are

long aad slender with tmpering end walls . In the tisane th«y

iV

Overlap •ach othar, and the slave areas are usually nusierout

on these ends. Ihey are rarely found in angloeperms with few

exeeptions l ike AW^r^t^UfYft ?flflnd?af (aalley and Swangr, 1949)

and ^2Cla|S aueunarla (Huber, 1939).

Xhe anglosperms sieve elemaits the 'sieve-tube members',

have certain highly developed s ieve areas invariably located

in the form of sieve plates which are formed on the end walls .

Xhe end vai l s vary from much inclined to transverse. These

units are situated end to wid, their cocmion walls bearing the

s ieve plates , thus forming a ccaitinuous tubular structure, the

'sieve-tube*, "^e s ieve areas located on the si'ie walls are

l e s s conspicuous or they are absent.

Hartig (1354) for the f i r s t time supposed that the walls

of the sieve-tubes in Cucurbita are perforated in a manner of

a s ieve . Von Mohl (1355) reported a thin swoibrane l ining the

pores. Nageli ( i 8 6 l , 1363) also followed the naae 'sieve-tube*

introduced by Hartig, since their end walls are perforated, the

•xaet tern 'si«v« area' was proposed by Cheadle and Mhitford

(1941). Ihe norphologie speeial isat ion of the s ieve-eleaents

i s represented by the developaent of the s ieve areas on their

walls and br the particular nodifieation of their protoplast*.

Sieve-areas are the wall areas with clusters of pores, throaiH

i Q

nhieli th« adjoining tUTo-olontnts ar« int«reonn«et«d bj

••ant of protoplaSBie strands. Ihiis tho sioYO aroas aaj bo

eoil^arablo to tbo priBarjr pit fiolds with plaSBOdosuta that

oeour in priaary vails of living paronehyoa eo lU . Qf eourso,

tho sioTo aroas aro spoeialisod priaary pit fiolds according

to Bsau (196Sa)«

Hill (1908) obsorTod no dlfforontiation botwoon tho

sioTo-plato and the siave^flold and he was of tho opinion that

8iore*fiolds connect tho sieyo-tubes with other sieira»-tubes

and also wil^ certain parenchyskatous cel ls of the phloem.

;>trasbttrger (1901) found t^at in gymnosperiBS ^ e closing

membrane of t^e sioTe-field remains intact, i^ilo in anglosperi

i t dissolves 80 that a single pore formation occurs. I t was

further supported br Sames and I4ac Daniels (1947).

Ihe devel<^iBent of pores in the sieve-areas was f irs t

studied in i i i l t (Hill , 1908) | Cueurbita (irroy-Wyssling and

Muller, 1957)} gttfittf&iU I U U M * M&IAIA* PftBagtglgJlt

TttfrMWi sp. and i i l U lilOllIA (£«oii §%, «!•» 3de2e){ Uim

l A U m i (Bottok and Crenshaw, 19«5)} ^fttc PlfiUlSftUJfcnm

(Northoote and Wooding, 1966)} MigotiauM (Mderson and

Cronshaw, 1970) | SOM woody species of dicotyledons (Svert

l i AL*» 3971) and CBBttrtito JiSlWk (Ssau and Choadlo, 1966$

BvortjiiAL., 19«6t Doshpando, 1976).

Iho dlaaetor of tho pores (without the lined up oollooo)

19

ia tti« siftve-areat •ari«8 In th« dlffertnt species fron a

fraction of a micron to 14/U uid avfla nore in soma dieotyladont

as raportod t^ Esao and Cheadle (1959)$ Khan (1977) and Khan

J l §JL. (1978). In yjiiiaiA Ymh9H^^t4 the dlamotar of tha pores

i s l e ss than 1/U, s l i j ^ t l y larger than 1/U in Pvrus sialic and

Pyrus eonmunis. 10.3/u in Cyg^rt^l^ spp. and 14.3/u to

Allan thus il1ris»lffia <K3a« and Cheadle, 1959).

a ley? pla^itt

lianstein (1864) applied the tern •sieve plate* to exhibit

the perforated end walls devoid of cal lose ( ca l lus ) . I^ageli

(1S61) found sc»ae s ieve-plate l i k e structures on the longitudinal

walls of the sieve tubes end he referred to them as the ' s ieve

f i e lds* . Hi l l (1908) reported that the sieve plate and the

s ieve f i e ld are the same structures and, according to him,

s ieve f ie lds ccmnect the sieve tubes not only with other s ieve-

tubes but also with certain parendiymatous ce l l s of the phloeou

Similar conditions were seen in gyimospermous sieve-elemwits

and albuminous c e l l s (Strasburger, ]B91| H i l l , 1901), Sieve

plate in s ieve tube of different regions of the phloem of

d i f fer« i t species varies in relation to i t s kind, number and

arrangement. A transverse end wall usually bears one s ieve-

plate and the inclined terminal wall bears several s ieve-plates .

Among angiosperas, the following two types of sieve plates can

be distlnguishedt

Zi)

a) Slapl* tl^ye plat« - tad wall baarlng a tlngl« siava

araa, l . a , , CMUrfr4ll»

b) Compound aleva plata - «nd walls baarlng several sieve

areas arranged In sealariform,reticulate or any other

manner, I . e . , USilSL a»d lUW. JBAll2!.<

Kaeh pore is lined ay callose, a carbohydrate* The

size of the pore is controlled by the amount of callose preskit

at a particular time. Callose is a carbc^ydrate - polymer of

glucose residues united into spirally coiled chains in 3 • 1-3

linkages (Kessleri 1958), ^ i l e cellulose occurs as straight

chains of glucose residues in 3 - 1*4 linkages and the proto­

plasmic strands of the sieve area are coiraaaly associated with

this type of oallose. Hageli (1861) for the f irst tine raf«rr«d

to i t as the *8oft substance*. Hanstein (1B64) used the t e n

'Callus* whieh was later replaced by eallose (Mangin, 1892).

Hartif (1S64) also studiad eallose but he did not name i t .

ikeoordlng to autsov (1881, 1882) i t i s present in about 800 tpp.

including anglosperasi gymospenui and eryptogaiui.

WllhelB (1881) reported that in soae eases aeeniiulation

of eallose r^prase&ts a teaporary ecssaSion of aetivit / t ••£•,

^ JULULf CtteiiPbita and ' l1*?^«Mft - J«no Zawskl (1881) also

obsarvad tba provisional eallose in winter in i^ttt||M n d soat

9 •-; L 1

monocotyledons. Howevtr, Jane ^vskl (2381) and ^trasburgar

(1B91) beile^ad ttiat la tha Btajority of plants the eallosa

aeeufflulation in sleva araas dapaads on the aga of siava tabas

and not on season. Ibis was also supported by Bsau (1965a}

and lahn (1967).

Ihe accumulation of callose occurs in two ways. In one

type the callcK»e mass increases within the pores of sieve-

areas and ecmstricts the protoplasmic strands. Deposition of

oallose also occurs rapidly on the surface of the sieve-areas.

Consequently, sieve-areas cease to appear as depressions in the

wall; instead they become thickened regions of the wall, wlhen

the sieve element reaches ^ e end of i ts activity, the sieve-

areas are blocked with bulging r&asses of callose i^ich mto

or may not be traversed by tenuous protoplasioic strands. If

there are nuBber of sieve-areas close together, the adjacent

oallose may fuse (Esau, ig65a). Since such extensive accumula­

tion of callose usually Indicates cessation of activity of

sieve elesents, the callose at this stage i s called 'definitive

callose* (ItaeoBte, 11389) • In other type, the aoeuBulatioa of

callose results only a tanporary break in activity is naned

as MoTBianey or provisional callose* (Esau, 1939, 1948).

Ihere is a controversy regarding the role of callose

in sieve-eleaents. According to sons earlier workers (Currier,

1957t Bsatt and (beadle, 1969t Crafts and Corrlar, 1963) which

Is the most popular view that i t serves to regulate the flow

Q C 5

Of tniittanee throtigh the sievo-artas ^sy narrowing th«

eonnoeting strands and also protects the leakage of sap

eonstltuwtits ^roui^ t^e side valls at the sieTe-areas.

Call<^6 is generally absent in tiie normal fimeticming sieve-

elements of Cueurbita but Partbasarthy (1968) correlated i t s

presence with the metabolic state of the sieve-elemmts,

Uae walls of the sieve-elements are eellulosio. ^o well

authenticated evidence for their lignificaticn is reported

{^BBM.^ 1965a). Ihe wall thickness varies, Ihe wall of sieve

tubes of primary phloem was studied in a l l plant groups by

Lesage (1391) and Leger (1397) and they observed thick glisten­

ing walls having pearly lustre. Ihey referred to thea as

'Macre*. Chau^aud (1397) found t^at these thickwnings develop

on the priisary wall of the sieve-tubes at ^ e tine of develop­

ment of siev»-plat« in i t . Ihis view was later supported \tf

many workers (Chauveaud, 1900} Chang, 1935| Esau, 1943 and

aehnke, 1971).

Ihere is soise controversy regarding the nature of the

cellulosie walls. According to Hill (1901) and c ehmldt (1917)

these *naere' walls are cellulosie in nature and beeoae ttln

as the sieve-elenent ages. 2;iBiQer8>ann (1922) described this

vai l as a layer laid over the priaary wall but he cota4 not

eonflrffi that nacre is a seoondary wall. In contrast, Sylces(l908)

23

d«ierlb«d the nature of those walls as attellagenoos and

sttggastad their occurrence in b&ih ^e si«ve>tobes «»d

parenehyna e e l l s . However, recently ^au (I965a) determined

the ee l lu los io nature of the walls of s ieve-elenents. The

nacreous wall shows certain special characterist ics . I t i s

not exceptionally hio^ly hydra ted but may shrink with the

ageing of c e l l s , ^though i t varies in thickness i t may be

so thick as to occlude the lumen very much, but i t does not

cover the sieve-area (Esau, 1965a). I t i s l o s t as the sieve

element ages. Presence of these nacre wall thickening i s not

a constant feature and Lsau (1939) reported that these thicken­

ings are completely lacking in V'itis (Csau, 1948), Whereas

i t has be<m seen in cryptogams and i t i s a characteristic

feature of the sieve-tubes of laminariaceae (Sykes, 1908) and

a lso in ^arfltllft (KUSSOW, 2JB72).

Nacreous walls show high chronaticity with stains l ike

bisaark brown (Chauveaud, 1900) or lightgreen (Change, 1935).

I t gives a posit ive reaetion to t e s t s for cel lulose and peetint

and stains v io l e t with nethylene blue (Sykes, 1908; Chang,

1935). ZiBUBemann (1922) applied a eoobination of haematoxylin

and eongo red, that stained the outer wall v io l e t and the

speel f ie inner one, red. These naere walls swell readily with

appropriate treatsM&ts (Sehaidt, 3917) and are highly hydrated

(Bsan, 193«, 1938a).

24

A prominent feature of the 8leve>elemeat i t i t s extrenely

unstable protoplast. Sieve-tube protoplasm has different

physical and ehesiical properties sioultaneously. Xhis gets

hydration and parietal protoplasm ean be eas i ly separated from

the side walls, although i t s connection with the end walls of

the 8ieve«tube i s strong in angiospenas. I t i s stimulated more

by mechanical manipulations, but physiologically i t i s rather

inactive. .Qie rate of metabolic act iv i ty i s very low, as shown

by i t s inabi l i ty to accumulate neutral red or to form zone with

tetrazoliuffi (Currier fli al»i 1955). Velten (1872) for the f i r s t

time reported the cytoplasmic streaming in phloem. According

to him this phenomenon was observed in mature sieve-tubes of

Sidft Wum& and in young sieve*tubes of MmiA. iS&ttt <?^8WfaUa

A nuBber of organelles such as mitochondria, plast ids

and starch, ribosoaes, diotjrpsomes, endoplasmic retioulun and

so on were also observed in the sieve-element protoplast during

eleetroo mieroseopie studies.

HA^9fiftfla4rii»

SAXBOB (1946, 1947} for the f i r s t t l M teteribed toae

2s

bodl«t in the sleT«-9leiB4mt eytoplatm, iiAileh sh« r«eogniz«d at

mitoehondrla. according to Mc aivorn (1957) they vere obseryed

in the sieye-element eytoplaim of JA^A YHiiftfifff CMttrmit

QoegypiuiB. Helianthus and Nieotiana but in a scattered forn

throu^out the cytoplasm of young sieye-el«nents. Ihey are

mostly spherical but some rod shaped mitochcmdria were also

seen in young sieve-tubes of Cuenrbita, rlc 3iyem (1967) also

reported that the s ize of mitochondria varies frcxn lyum or l e s s .

I t vas noted to be fa ir ly cmstant for each species*

However, /iiegler (1960) and Hohl (1950) have fai led to

locate mitochondria both in HgragXy ffi and Jfliaca but Kollaiann

(1960) and Juloy £ife QX.» (1961) observed these in the mature

sieve-elements of Pa^^^Hora and ffttCttr^l-la SlSR^ respectively

under electrcm microscope*

I t was Brioti (1373) who gave the firs^t indirect evidence

for the existenoe of plastids in sieve-tubes and observed tbe

presence of stareh in about 1 9 species , which are noetly

dicotyledons, Kuttow (1382) and fiseher (1886) reported that

the some dicotyledons and a few monocotyledons possess stareh

in ttielr sieve-tubes*

Xhe presenee of plastids in sieve-tubes for the f i r s t

t ine vat deteribtd by Stratborger (1891)* He tuggetted that

2 (J

ttaroh gr&ins d«T«Iop In ieueoplasts in some plao«8 and tamad

them as 'Starkablinder' (Starch-blinder). Ihey are observed

in the sieve*tubes of H i i t t MaHHiUkt ^ ^ » members of Magno-

l iaeeae , Hanunculaeeae, NTmphiaceae and XUJJL (Xil iaoeae). He

also confirmed the presence of the Ieueoplasts lacking starch

grains in the sieve-tubes of Aristoloehia and iifft.

iiehnke (1971) depending on the electron microscopic

studies , c lass i f ied sieve-tube plastids ihto two main types,

visj . , *.->» type plast id (storing only starch) and 'i** type

plast ld (having elaborate protein inclusions and often also

stwrch grains).

xlibosoaes are said to be the globular macroicolecules of

ribonucleoprotein (£>itte, 1961). !Ihey are found in tlie

cytoplasis either in free fom or associated with andoplasaie

reticulum. According to Esau (1972) in young sieve-eleniMits

o^ IL&l&i&» there Is typical polysomal arrangenent of riboeone

on the endoplasmic reticulum, auvat (1960) and Dnloy tJ^ §Xi»

(1961) could not report ribosomes in sieve-element protoplast

vh i le Singh and Srivastava (1972) observed them in corn phloem.

Later, several workers reported that mature sieve-elements laok

ribosomes (^gleman, 3»66$ Wooding and Northeote, 1966| Behnket

1967, 1962a, bt 0* Brian and Zhlmann, 1967| Esau «id Cronthav,

196Bt £ • • and Chambers, ]968t Evert and i>eshpande| 1969|

Zl 't

li»au, 1^72| Kv«rt Ai AI** 1*^73J Burr and Ev«rt, 1973; «ifariBbrodt

and Evartf 1974$ Kruatraehua and Kvart^ 1974X^

Many workers oisservad a number of ve i l differ ant la tad

dletyosoines in early stages of development of sleve»elements

which have retained a delimiting tonoplast (Schumacher and

Kollmanni 1959; t sau and Cheadle, 196ga, b| i::sau fil AI*« 1^62;

Kollmann and ochuroacher, I9625 iSau, 1B63, 1^72). Recently

iiingh and •irlvastava (1972) also recorded numerous dictyosomes

in com phloeci, Dictyosomes are stacks of flattened sacs or

c i s t emae , approximately circular in outline, and each surround­

ed by a vesicle* 1!hey disintegrate during the differentiation

of sieve-el^aents and may even be absent in mature sieve-elementSt

iaeveral workers have reported the presence of endoplasoie

reticulum both in young and mature sieve-elements (Buvat, I960)

Ottloy Ai Al»t l^^lf ^sau and Cheadle, 196ga, b$ li sau l iA i« ,196S)

Falk, 1962) Cheadle and Hisley, 1962{ Kehrioh, 1963) Bsau, 1963),

but tile condition in «diieh i t occurs i s rattier controversial,

Keoantly Singb and Srivaatar* (1972) observed sore endo-

pla ia io retieoluB in the developing sieve*elettents along with

2;i

the rlbosc«i«8 and dlotyoscMBes than in the matured ones,

^-Protein or a i lae bodvt

Ihe occurrence of P^proteln as densely-stained substance

in the sieve-element protoplast has become the most prominent

and controversial aspect for botanists, M'ilhelm (1830) for

the f i r s t time observed l^ese structures in Vit i s and Cueurbita^

and named tbeo as 'Slime drops', aussov (1332) also recorded

the slinie bodies in many dicotyledons, monocotyledons and

pteridophytes. Jut in monocotyledons and pteridophytes, he

noticed them as somev^at watery sieve-tube c<mtents, Stras-

burger (1391) studied the slime forraaticm in leguminosae and

terned them as 'olime bodies'. However, in a number of vascular

plants , l ike ^Qtyp^dlffla mLXMUL ( ' axe, 1966); llwdtuqi lUlSACft

(Kvert Ai AL»f 1971); yiitoffftflU9ta tBlTtttiUU (Evert (|i &!,,

1972b>; and ^/ui^ SMM. (^ingh and iarivastava, 1972). P-proteto

or sliaae bodies are considered to be lacking. Similarly, i t

i t not seen In aany lower vascular plants other than P^lvp^iny.

P-protein was not found at any stage of differentiation, even

in tibe sieve c e l l s or in parcndiyma ce l l s in secondary phloem

of ikuillB latBfll (Paliwal and i3ehnke, 1973).

Ihe role of P.protoin in s ieve-eleaents has attained a

eontroverey, Thaiae (1962) and Canny (1962) suggested that i t

play* an laportant role in the traneloeatlon proeess whiU

• o t t of the anatonistt refuted to aeeept thlt ttatoaent t laco

2:)

i t thown at l eas t two types of transloeatlon proeosses, one

involving the plants having F-protein and the other n^ieh laeks

i t . Recently, Evert, Murmanis and aaehs (1&66) found that in

Cueurbita the s l ine bodies are enclosed by a double membrane

and hence i t was doubtful that slime bodies are responsible for

the synthesis of slime* ^lime i s i jore or l e s s viscous substance,

which stains readily with cytoplasmic s ta ins , i s present in

the sieve-tube siembers of dicatyledons. ihe slime i s of a

proteinaceous nature*

^ilheln: (13S0) was the f i r s t to report the absence of

nucleus in the mature sieve->elen)ait protoplast. Tliis view was

furttier supported by a number of workers (Janc^awski, ISSlj

Kttssov, 1882| Strasburger, 1B91) Artschwager, 1924| Crafts,

1939 and Salmon, 1946, 1947} in various groups of vascular

plants.

According to Crafts (1934), Esau (1938a), Abbe and Crafts

(1939) the mechanism of nuclear disintegration occurs in two

stages involving the enlargement of the nucleus followed by the

lo s s of i t s chromatieity. iSarlier, Ssau and Cheadle (1966)

studied the aaelear disintegration in detai l and observed

f i r s t the loes of sUinable contents, followed by the f inal

die^appearanee of nuoloar membrane* Ihis observation hat

been reeently eonfirmed by Krvin and Evert (1967, 1970),

30

i3«hnk» (19e9b) and Shah and Jacob (1969). Contrary to th«

above findings, th« presence of nuclei In mature sleve^eleoents

was noted In number of plants such as Urtlca (I'lscher, 1886),

Cueurblta^ ImpatlenSy Vltls and MaeroDloer (Lecomte, 1889)

and In Strvchnos (dcott and Brelxier, 1889). Some recent data

In this respect have been given In Table 2«

In some plants tiie presence of nucleolus was observed

even after the dls.appearance of the nucleus In the s ieve-

element protoplast, Ehgard (1944) was the f i r s t to report the

extrusion of the nucleoli in Rubus and referred to these

intrusions as slime bodies. Lecomte (1889) named them as

-albuminous glcdiules. Crafts (1939) studied these structures

in ffaswarlUftf Euealvntus and Oossypium and termed them as

sculptured spherical bodies. Engard's finding of extrusion

of nucleoli in Rubus and Gossvpium was later followed by £sau

(1947). She also described that the extruded nucleoli can be

distinguished from the slime bodies and that both kinds of

inelusioDS may be present in the same element. Zahur (1959)

reported the extruded nucleoli in 14 species of dicots .

Sxtruded nucleol i having rod-like coaponents in quasierystalllne

aggregates was recently reported by Evert ^ ^, (1970).

CQMPAyiOB gft^ t

Sie specialized parenehyna c e l l associated with sieve-tube

nembers, i s termed as the conpan&on eell« I t was considered

lABLE 2

S L . NO,

??AHK or ap'CiEa MOHPWOLUaY UfOKKKR'S NAME WITH YKiiR

1« ^»fft^§

b) iAfiCU i:^£iilU&

3 , Pinus strobttB

5 . Pinal nin«a

6, S«aml>

8 , IQUm. ^••yjg'n'fc

9 .

10 . Sai l^x hianidM

necro t ic ft

ti

tt

ft

It

*

If

ft

II

H

H

NoraU.

N•erotic

Nomal

N Mr ot ic

Pao lUlo (1963)

Kvcrt and i i l f l e r i (1965)

-do-

-do-

"dOm

-do-

-do^

Murffianis and Evert (1966)

Srivastava and ©•Jirien (1966)

wooding (1966)

0* Brian and niiaann (1967)

Shah and Janat (1968)

Kvart and i>oahp«nda (1969)

SriTaatava (1970)

Irvin aad Svort (2970)

Par«i«aarlhy (19«4: (roforrad tar Bvort

3J^Mt^ 2 (coQtd.)

NO.

12 . a)

b)

0 )

A)

8 )

f)

e)

h)

1)

J)

NAME 01 SPKCXi^

ttlftl,ifi}2JLL&

;i«auola s«BBQ«rvir«i8

XaxodiuK dlatichuffi

ACT nagttndo

iiSSL «ftcct!iri^m

S,9niM», ric?iB<?».»

Q2m^s n9lmU^vA

Jttglans n i g r a

ESBUllll. l ^ B M l ^ U i ?

SaauataiM

)c) auit lUbi£&

1)

B) f i i J A m r i B i n i

n)

o)

i « U

IQjHE. ^^riLflMi

Illi&iifiicla

f t i l i t • « * * • • * •

MOHPHOLOaY

Normal

II

H

Clear t o dense s p h e r i c a l body

Lilear s p h e r i c a l body

wClOo

Dense s p h e r i c a l t o elcmgate body

R e l a t i v e l y dense S p h e r i c a l body

Spherical body soffletimes s w o l l o i

Clear s p h e r i c a l body

Clear to dwise spher ica l body

Nomal to dense •pher ica l body

Clear erunapled body

SoTMal to clear *pherlcal body, •onetines swollen

Horiftl to elear •pherleal body

smmnhmB^ graavla

waHKKR*S NAME Wim Y£iiR

i ivert e t a l , (1970)

-do-.

-do*

- d o -

- d o -

- d o -

- d o -

- d o -

- d o -

—do-

«do-

- d o -

• d o -

—do—

- d o -

T • d o -

3i

as th« general feature of netaphloeo and seeondai^ phloen of

angloapexiit but eould not be reported In the angioapermoua

protophloem (Beau, 1939). I t vat not also observed In primitive

woody dicotyledons (Bailey and Svaay, 1949)• The number of

companicxi c e l l s associated with a sieve-tube member varies from

one to several in different species and may also be variable

in the same plant (Cheadle and i^au, 1953| ^ ahur, 1959).

according to Kumar (1969) in I . Prosopis spieigera L. the

number of ccHnpanion ce l l s associated with sievfHtube element

varies fron. oae to three. CcKspanion c e l l s also vary in s i z e .

3ome are as long as the sievt^.tube members with which they are

relatedf others are shorter than the sieve-tube members* Ihe

co£Bpanl(%i ce l l s of a sieve-tube element ;:iay occur on various

sides of this element, or ttiey may for^ continuous longitudinal

ser ies on one side of the sieve-tube element* In SOBW

herbaceous dieotyledcms and in many monocotyledons having l i t t l e

or no phloem parenchyma, the conipanion c e l l s of the superposed

ser ies of sieve-tube members form continuous longitudinal

ser ies (Strasburger, 1B91), but in other plants the companion

c e l l s of different elements are oomnonly not in contact with

eacdi other.

2he va i l between the eoopanion c e l l and sieve eleaeDt i t

either taiformly thin or has depressed areas, primary p i t - f i e l d s .

Itader the eleetron aieroseope plasmodetaata are evident in

these wftUe (Btau sad Cheadle, 1:902b).

32

In oontrftst to th« 8Uv«-«l«stfnt, th« eompvaica o«Il

retains Its nueleus at maturity. At the height of act iv i ty

i t s protoplast laay s ta in more heavily ^on that or ordinary

parenchyoa c e l l s , and i t was noted that this chromaticity

increases after the con|)anlan c e l l develops beyond the meriste-

matic s ta te . Esau (1947, 1948) observed s l ine bodies in the

companion ce l l s of l i i i £ , Mbi&lA and tUM^aad noted that the

chromatidty of the companion c e l l protoplast increases after

the dispersal of Imese bodies. Ihe mature companioci c e l l s

possess diotyosomes, mitochondria and endoplasmic reticulum

(Ksau ana Cheadle, 1961, 1962ii), while they are devoid of

starch (B^au, 1965). heoent findings of Kvert and Deshpande

(1971) had shown that plastids also occur in the conpanion

c e l l s . The sieve-tube elements and their companion c e l l s appear

to be closely associated not only ontogenetically and morpholo­

g ica l ly tmt also physiologically. When the sieve-tube proto­

plasts are disorganized at the aid of i t s ac t iv i ty , the

associated ecnvanion c e l l s die a l so .

Zhe s ieve e e l i t of gyanospermi and vascular cryptogams

have ao eonpanien e e l l s . However, in QiaJLil ^<i conifers

eerUin ray and phloea parenehyaa ce l l s are c losely associated

norpholegieally and physiologically with the s i e v e - c e l l s

( S s a n ^ j y ^ . , 1953| Qrilloe and Smith, 1959; Srivastava, 1963a,

b) . Ihese parenohyma c e l l s have been termed as albuminous ee l l t*

because they frequently s ta ia deeply with eytoplasmie s ta ins ,

«• thottgh ^ e y are partieularly rich ia protelaaeeooe aaterlmls

f>

(iitrasburg«r, 1391). Whan albuminout e t i l t oeeur in Tf^

they art usually loeatad at tha narglns of rays and eonstltuta

tha araet ray calXs which ar« ta l l er and of Siaaller traaisvers*

diflusteters than the proeumbent TBJ o e l l s . iilbumlnous ce l l s

inclined among axial parendbiyma oe l l s appear to be oostly

mmibers of declining t iers ( ir ivastava, 1963b}, the vai ls of

the s ieve c e l l s facing the albuminous oe l l s have conspicuous

s ieve areas. Typically, albuminous ce l l s contain no starch

and they die ^Ai&a tdie sieve c e l l s are disorganised,

Nagali (1858) observed parenchyioa ce l l s in the phloeoi

t issues and referred to them as 'bast parenchyma*, according

to Ssau (1939) the phloem parenchyma forms the characteristic

feature of tiie primary and secoadary phlo«& of angiosperes,

gyonosperns and eryptogas*.

In primary phloem the parenchyma c e l l s are elongated

and oriented in the SAM dlreetion as the s ieve-eleaents , «hi l«

in the seeondary phloea, pareoehyna occurs in two systeB»{

the axial and the ray system, Ihe ray parenchyma wi l l be dealt

with under 'Phloem ray * in subsequent pages. She also

reported that the walls of phloem and ray parenchyma oe l l s

have primary p i t f ie lds whieh interconnect the ray and axial

parenehyaa o e l l s with one another and within themselves, while

34

in the aetlT« phlOAin, th« phloem parenehyna and the ray ca l l s

obviously hava priciary unligniflad val la . After the oessation

of the t issue aet lTlty, the parenehyoa c e l l s aiay remain

re lat ive ly unchanged for a long time, or tibey develop into

s d e r e i d s ,

Later, i';sau (1965a) noted that the phloeBi possesses

variable number of parenchyma ce l l s other than albuminous and

ccnopanion c e l l s , <^ccordiag to KuiBar (1369) the phloem paren­

chyma ce l l s In rrosoois soieigera L« are quite small and occur

in bands of three or four layers on tho lower side of the fibre

bands, ihey also forau crystal l i ferous ce l l s on either side of

the fibre bands,

'!he occurrence of nuclei , endoplasmic reticulum, ribosotaes,

p last ids , mitochondria and occasicmal dictyoscnses vas studied

in the parenchyma ce l l s of secondary phloem in Ti l ia i iwrAmil

by Evert et a l . (1965). Mitochondria, chloroplast, dictycwoaes,

end mierobodies are found to be the important c e l l components.

In some phloem parenchyma c e l l s of EugalvntuSy slime l ike

sttbstanee was reported hy £sau (1947), Becently the preseoae

of P*prot«iii in the secondary phloem parenchyma c e l l s of

FirttiBfffllitM i U i l S A sad l i l i t fiPAfia has been confirmed bjr

Davis and £vert (1970),

Aofardliif the funetion of phloem parenehyma c e l l s , Bsan

(1999) ropertod that thegr are ecnoemed with many of tho

ao t iT l t l o i smoh as tho storofo of fa t , ttareh and other orgaaio

food matoriolA and aoewnilatloBS of rosins «id tannins, Cortala

3iJ

parweiehyBtt c«l l* in th« ttoondary phlo«a aeeuiaulat* crystals

or r«sIns but die afterwards (^bbe and Crafts, 1939)•

In many plants a phellogen is eventually formed in the

phloem. I t i s produced by phloeia parenchyma and ray parwnchyma

respectively* Phloem parenchyma may or Daay not have i t s origin

fran the &asse mother ce l l s whidti give r i s e to the sieveoelements.

Parenchyma c e l l s ontogenetically related to gieve^elements, die

at the Same time as the nearest sieve-elements die (Cheadle

and i^au, 15?58| Svert, lB63b| arivastava and ia i ley , 1962),

But recent findings of jisau (1970) suggest that whether the

parenchyroa c e l l s are cmtogenetically related or not, they do

not degenerate when the sieve-element ceases to ftaicticxi,

f ibres are the z^st ii^ortant components of phloem,

mainly in the secondary t issue (bsau, 1965a). 3!he term phloem

or phloie fibres i s applied to fibres originating in primary

or secondary phloem. Ihey differ considerably in length and

are usually many times longer than broad.

Ihe extraxylary fibres are sometimes combined int« a

group teraed as 'bast fibres• as in ^ lam Msltatiseimpm and

Helianthna^ Ihe word bast was original ly applied to the

extra-oeablsl region of dicotyledonous stems (Haberlendt, 191t)»

Ihe fibres of the extra-aaBblal reglcn of diootyledenoiM i t«M

btlong, In meet inttanees, to the phloea. Ihe tera baft fibres

3C

i s S t i l l us«d for phloem fibres in refer«neos dealing vith

the economic use of plant fibres (Harris, 1954). The extra-

xylary fibres are eategorissed into tiie following headsi

Phloie fibres i , e « , fibres originating in primary or

secondary phloem and peri-vascular fibres (cort ica l fibres)

i . e . , fibres originating in the cortex.

1 ibres are elongated elements with both ends tapering,

narrow luisen and thick secondary walls (Esau 1966a, lahn, 1967;

Cutter, 1BG9). according to Ksau (1965a) they my be liipfiified,

but In sc»iie cases they are not so . ihe pi ts in their walls are

usually simple, but so&etimes they nay be s l i gh t ly bordered,

as reported by ^ a u (1965a).

Ihe prib.ary phloeia fibres occur In tiie protophloem but

not in metaphloeiu and they develop *4 en t^e shoot exhibits

aaxiauifl rate of growth (iisau, 1950). Occurrence of fibres in

primary i^loem i s a rare phcaiou>enon. However, these fibres

forming bands provide a characteristic appearance to the

seeondary phloem, ii^i<^ i s an ii^ortant feature for a genus

or speeies , e . g . , gjlfatrgUt some Bttfialyptttf spp* (Chouse and

TUniis, 1974{ Khan Ai i L . , 1978). Qie presence of fibres in

priwury phloem and seeondary phloem was also studied by

Holdhelde (1951) and Zahur (1959) in soae species of woody

dieotyledons. Qyanosperms usually have no fibres in the

primary phloem, but many have them in the secondary phloem.

Xhe seeondary phloem fibre elements in angioeperms and

gymosperms are variously arranged, e . g . , l a fiifiCA, the fifer«t

'J'-1 3v

eont t l tuU the largest portion of the phloem and th9j •nelose

among them seattered grotqis of the other phloem elements

(ATtsohvager, 1950) { in HiiUL* the f ibres, a great proportion

of which are septate (Strasburger, 1B91; is^au, 1948a), are

arranged in tangential bauads which alternate with bands of the

sieve-tubes, ootspanion ce l l s and phloem parenchyma; and In

^aurugf i^icotiana and >itgnQl9>4M tti«w are few fibres and they

are scattered among the rest of the phloem elements of the

vert ical systen^

^^ Pfuni s there i s no aclerenchyma in conducting phloem

but i t differentiates into fibres and solereids both ^sa

phloem ceases to functi^m. Recently, the distribution pattern

of phloem fibres ia reported by C iouse and Yunus (1974) In

iJalbergia^ Khan ft! jJL* (1973) in some Eucalyptus spp.{ GBiause

and Uashmi (1979) In tropical trees such as Dalonix regia^

PMlranJiu y^rtargfa ll snd pti ypftwuffi ftrrmtotittffl* Khan iii l i . (1977) fotad that fibres are to ta l ly missing in secondary

P^lO»u of iflftgiftfiHUt £lldittl» tMiSUA XfilttHLUUlf Clerodandron

iBflOtti Citr94tnar« ffPlfBd»f» OIUUDM Yar4tffili» SMM^U

arbortriltif m d XLItS negundo.

In some species the secondary phloem fibres aature in the

eondiMting phloem as specialized mechanical elements as in

^^^U- ^B other plant species they have primary vails md,

o

aativ« protoplasts in the fUnetioalng phloem end differentiate

•s fltoret only after the sieTe-elements cease to function, e.s*i

pfMBiig (Schneider, 1945). According to Esati (1948a) eTcn ^ e

s«e<mdar7 phloest fibres, like xylem fibres, may remain allTe

and store starch as in Vitis.

Phloeia fibres which do not develop directly from fusiforn

eambial init ials but fros paxenchyoia cells of non-conducting

phloen have been termed as fibre-solereids (hsau et al»,1953).

I ibre-osclereids are reported in Pvrtm malus by ^.vert (1963) and

in Acacla catechu by Clause jai ai,, (1979). iiucilagenous fibres

have also been reported in the secondary phloem.

Intrusive growth in phloem fibresi

Intrusive growth is the apical elongation or the lateral

expansion of cells which they undergo after completing their

primary phase of growth. Many workers including (Schleiden,

1B42| Kundu, 19421 3ehoeh-iiodoier, 1960; Kundu and Sen, 196l|

Liese and Parameswaran, 1972; Qhouse and Sabir, 1974; CSiouse

and Yunus, 197S| Qhouse t i t ] , . , 1975b) ;iiddiqul jjL i l . , 1976;

Ohouse and Slddlqol 1976a, b; Ohouae and Uashmi, 1977, 1978)

have reported the phenomenon of intrusive growth la phloem

fibres in a ntuAier of tropical trees.

Lieie and Parameswaran (1972) also reported that the

phloem fibres not only differ la length la a l l plaat ipeelet

3!)

•tttdl«d 1^ then but also show diT^rs* apieal struetur*. %ou9«

•ad Sabir (1974) later eoofiraad this Idaa and suggested that

f l toe eleffi«3ts censlst of rarloos types of apieal features

sueh as bilateral and unilateral forkings, subaplcal branching,

serrations, and deep depressions due to special growth oharaeter-

i s t los . Ihe Intrusive development In phloem fibres takes place

by one or both ends resulting in mono or bipolar growth, Sie

intrusive growth is very coiinion in vascular elements of pri'riary

and secondary origin. Ihe study of Cheadle and Ksau (1964)

fonss the Isolated report of the literature regarding the

absence of intrusive growth to ; At3rP4ffll4roi;»

Phloem rays are the Important constituents of tti9

Secondary phloem and cmstitute the horisontal system. Xhe

phloem rays developing outward are continuous with Ihe xylem

rays sines both arise froa a coonaa group of ray init ials in

the oaablam. Ihe phloem ray and th% xylem ray together eonstl-

tttte the vascular ray. Near the cambium the phloem and the

xylem rays having eoancn origin are usually the same in height

and width. However, the older part of the phloem ray, lihleh

i s displaced outward by the expansion of the secondary body,

may increase in width, sonetiaes very considerably (Holdheide,

1951). Phloem rays are uniseriate, biserlate or multlserlato

•ad th«y i^ry in height. Qie rays may be composed of one klad

40

of o«lI , ! • • • , hoffloganeous, or they may contain two kinds of

e«II , proeumbant and eraet, and ara known as hetaroganeout

type of rays. In dicotyledonous secondary phloem, crystal

formation i s very coiaiaon and occurs in the rays , while the

seccmdary phloem of conifers s ay contain resin canals vfhic^

occur in the rays in tXssA canadensis (Ihomson and Sifton,

1925)• Aie di latation mechanism of the rays in non-conducting

secondary phloem was studied by Holdheide (1?S1) and Schneider

(1952). J3ie di latation of phloem rays, a form of non-caatbial

secondary growth, i s considered a coauawi feature of various

Species of Citrus and 111 la C^chneider, IBSti^ 1354, L955 and

iisau, 1966b), Sisau (1965b) differentiated such type of growth

with cassbial secondary growth by introducing t^e term inter­

calary, secondary growth to dl latat ioa of rays in the stems

and roots outside t^e vascular eambium,

%e degree of ray dilatation highly varies from spaeiat

to species , witiiln a tree soma rays baooma dilated i4iile o^ars

do not, or ca l l divisions nay be seen only in a part of a ray

(Schneider, 1955). In fiUail. JUOfl&iil. the phloan ray ca l l s

f i r s t stratoh tangantially and then undergo divisions by radial

walls to form masses of paranehyma tisaua (^chnaidar, 1952),

Aoeordiag to ^ehnaidar (1955), fomatioa of aipansion

t i s sue takaa plaea hy the act iv i ty of a naristam foraad in the

rays, ihiali h% eaXlad *dUaUtion aarlstaa*, l i i i la in Pyaaonla

4i

•nieigaya L, no such iMriatttm was obs«rT«(l (Kumar, 1969). The

forffiatlon of •xpazxBian t lssu« in this plant i s due to the

tangential stretehlng of the ray ce l l s followed by antic l inal

div ls ioas . m i s t issue has been termed variously by different

workers, such as 'di latat ion tissue* (Schneider, 1955) and

•phloem expansion' (nihitiaore, 1962a, 1963). But i t wi l l be

apt to terffi this t issue as 'ray expansion*•

uclerelds are 0X>at widely aistrlbuted in the plant body

(Je iary, aS34| Haberlandt, 1914). Ihe cortsx and the pith

of gyn»iOsper£tis and dicotyledons often contain aclereids,

arratiged singly or in groups* ^clereids are also consnon

components of phloem v^ere they iaay iiitergrade with fibres

(Ssau, 1965a; Ohouse jftt ^ . j 1975a, c; f ^ i a n ^ ^ . , 1976, 1977,

1978; iChan Shahnax £^ jJL*, 1977}* In oany plants the inter­

fascicular parenchyma ce l l s located between the strands of

primary phlo«n fibres develop l ign i f i ed secondary wal ls and

differentiate into solereids which, together with the fibres

form a eontlBUous sclerenehyma cylinder on the outer periphery

of Taseular system, Ihe plants in which a continuous scleren-

ehyma eyllnder i s present in the primary s tate may show a

di8rtq>tlan of this cylinder when the Taseular system surround-

•d bgr the selereachyaa l&ereases in eireumferenee throu^

•eoondary grmtttkm Oie toeeki in the te levtn^yaa e y l l i i 4 « are

42

f l l l«d vith par«nehyna eel l t whioh lat«r may differ«ntl&t«

Uit& fteldselda, a.g*, Ariatoloehla.

ael«r«id8 vary widely in shape, size and eharaeteristies

of their vai ls , Coianonly the selereids are bradiyselereidSi

l,e.,3t(»ie ce l l s , short, rou^ly isodiametrie selereids,

resembling parenchysia cel ls in shape, widely distributed in

cortex, phloem, pith of stems« 3clereid forms may be oonsi*

dered as the characteristic of species and isay therefore be

of taxcsaoiaic value ( Oarua and Datta, 1959j Gfcouse ^t e^l., 1976a,

b ) .

Ihe secondary walls of the selereids vary in thickness

ia niany species and are typically l ignified. If the walls are

relatively thin, the selereids cannot be definitely separated

from sclerotic parenchyma and fibres. In roost of the selereids

the luffilna are alniost obliterated because of massive secondary

wall deposits, and the wall shows prominent pits , often with

ramiforn eanal-like cavities, Ihe pits are conaonly siaple,

but soBetiaes the secondary wall slightly overarches a snail

pit ehaaber. Ihe seecndary wall often appears eoneentrieally

lamellated in ordinary and polarised light. Ihe lamellatlon

may be the result of an alternation of Isotropie layers of

eelluloae (Bailey and Kerr, 1935). Crystals are enbeded la

the seeondary wall of the selereids in eertain species (Bailey

•Ad l«0t , 1948| Xnaodar and Otngadharan, 1974). In soat

telert idt tiie Aeposition of seoondanr walls is uneven.

SelemiAs MT either roUln their protoplasts on reaehiag

43

0«splte the avai labi l i ty of •oluoinous lit«ratur« on

cambial act iv i ty our knowledge i s scanty regarding the seasonal

productloa of secondary phloeis. iiccording to Esau (1966a)

this might be due to the lack of exact technique required for

i t s investigation, Soine eMnent workers including Janesewski

( l a a i ) , ilussow (1832), iitrasburger (1B91), Knudson (19X3),

Schneider (1S17, 1945, 196ii), dwarbick (3527), Gill (1932),

Priest ley (1935), i i l l i o t t (1935), Kaau (1939, 194£), liubor

(1939), Holdheide (1951), .<ilcox aSt a l . (1355), Jri l los and

umith (L959), invert (1^)60, 1963b), Derr and ,.vert (1967),

Al f ier i and ^.vert (196B), Xucker (1968), iJavis and i.vert (1968,

1970), rucker and ^vert (L969) worked <m this aspect and

confined their studies to only seasonal cycle of phloem in

tesyierate species , iut the information about ttie seasonal

developBont of secondary phloeu in tropical trees i s scanty.

IfawtOQ idad Lavton (1971) studied t ie seasonal v&riation in

the secondary phloem of some tropical forest trees from Nigeria

(AlbJgala ttf4llltt\tf?ltii Antiaris af^ieanay aombax bucpopoaenaey

^9lirrh«ii fliWllwn^it 4kte9a<n4r9B hi^tlfttiai and itfil&sBft l£andU)«n<l fouB<l that only in two genera (ItftiSSBA and UsJS^

dendroB) the eaabial act iv i ty ceases during the dry season af

Indieated bgr the laek of aetlve phloea while in a l l other eases,

the active phloea vas present throughout the dry seasoa.

44

S«asQn«I produetlon of secondary phloem is also reported in

Miaaeaae elMigi Xxy €house and Hash mi (1979). Sotae more

ifflportant findings regarding this aspect are suimarized in

Table 3 .

In angiosperms the seasonal tiialngs of production and

differentiation of various elem^ts of tiie secondary phloem

•ary considerably (iCozlowski, 1971),

«

0*

%

r/5

Q

a.

CO

a 8 OS

^

is

u

It

I SI

I

lis 55S

I 4 > <«k 4 ^ *Mr

I I :5 i

2 8 8 I ^ *» s I I 5

I 5

I 8

o

52^ ^ S

I

1 a 51 • 4» It •4

1 a z to

III 8 S I Q Q n

5 H H H >> ^ ^ «rl •-< »« •« l« *g ^ ^ •9' m

I 5

o 4«

! l

n 4*

I Si

lis lie

4»

§ 1

•o •rl X

1 *t H

m

a 8 4» A • ta

d

4*

3 DO

3 1

n •H

«4 t4

^

1

I

e I

^

t i 1 I

B o

Si

^

Its

8 Ss

IK

4*

I

2 |

In xm a ss*

O 01

a

o

IIS 1

I r-i

4» m

I o

M •.7 SI a

I l4

I CM

4 . *

I I I f 8

»4 ^

Mst

u

I

si

I I CB (9 4B IB IB B §* o

aj

a

o

I I - I

I o 6 In

11 I

O S

51. 2

S • A M 1. ^ s l

f-« N

4 10 'O'O t4<t( i 01

o

a •& 4«

o

• o

•Otf

SI

• 0 I" a ai

A. 09

3 ' *

4;j

Sine* lmotfl«(ig« r»gardiiig the produotlon of Mcondary

phlo«B is m«agr« •••n less la known about tho longovitjr of

tho socondary phloon (Priftstio/, 1930{ Bannan, 1955| Warolng,

1953a, b| Chovdhury, 1968| Falival and Prasad, 1970{ Fallval

tiftiUt 1975} Xunus, 1976$ Hashnl 2977 and Khan, 1977)* Iho

term longevity applies only to the funeticoiing of the sieve

eleaents,

'Sie Icmgevil^ of the phloem varies fr(»3 species to

species. In majority of the dicotyledons the functional part

of the phlegm i s confined to the secondary phloem t^ich is

produced in last grovth season (lahn, 1967)• Hovever, in some

plants the entire phloem produced in the previous season ceases

to function before the cambium begins to produce new phloem

in the current season (Esau, 1965a). I^XlSL snd Vitia were

fotmd to be two exeepti(»s in whidi the phloem bears longevity

of more l^an a year. Phloem was reported aetive for 10 years

^ Ti i ftSCdAlyiL Mill. (Holdheide, 1951) and 1-5 years in

T^lia ameyjaana I«, (Bvert, 1962). In j m i £ i t remained aetive

for two seasons (Esau, 1948).

In plants having anomalous phloem,e.g., flgBgitoTUltft

and some woody species of Chenopodiaceae phloem remains aetive

for many years (Fahn, 1967). While In most conifers the

longevity of the phloem i s restricted to a single growing

season (Bsatt, 1966a).

4(;

In th« ttajority of th« plants studied, th« phloen b«eoM«t

non»f«ietioaaI in th« IMM scatoii In vti l^ It was darlTsd from

ths oanblUB (isisau, 1939, 1945{ Hubar, 1939; Holdhalda, 1951|

Davis and Srart, 1968, 1970}• Savaral vorkars had danonstratad

that in S08MI woody diootyladons, oocasionally small siava-

al<nBants ranain functional tmtil new phloam was produoad in

spring. In soma speoias such as paean (iurtschwagar, 1950),

yallow birch (Mfileox Ai Al«t 1^56) and soi&a oonifars aspaoialljr

in PiPPS s^yobus (Strasburger, 1S91) Jrown, 1916) Abba and

Crafts, 1939) Grilles and Smith, 1959| Srivastava and 0*Brian,

1966) the later formed sieve-elements at ^ e close of the seascm

undergo only partial differentiation and overwinter in tiie sane

condition so that these undifferentiated phloem c e l l s take up

the function after the differentiated phloem becomes function-

less and before the new cel ls are formed l^ the a c t i v i ^ of

cambimi. Ihis typa of partially differentiated overwintering

phloem was not reported by Alfieri and Bvert (1968) in EiQBl..

Many workars including Coekerham (1930), Ell iott (1935) and

Hubar (1939) did not find any differentiating or partially

differwutiated sieve-elements in winter in ease of Mftt* ^n

addition they also reported the complete abswioa of the

funotlonal sieve-elaments at the time of eambittm reactivation

in spring.

According to lueker and fivert (1969) sieve-alemiot

differentiation occurred the year round in A^^T naauniio-

47

P«ri<l«rB is a prot«etiT« tlssu* of ••eondary origin*

Porldarn fomatlon is a eoomoa phttnoDanoo in starns and roots of

dieotylodons and gymosparns that ineraasa in thieknass by

saeondarjr growth* "She paridars usually consists of following

thraa eoopon^it alamantsi tha phallog«a,or eork eanbiun, t^a

naris ta% producing the peridarmi tha phalXam, cosimoaly eallad

cork, producad by tha phallogan toward the outsidai and th9

phallodarsi, a tissue Idiat resembXas cortical paranchyaa and

consists of the inner derivatives of the phellogim.

Iha term periderm can be distinguished froa tha non­

technical term bark, AS discussed earlier, Uie tern bark in

i t s restricted meaning refers to a l l the dead tissues together

with the layers of peridara. If bark is used with r e f e r e n t '»<

to a l l tha tissues outside the vascular eanbiua, the paridara

and tha tisanes of tha axis isolated by i t aay be eoMblnad

imdar ttim designation of outer bax^* Iha ta<tfmieal term for

tha outer bark ia rhytldOM (Da Bary, 1B84). 2his tarn la

darivad fron tha Oreak word aaanlng wrinkle and refers to tba

appaaranaa of the outer bazic irtian i t ecmsista of layers of

eork alternating with layors of tissue cut off bjr the eork.

Paridara eharactarittieally appears on the surfaea of

roots and staas and thair branches la gywioaparas and woody

diootyladoBS that uadarge a eontinuoaa pronoonead iaeraata la

thiekn*»s by Stteondarir growth, P«ri<l«rB occurs alto in

herbaceous dieotylodoiit, In which It i i tonctioAii l i s l t«d to

th« oldost parts of stoa and root. Monocotyledons rarely

dsTelop a protective tissue comparable to the peridera of

dicotyledons. Foliar organs normally produce no cork, but

scales of winter buds in SOILO gymnosperns and dicotyledons are

an exoepticm. In t^e imderground stcmis of some vascular

cryptogams, the epidermis or the outer cortical layers becone

suberised.

£sau (1954) reported that the external appearance of axes

bearing periderm or rhytidome i s highly variable. Ihis varia­

tion depends partly on the characteristics and manner of growth

of the peridera i t se l f and partly on the amount and kind of

tissue separated by the peridera trom the axis.

If the plant has only a superficial perider% a relatively

small amount of primary tissue is cut off, involving either

a part of or the entire epidermis or possibly one or two cortical

layers. Xhis tissue i s eventually sloughod away, and phellem

i s exposed and the stem in this instance i s considered to have

no rhytidcme. If the exposed cork tissue i s thin, i t commonly

has a smooth surface and i f i t i s thick, the surface is oraeked

and fissured. Massive eork usually shows layers that seem to

represent annual inereaents.

4;i

Ih« d«*p«r p«rid«rBUi cut off larger anoantt of tho

original stea tiasuos and usually form a rhytidoaa. Iht

eharaetaristies of the rtiytidom dapand on tha nannar in lihieh

It faparataa ont from tha traa (Ssau 1964b). In SJJBUL

ftaarieanaf Magnolia MWUliti and Cilg rffBJ f» certain rhytidomas

ecmsist mainly of par«a<rfiyma tlseue and soft oox4c ce l l s , i4iile

in others, a.g.t species of Qneroas and QaiSA ^ « riiytidcae

ecmtains large amounts of fibres (mostly phloem fibres) vhich

are associated witih, hard cork cells« Ihe manner of formation

of the periderm influwioes the shape of the bark in g wieral

and of the ziiytidoine in particular* When the subsecpent

perideriss develop in the form of overlapping scale-like layers

or shel ls , the outer tissue breaks V[p into units related to

the layers of the periderm, and the resulting outer bark i s

referred to as scaly bark. jQiia type of bark occurs on

relatively young stems of £i&)at EJOM. OSmmil «nd ££2tfiftii

spieigera L. (Kumar, 1969) while in HUs. , Lonieera, Clematis

"^^ Cunrassns, the subsequent periderms are formed as entire

cylinders and so the dead outer tissues are sloughed as hollow

eylinders from the stem. Ihis type of bark i s termed as

ring bark. Zhe bark of Platanua^ iupbutas and species of

gUgitTBtlii i* considered to be intermediate betwe«i the scaly

bark and the ring bark as in these plants the outer layers of

the bark peel off in the form of relatively large sheets.

She manner in i^lch the dead tissues separate from the

stem is determined also by the nature of the periderm ( Oe

so

Bar/, 1B84{ Mtihldorf, 1926} Pf«iff«r, 1928) • In • on* plants

the ••paratloQ oeenrs through thin*walled eork ealls^e.g.,

4gbtttu» and Platanni in which the dead tiaiua separatat from

tha parldern In tha forn of large, thin seales through the

outer thin-walled layer of cork, while the subjacent thick-

walled cork cel ls renain attached to the stem ^ i e h , therefore,

has a smooth surface. In species of Euealvntus the sheets of

dead outer tissues of the bark exfoliate through layers of

parwichyna cel ls with onthickened walls, which occur on the

periphery of the phellem (Qhattaway, 1953; Khan and Ohouse,

1978)• In some trees^e.g*, tMM,* ^ ® inner bark grows slowly

and, therefore, louch expanslc«i tissue is formed* In this ease

the subsequent periderms cut off sunall uKOunta of seoondary

phloea and the sloughing of the outer bark is slaw, resulting

in the fal l of minute scales and even powder (Whitnore, 1962b)*

In many plants the periderm cel ls show considerable

cohesion and the succeeding layers of rhytid(MBe adhere to one

another and remain as sudi on the stem for many years* In

this way, the outer bark becomes very ^ ick and deeply grooved*

Such type of bark occurs in Sennoia seaBerwirana (Isenberg,

1943), certain species of j&iuuuua, jfjtlil&i ^iU&t AsULlUiL snd

ZJUUiSL, Ihe loose and fibrous bark is found in certain

fimlyplli species (ChatUway, 1955)*

0 i.

Ph«llog«iii

%• phellogcm i i a secondary nerlsteoatio tissue. I t

originates from ce l l s t^at have undergone differentiation and

i t produces tissues that comprise part of the secondary plant

body. Jhe phellogen occurs as a lateral meristen because i t

results in an increase of the diameter of the axis by periclinal

divisions in i t s ce l l s . Histologically the phellogen is sinpler

than the vascular oanblum as i t consists of tmly one type of

in i t ia l s ( ce l l s ) . Ihese phellogen cel ls appear rectangular in

cross-section and somewhat flattened radially, and In longitu­

dinal tangential section, they are seen in the form of regular

P<^ly^QaM, Iheir protoplasts are vacuolated to varying degrees

and fflay c<mtain ehloroplasts and tannins. Ihere are no inter­

cellular spaces in the phellogen except In those regions vrtiere

lenticels develop.

!Die phellem c e l l s , i . e . , the cork cel ls are usually poly­

gonal in tangential section, while in eross-sectlcn titiey are

flattened radially. In eross-seetion, the cork eel ls are usually

arranged in oMipaet radial rows which are devoid of Intercellular

•paees*

Gotk ee l l t are known to be dead ce l l s . Various types of

«Ofk e«IIs can be distinguished and in a f«v plants crystal-

containing cells and sclereids may also be observed among ttie

coriE oel i s . ^luetines noa-snberised ce l l s , i . e . , phelloids

occur in the phellem. Xwo coonoa types of cork cells are those

which are hollow, thin-vailed and soiaewhat widened radially,

and those which are thick-walled and radially flattened. 3he

cel ls of the latter type may often be f i l led with dark

resiniferous or tanniferous substances as reported in liuealvDtus.

ihese two types of phellem cells may occur together in the saoe

plant as in iirbutus and Jetula where they occur in alternating

layers.

Jhe primary wall of the phellem cells consists of cellulose

and may scmetimes also contain lignin or suberin. Internally

the primary wall is lined Iqr a relatively thick layer of

suberin, trtiidi is termed as the suberin lamella. In certain

plants a lignified or thin cellulose layer may be present on

the inside of this lamella. In the thin-walled phellem ce l l s ,

this inner layer of cellulose is absent (Eames and Mac Daniels,

1947). Ihe suberin lamella is impermeable to water and gases

and witiistands the acticn of acids, Ihe protoplast of the

phellem cells i s lost after the various wall layers have been

formed and the ce l l lumen becomes f i l led with air or the

pigmented substances mentioned above.

la the phellem of sane plants, e .g. , species of Hologvlon

•nd inihitUi bands or large groups of hollow, thi^-vaXIod

ee l l s oeeiir among the usual thin-walled cork ee l l t . Hies*

do

e«Ils hm a llgnifittd prlnary wall and a thick outar layer

of aaeondary wail on tha inalda of wliieh i s a ralatlvaly thin

subarin laaalla. Ihis subarin lanalla i t liaad by a thlnniah

eallulota layar which may sometines be lignified.

PhillpgtrBt

Xha phelloderra colls are the living cel ls with nan*

suberi2ad walls. Xhey aro similar to the par«iehyna cel ls of

the cortex, ait they are nstially arranged in radial rows, i f

the phellodeno i s multiseriate. In certain plants t^e phello-

derm cells contain ohloroplasts and are photosynthetie. Zhese

ce l ls may also store starch. Sclereids and other special cel ls

are sonetiiaes observed among t^e phelloderm ce l l s .

With regard to the origin of the phellogen, i t is

necessary to distinguish between the f irst peridem and the

subsequent perideras which arise beneath the f irst and replace

i t as the axis continues to increase in circmference. In

the stea the phellogen of the periderm may be initiated at

different depths outside the vascular caabiun (Metcalfe and

Chalk, IdSO), In nany steas ,e .g . , Sjdaum MftUMkTtt SauaQO,

£iltt£ »A l i c i m olaander^ the f irs t phellofen i s foraed in

the epidemis i t se l f . Kere eoaioBly ^ e f irst phellogen

develops in the layer of cel ls iHsediately below the eplderaia,

'di.i

• • S M PopqlM*« JgglMis and ]|]jBi!.* In th« potato tubar, tht

pballogen davalops In tbt •pldermls as wall as in the sub-

•pldarmal lajrar, but tha phallogan fornad in tha apidarnls

does not eontlntia to ftnotloa after i ts formation. In the

stems of certain plants,e.g. , iallilLUk Plf^49igMU> spscies

of itf4'»Wl9A;^ft «n<i OaU&9 toe f i w t phellogen forms In the

second or third cortical layers. In IhoJa^ Punieay Arbntas,.

Vitls and ^abasis the phellogen arises near the vascular

region or directly within the phloem. In roots of lormnosperBS

and diootyledcms, the l i r s t phellogen i s formed in inner layers,

tasttally in the perioycle. kAille in the roots of monocotyledons

the f irst phellogen usually develops in t^e outer layers of

the cortex.

If 8Ubsequ«3t periderms form one witiiin the other, one

additional phellogen may be produced in each growth seascm.

Thtt later*formed periderms each develop deeper in the cortex

or primary phloem and, with continued seecmdary thickening,

deeper and deeper within the secondary phloem, JUM a result,

two types of formation of subsequent periderms may be dlstiag*

uished. In those plants in which the first-formed periderm

develops in an inner layer,e.g. , SliX»^ the additional

periderms usually form entire cylinders similar to the f irst -

formed periderm, while in plants in which the f irst periderm

i s formed in the epidermis or the outer layers of the eortex,

• • ! • • E BttLt the additional periderms develop in the form of

•eales or shells , the eoneave side of whieh i s direoted outwards.

dU

In e«rt«in g«ii«ra the subsequent peridens slres<iy begins

to develop in the f irst year of growth of the stea or brsn^.

In apple and pear trees subsequent periderms begin to develop

in the sixth or eighth year of growth and the f irst phellegen

may even remain active for about 20 years (Evert, ir)63). In

Pnniea and in a few speeies of Populus and HcmM the f irst-

formed phellogen may remain active for 20 or SO years, while

in iiiufiXSBS. wmaZf some species of lasMt «« &>?4f and EftlSaxlSD.*

and in a few o^er genera and species, normally no subsequent

phellogens are formed during the l i f e of the plant. In Quergus

suber and other species of f irst phellogen is active throughout

the entire l i f e of the plant, or for many years, there are

seasonal differences in the types of phellem cel ls produced,

AS a result of tiiis, bands, i^ich can be regarded as annual

grow^ rings of phellem, develop. In bottle corks sudi annual

rings can be sewn.

With the formation of each subsequent periderm the

tissues exterior to i t become cut off from the nutrient and

water tupply and, therefore, die* AS a result of this a hard,

outer crust develops on the periphery of the axis. Ihis crust

inereases in thickness due to the addition of further cork

layers which enclose pockets of cortical tissue and dry phloem.

All the eoric layers together with the cortical and phloem

tlssueSy •sternal to the Innermost phellogen are termed as

rhytldome or outer bark. The living part of the bark insid«

the rtortUMM U often teraed as the inner bark, tflth the

di)

lner«as« in dl«a«t«r of th« ••eandarjr zjrl^a th« elreiUif«r«iio«

of tho eaabial oyllndor onlargo*. AS a rotnlt of this , tho

ii«w-foni«d lajort of ioeondur sqriom ar« largor in eireiuiforoBeo

than tho outor lajrors of tho Innar bark uhleh aro» thoroforo«

brott^t undor strain. 3his strain is aoeonaiodatod bjr tho

prodtaetian of oxpansioo tissuo and proliferation tissuoCWhitaoro,

1962a). Kxpansion tissue i s an intercalary tissue fozved

oainly by the phloem rays, and proliferation tissue develops

as a result of l^e proliferation of axial phloea parendiyna.

DEVSLOPKMir 01 P/illDERMt

The cel ls of the epidermis, the eollenohyma, or th9

paren<^ysia that init iate per idem are living ce l l s , and their

change into phellogen is an expression of t^eir ability to

resuae aeristeoatic a e t i v i ^ under appropriate conditions.

Ihese cel ls undergo periclinal division, itfith title coBB«ieeBent

of these divisions starch and tannins are gradually lost froa

those cel ls that contained them, AM a result of the f irst

periclinal division two siailar cel ls are foraed. Frequently

the inner of these tvo cel ls divides no further and is then

regarded as a phellodera ce l l , while the outer functions

as the phellogen cel l and undergoes a periclinal division

resulting la the foraation of two ce l l s . Ihe outor of these

two colls differentiates into a cork coll and the inner cel l

constitutes the phellogen in i t ia l and contiaues to divido.

r • r<

Sea«tlB«s th« Gork and ph»Ilog«D oellt art foraad aft«r th«

f irs t dlTUioa and then no pbellodarin call Is fornad. In

addition to parielinal divisions tha ini t ia ls of tha phallogan

undargo occasional antielinal divisions so that tha oireunfaranea

of tha oork cylinder ineraasas continuously,

fha nunbar of pkellaiB layers i s usually graatar than

the number of phallodarm layers. In certain plants, thm

ph e l l Oder a i s couple taly absvit but in many plants i t consists

of cme to three layers of ce l l s , i^i le in a few other plants

i t i&ay be upto six layers ^iok« She number of layers in

phelloderm may also alter vith ^ e age of the plant,e.g. , in

r i l ia , the phelloderm may be <me ce l l deep in the f irst year,

two in the second, three or four later, Xhe subsequent

periderms fors^d beneath the f irst in later years, contain as

nueh phelloderm as the f irs t or l e s s , Xhe number of layers

of phellem cells produced during one year varies from 2 to 20

in different plant species. If the first>forffied periderm

remains on the axial organ for many years, the outer layers

of eozic become eraekei and are shed so tiiat the layer of oork

remaining on a plant is of more or less constant thickness.

In aertain planta, such as Quaraus £BlttIL u d Arlatoloahia,

thick layers of cork are added to the surface of the stems,

Ihe first*formed periderms, which ara replaced bgr more internal

pariderms, are relatively thin and contain only a faw layers

• f eork ealla. In moat dieotyledons aad gynoaperas the f irst

p«rid«rai ttsoallr d«T«l^t in th« f irst fmr of grovth of tlio

axial organ on those portions whidtt havo ooasod to olongato,

Paridorn fornod on young organs dovalops at tho SAM tiaa a l l

ovar tha eircunfaranoe to form an mitira e7lindar« whila

periderms that ar* foriaad on older organs usually start to

deYolop in isolated pat^es fros vhieh the activity spreads

laterally, and i t aay take some years t i l l the eork tissue

foriBS an entire cylinder.

2!he classical studies on periderm pri:arily dealing

yith the naorphologieal and anataoioal aspects of the cork

ce l l s , generally neglected tho origin and mode of development

of the cmoerned meristem* ^ a result, the literatiure

regarding the developm^ital details of the periderm is s t i l l

meagre. Ihe limited work done in the past (Ghattaway, IdSS,

1955I Liar, 1955I Schneider, 1955t 3owen, 1963; Ssau, 1964b|

iahn, 1967| W a i s e l i i j l . , 1967; ArxeeAliAl«« 1958, 1970{

Ahmed Ai Al*t 19< f Ohouse and ITunus 1975| Khan and Ohouse,

1978) elearly indicates that s i t e , time of origin, duration

and mode of activity of phellogen show quite a vide range of

diversity amoog the different plants. Hmee there is Moh to

be known to mderstand fully the phenomenon of phellogen activity,

59

For each tr«« tpselM aTallabl* around Si opal at loaat

five, adult nornal troos of con^arabla aga and •igour v i l l b«

soltettfd within tha foratt area and tagged for nunbering

aerially. The trees with abnormal growth due to shady or poor

so i l conditions wil l be avoided since they are likely to show

alow growth and ot^er physiological disorders*

Ihe bark samples will be collected In tixe form of blocks

of 1 to 2 cm square, covering the east, west, north and sooth

sides of the selected trees f om the main trunk at chest height

(about 1.6 metres from the ground level) vitii the help of

chisel and hammer, li/hile removing the blocks, care will be

taken to get the cambium intact alongwith some sapwood. Gara

will also be taken to engage smallest possible area on the

bole so that minimum injury is caused to the tree. Colleetions

will also be made from small twigs for the sake of eo^^arison.

Sie bloeks, soon after their removal from the trees v i l l

be fixed, on the spot, either in F.A«A«^ or Craff IXZ « Ihey

wi l l further bo aspiratod for the free aeeess of the fixativw

60

into th« dMp-Iyinf tltsuos aft«r reaching the laboratory.

Aftor a wook, tha f ixad oiaterialt will ba tranafarrad to 70J(

athanol or to a Aleo-glyearol ndztura^ for prasonratioo,

Xha fixed naterlals v i l l be washed thoroughly in running

water and sectioned in the following two ways depending on tiie

nature of the material and purpose of study»

a) Xbe woody hard material will be seotioied at a thickness

of 10-12/u in transverse and radial and tangential longitudinal

planes on heicherts slidin,^ roicroto^e.

b) aectloning on rotary ndcrotome after conventional wax

embedding will be undertaken to study the details of phioio

coipciients in series.

1* I'.A.A. (fornalln-aoeto*alcc^ol)t

Olaeial aoetie acid ••• 5 ee Forvalin ••• 5 ee Kthyl aleohol 95j( ••• 90 ee

2, Craff n i t

li Chromic acid . . . 30 ee 10^ Aeetle aeid (not glaeial) ••• £0 ce Foraaldehyde 37-40^ (aquous) •• . 10 ee Water . . . 40 ee

8* Aleo»glyeerol aixturet

This aixtare eOMprises of eqnal vol^aes of ethanol and glyeerine.

Ox

Stainst

'Ih«r« «r« a larga naBb«r of stains that will ba ttssd

alona or in oonbination dapanding on tha natura of tha plant

aiatarial, 3ha following ara tha eomon stainst

it., iQT tha study of baxk

( i ) naming's Xripla stain

(11) Haidanhain's Iron Haamato^lin acd iiismazk brown.

( H i ) Haidwihain's Iron Haamatoxylin and aafranin.

(iv) Xamiie aoid-i''errio ehlorida-iiacmoid*

(tr) £>afranin and 'tast gra«i*

(vi) FOB tar* s Tannic acid-Ferric chlorida.

B* For the study of maearatad tissual fibras and siaTa-alemants,

For Biaearatad phloem elements, Astra blue will be used

for sieTe-tube elwaoits and Safranin for fibres.

In a l l tha eases the material will be d^ydratad in ethyl

alcohol series and passed thr<Migh clove oil-xylene for mounting

in Canada balsam.

Ihe schedule for staining with differant stains i s

described belowi

( i ) Fleming's Triple iiUin (Johansan, 1940).

I t i s ealled triple stain because i t iaTOlves three

stains, Safrania uhieh stains lignified call walls red{ Orange

a nhiA sUins oytoplasm orange and Crystal violet that stains

e«IlBie«« wall* rlolat.

62

Svetiont In 30ji aieohol

Safranln^ < 4.24 hrs* )

1 30^ alcohol

Crystal violet^ (10 min.. 1 hr. )

i 3-dbiangas with water

30; aieohol

50^ alcohol

70^ alcohol

90^ alcohol

i 9Sli alcohol

I

Xjrlana 2-ehangaf

Hinsa with xylana (plppatta)

iiinse with elova oi l • Xylena (pippetta)

Diff«r«ntiata undor aderoaec^a

Hood with fVaah elova O I I A

y ashing with used elova oil*

Drain off orange S

r Hood aeotions with

Orange CP /

^ 2*3 changes in . anhydrous aieohol

2, Safranlni Solution will be aade bf dissolTing 2.25 ga dye in 225 ee of 95^ aieohol and diluted with equal amount of water when needed. I t stains eutin, ehroaatini lignln and in soae oases ehloroplasts.

3* Crystal Tiolett iS solution prepared in dist i l led water i s sufficient to work. I t stains tlie cyt^Iasn and nucleus. I t frequently overstaias the cellulose walls violet .

4« Orange Ot i f solvtlen of the dye prepared la absolute aieohol i s used. I t is a useful eytoplasaie stain*

(11) H«ld«nhaln*s Iron Hatnatoxjrlln and BUaark broim

(Johansan, 1940).

Saotlona In dl>tIliad vatar

i Iroa aluo^

(S 2Blnuta8)

I 3~diangas In dlstl l lad

Xylena

I 1 iilestalnlng vlth 2%

Iron aluQ ( 1-2 mln.)

3«changes In dist i l led water

30^ alcohol

50^ alcohol

I 70^ alcohol

i , Blsaark brown' (UlSi hra.)

Cloye oil • Xylene

Absolute alcf^ol *• Clove oi l 4* Xylene

Haldenhaln's^ Haematoxylln Absolute alcohol

t 90;l alcohol

Washing with 70| aloc^ol

5. Iron alUBt (lerrle AmonlttB sulphate}! To be used as aordvit or a fixer to the stain* 2% solution of Iron alua Is prepared in dist i l led water,

6. Heidenhain*s Haeaatoxyllnt 0*5% solution of the dye will be prepared bgr distolving the stain in 100 ee of dis t i l led water* Ihis stain has l i t t l e or no affinity f»r tissues unlets iroa(always in the farrie fbras) or aluniniun i s prasent la the later«

7. Bis rk taroini Salution will be prwared bgr diasalviAf 1 t of the dye in ]0O ee ftf TOJI aleeiial* St t talai tHa

f ' 'i

( i l l ) B«ld«nhftiii* 1 Iron Hasoatoxylin and Safranln

(Johanstn, 1940).

%• tehvdule for this stain i s the same as the previous

one exo^^t ^a£ the disoark Inrown i s replaced bgr aafranln

1*12 hrs* after treating the sections with 30^ or 50% alcohol,

^afranin will stain lignified walls red while Haeoatoj^lin

stains the oytoplasoi grey and middle lamella black or bluish

black*

(iv) Tannic aeid^ •> lerrlo d^loride-Lacmoid stain

(Cheadle et B1,^ 1053)

Ihis staining schedule will be used for the detection

of oallosei where Tannic acid, ierric diloride stain the

cytoplasm brown nAiile Laomoid stains the callose and lignified

walls blue«

Sections in dist i l led Xylene III vater ^

i 1 Tannic acid Xylene II

! (5«10 ain«)

i 3 washings with xylene dist i l led water | ferric chloride^ Carbol.xylene

(1*5 Bin.) i t

3 washings with Abs, alcohol dist i l led water |

Sodlua hydregen^carbonate 90)( alcohol in 80% aleohel (2-3 ain.)

(30 ain*) t I , , 70J< alcohol (2*3 ain.)

Itaeaeld^^ t (12.84 hr«.) 80% alcohol (8-3 ala.)

L f

6:i

(•) Safranin and Fast grean^ in eonbinatloii.

I t i t a mry eoiBBOn ttain. In this staining sohadula,

f irs t tha saetions v i l l ba troatad with aquous iiaf^anln for

g-12 hours and than dahydratad. Aftar 95 | alcohol, saetions

v i l l ba treatad with Fast graan in clova oil maditm for

'd»3 minutas, ma axeess of Fast grasn v i l l be romovad

in olova o i l . i«atar, tiie saetions will ba elaanad in

xylana.

B, Xannie aeidi

1^ Xannic aeid solution v i l l ba prepared in dist i l led vatar. Xannie aeid acts as a mordant in this eonbina-tioa. ihis stains al l the primary vails brownish blaek.

9* Ferrie chlorides

3% Ferric chloride solution v i l l be prepared in dis . t i l l ed vater. I t v i l l stain the primary vails brownish blade*

10. ^odiuB hydrogen earbenatei

IS soltttiOD of the dye can be prepared by dissolving 1 gm NaHCO in 100 oe of absolute aleohol. this i s used as a aordant for Laeaoid. I t gives brown stain to the tissues labruded grey by Ferric chloride*

11* i«aaioid or (Basorcin blue)t

IjK selttticn can be prepared by dissolving 1 ga of the •tain in 300 ee of absolute alcohol* This is used as a ttiero oheaieal reagent for callose*

12* Fast greaai

0.6 to Ui solution of Fast green la absolute aleohol v i l l be obtalaed and equal voluae of clove o i l add«4* I t i t «••« f«r diffWMitiatlag Safiraala. I t t U l a t mam •r UM a l l a«i»ligBiflad tissues.

^\l

(TI ) Foster'! Tannie aeid-Fsrrle ehlorld« (Iotter, 1994}•

SoetloQ in dtistillod wator

i Tannie acid (5*10 nlnutes)

\ Washing with dis-t i l lod water properly.

Washing with dist i l led vater and study mder Bieroseope for teaporary mounts

Ferrie chloride (2-5 minutes)

For pemanent preparations ^ e sections wil l he dehydrated

in customary ethanol series and mounted in Canada balsasi. Ihe

properties of this stain are described previously. I t stains

a l l the primary vails brownish black.

l e study the morphological Yariations within tho barks,

the fibres, sclereids and sioTe-elements will be separated by

maeeratioii process following Ohouse and Ktnus (1972) and

Ohottse l i AL« (1974).

Xhe blocks of the bark containing the cambium, fixed in

F.A.A. for 3-5 days, will be sliced tangeiitially at a thickness

of about 0«6-l«0 mm. Ihese s l ices will then be treated with

6.0)( RaOK solttticm for abcrat 5 days at 4S-60*C, to softma

6V

th« tissuat, AfUr 72 hcnirs* th«s« will b« trantf«rr«d to

frttth RaOH solution of the sano coneontration* Poriodioal

choeklng will bo dono to know tho coodition of tbo troatod

• l ioot . 3ic troataont will bo eontinnod t i l l tho oollt of tho

• l ieos boeone sufficiently looso to allow tho soparation of

tho individual olomonts on a slido on applying a sl ight

pressure over the coverslip during mounting.

After reaching the desired stage, the s l ices will be

washed thoroughly on a funnel and stained in haematoxylin

for general tissue and in iS aqueous solution of either astra

blue or lactuoid for 3^-24 hours, for sieve>elem«its.

ilfter dehydration of the material in ethyl-aloc^ol

series , aounting will be done in both cases either in Si

glycerine or in Canada balsaou

6?]

Zhe present work '*3tttdi*a on the bark anatony of

different forest trees of Hadhya Pradesh around aiopal"

is an atteo^it to find out the different bark components of

the different species of forest trees. On the basis of the

above a diehotomous key will be prepared for the purpose

of identification. Ihe following aspects of th© given problem

will be studied as described belowt

I . i^oroseopio stu(^t

(a) i ^ e of bark (i^^temal appearance)

(b) Colour of bark

(o) I'ejcture of bark

IX. Hicrosoopie studyi

(a) Structure of bark before periderm formation and

radial growth.

(b) Oeneral study of bark after periderm formation

and radial growth.

(e) Detailed study of secondary phloem.

(d) Detailed study of reo^ining components of bark.

III . Results.

6JJ

Abb«, L.a. and Crafts, A . S , (1939), Phlo«m of whit* pine

and oth«r eoniferous species . Sot, 9as« 1001 e95->722«

iihiaad, ^*^ Janal, A. and Ohotise, &.K.M, (1975). Oistribution

pattern of phloem fibres in some Cassia spp. Eeo* Adv.

P i . Bel. iUlyani. p. 111.

y Khan, M.I.H. and Begum, H. (2969), Periderm in

some perennial plants, "In recent advances in the

iinatoiqjr of tropical seed plants", ed. Cbowdhury, K.A.

Hindustan Pub., New Delhi.

^ 3iddiqui, F.iU, Jaoal, ii. and Ohouse, M.K.M. (1977)

Amount of active conducting area In secondary phloen of

soae Indian tropical trees . Synip. Hec. Hes. PI. Sci . p.20.

Al f ier i , F.J. and Kvert, B.F. (1968b). Seasonal developnent

of the secondary phloea in Pinus. Am. J. Bot. 551 518«628.

Anderson, O.B. (1927). A ailerochemical study of the strueture

and developnent of flax f ibres. An. J. Bot. 141 187-211,

Anderson, B.J. and Evert, H.I. (1966). Soae aspects of phloen

development in iiJIliaill. All»« Am. J. Bot. 521 687.

Anderson, R, and Cronshaw, J. (1970). Sieve plate pores in

tobacco and bean. Planta. 91i 173-180.

Artaehvag^r* £• (1924). studies on the potato tuber. J. Agr.

Bet. 27t 809-836.

................................,^ (1950). Hie time factor in the differentiaticB

of ••eoadary xylem and phloem la pecan. Am. J. Bot. 841

63»^*

7d

Arx««, 7», Liphsehits, N. and Malsel, Y. (1968). lh« origin

and d*v«Iopnent of ph«llog«a In Hoblnia PlsMSififtftJA L*

Nov Phytol. 67I 87-93«

, W«i««I, ?• and LlphtehlU, N, (1970). Perldtrra

development and phellogen act iv i ty in shoots of iiUUBJLt

raddiana dg^vU New Phytol. 691 395*393•

bailey, I»hf« (19dl)» Coisparative anatooQr of the leaf bearing

Cactaeeae. IX. otrncture and distribution of scleren-

chyma in phloem of tSLSMiUAi fWt»*S^9Bf j f and Q\tUbintU«

Arnold Arboretum Jour. 42t 144-150.

and Kerr, 1'. (1935). aie v i s ib le structure of the

secondary va i l and i t s significance in physical and

eheraical investigations of tracheary c e l l s and f ibres .

Arnold itrboretum Jour. I6t 273-300.

., and Nast, C.CJ. (1948). Morphology and relationships

of UUfillUI* §8bliinara and ^sj l iuu. I . stem and leaf.

Aitiold Arboretum Jour. 291 77-89.

__..^ and Sva^y, a.O.L. (1949). The morphology and

relationships of Austrobaileva, Arnold Arboretum Jour.

301 211-226.

Bannan, M.W. (19S6). Xhe vascular cambium and radial growth

In JBUUA oaaidantalia L, Can. J. Sot. 33i 113-138.

Barua, P.K. and Dutta, A.C. (1959). Leaf sc lereids in the

taxonomy of JSttft aamelliaa. I I , C«if^|^ft sinensis L.

Phytomorphology 9t 372-382.

Begum, H. (I960). Anatomieal study of bark of iAAftlA nitlotiea

(I*lBa«) aad some a l l i ed speeies such as iSMiA dmmwrrmkm^

n

Willd. and j fiAftU •oIi«»i"fc Willd* Ph.D. fh«tis tubalt .

t«d to A.M.0., Aligarh,

«ilahnk«, H«l>. (1967}. Ob«r dan j^ufbander isiabalamantplattidaii

ainigar Dioseoraaoaan. L, Pflanaanphysiol. 57t 243-254*

» (1969a). Qle Si«brokran<->pIa8tldan dar Monocoty-

ladonan. Planta 841 174-134.

* ( laeab), Obar daa felnban und dia ausbraitung

dar <iiabrohr«:i-pIaSffiaflIaiaenta trad uber iiau und

Oiffaransiarungdar Siel^orex bel alnlgen Hanoootylan und

^9i IBBIUXI* ProtopXassia 681 377-402,

» (1971). Zunilainban der Slebrdtiran-Plastidan

von ^Ifft9l.9fitllft and j^ftCM (Arlstolochlaceae). Planta

97» 62.69.

iilooh, R« (1946). iJifferantlation and pattern in Monaatara

daliolo^f. Ttia Idioblaatic davalopoent of tha trlcho-

selaraida in the air roots* nm, J, 3ot« 33i 544-551,

Blyth, A. (1958). Origin of priaary extraxylary atea fibraa

in dieotgrladona. Calif. Univ., Pubis, , Bot. 301 145-232.

aorsdorf, M. (1976). Contributions to the anato^r of saeondarx

phloas and rhytidoma of cultivated poplar sorts . I-lora

(Jana) 1651 325-353.

aouek, O.B. and CronahaV| J. (1965). Iha fine struetura of

differentiating s ieve tube eleaants. J. Call. Biol. 25i

79.»6.

Bovan, V.B. (1963). Origin and develepatnt of winged eork in

BlifllTM l l limit Bot. Qas. 124t 256-261.

• Briotif 0. (1873). Ub«r alIgeiB«in«8 Vorkontn TOU Stark* in

d«i Sittbrohren. 3ot. Zaitung 31i 306-314t 322->d34|

338-344«

«_ .(1B79). tlb«r allgtiMilnea Vorkoinnan von starka In

dan diabrohr^i. Bot. ;>^alt. 31i 305.314 & 321>334 ft

33*7 «344«

3roim, H.P. (101S}« Orowth study In forast traet , I I . tjUDSOL

atrobns i.. flot, Oai!. 591 197-294,

3rovna, r.O* C1955), loraat trees of Sarawak and Brunei.

Kunching,

Jurr, r,ii» and i:.^ert» a . i , (1973), 3on© aspects of sleva

elemexit structure and development in delaginalla

Kranssiana. Frotoplasoa 78t 81-97.

«duvat, a. (1960). Observations sur las infrastructures du

cytoplasme au cours dela differentiation des ce l lu les

eribless de Cuenrbita £ t s£ L. C.a. Acad. Sei* Paris, 2St

1528* 1530.

• (1963). Inffastrueture et differeneiatlon des

eelltt les eriblees de Cuenrbita AiBfi* Svolution du

tOBoplasts e t s ignif icat ion du eontenue oel lulairefinal*

C.R. Acad. Soi . Paris. 258t 5193-.5195,

Chang, C.7, (I936)* | | ) i f ferent lat ion of protqi>hloem in the

angiospers shoot apex. Mew Phytol, 34t 21-29.

Chattaway, M,M. (1953). Ihe anatosgr of bark. I . Ihe genus

SttaMlvBti^a. Alls. J, Bot* l l 402-433,

.........................^ (1955a). Ihe anato^r of bark. T. BuaaivBtna

•peeias vith stringy bark, Aust. J. Bot. 3i xm^xm*

7.i

.(1959}« Ihe anatongr of bmrk, VII. Species of

Kttgfiia (c«nsu» l a t . ) Tropieal Moods at l * ia .

«Ch«UTMUd, 0. C1B97}. Sur l*«voIntlon d«s tub«s oribl«s

prinalres. Compt. Hmd. Acad, />cl. 1251 546-547,

• (1900). Htserches sor Lemod© d© foriaatlon dos

tubes eribles dans l a raclne de diootylfidons, itnu,

Sei , Haatt aot, l^i 33-394.

CheadlOi ?,I , and Ksau, K, (i;95d), :jecondaiy phlciea of

Calycanthaceae, Calif, Univ. Pubis,, io t , 291 397-510.

, and (1964), secondary phloem of Lirio-

dendron tullpifgr^> Diiiv, Calif. Pubis,, Jot. 36s 143-252,

^ l i f ford , £,:4,, Jr, and Ksau, I , (1963). A

staining combination for phloesi and eontinguous t i s sues .

Stain Technol, 28t 49-53,

. and Dhl, H,W, (1948), The relati<ai of metaphloea

to the types of vascular bmidles in the XcKioeotyledoneae,

iiiD, J. 3ot, 35 s 578-583,

_ _ _ _ ^ and Whitford, R,a, (1941). Observations on the

phloea in Monoeotyledoneae, I . The oeourrenee and

phylogenetie special isat ion in structure of the s ieve tubes

in the •etaphloen. An, J, 3ot, 281 ^3-627 ,

Chovdhury, K.A, (1968), History of botanical researches in

India, Buraa and Ceylon, Part X, Wood itfiatoiqr, Aligarh

NusllB Ifoiversi^, Aligarh.

Coekerhan, 0, (1930), Soae observations oneaabial act iv i ty

and seasonal s t a r ^ content in Syeanore (AftfC ttUMdft*

platanns). Leeds Phi l , Soe, Proc. 2t 64«80,

'V A 7

Crafts, A^B, (1932). PhIo«m anatony, exudation and transport

of organic nutrients in cucurbits. Plant Pb/s io l , 7t

183-225.

.,........_,_.._.,,....».... ^ (1933). Sieve-tube structure and translocation

in the potato. Plant Physiol. 8t 81-104,

(1<?34). Phloem anatomy in two species of Eifi2llaa&»

with notes on the interspecif ic graft union. 3ot. Oas,

95J 592-808,

.______„ (2339), 'Iha reXaticm between structure and

fmiction of the phloem, inr.. J, 3ot, 26i 172-177,

...,.„..__„ (1943a), Vascular differentiation in the shoot

apejE of iftfljiaia ifiiaaixiisai. ^ . J. aot, soi 110-121. „ . . . _ _ (1943b). Vascular differentiation in the shoot

apices of ten coniferous species, am, J, Sot, 30s 382-393,

and Currier, H,3, (1963). On cleve-tube function.

Protoplasma 57t 188-202,

Cronshaw, J, and Ksuu, K, (1968). P-protein in ttkn phloen

of eucnrbita. I , fhe derelopsient of P-proteln bodies.

Jour. Cell a io l . 38t 25-39.

Currier, H.B, (1957), Callose substance in plant c e l l s , An,

J. Bot, 44s 478-488,

T Ssau, K. and Cheadle, V,I, (1955). Plasnolytie

studies of phloem. Am, J, Bot, 42s 68-81,

_ _ _ _ _ _ _ and Stragger, S, (1956), ^ilin bine sad

flQoreseeaee Bieroseopy of eallose in bulb seales of

hlXlM. MUUk ^' Protoplasna 49s 522-559.

Cutter, S,a. (19i9), 'Plant anato^r*. Part I. Cells w d tissues.

Sdward Arnold Ltd. I(«iid«i.

v:>

i>«vls, J.D, and fiv«rt, R,!. (1968), i>«asoa»l d«v«lopiii«tit In

the secondary phlo«ii In EfifittiM fagBttlftlfiti* Bot. Gas.

i;29t 1>8,

_ _ . . . _ (1970}, Season&I ojrcle of phloem developtaant la

woody vines. Bot. Gax. ISlt 1E&-13&.

Do dary (1B84). Coc^arative anatomy of the vegetative organs

of the phanerogams and i e m s . Glarendcu I r e s s . Oxford,

ilerr, ttf.I* and i!.vert, K.>« (1967). &i% cambium and seasonal

development of the phloetu in iiobinia p^eudoaoagja. AO.

J. dot. 541 147-153.

Jeshpande, B.P* (3^75;. i>ifferentiation of the s ieve plate

of Cnearbita. k further view, *inn. Jot. 39$ 1)315-1022,

Duloy, K., Hercer, F.V. and Pathgeber, H. (1961). Studies in

transloeatioQ. l i . lautsbierosoopio anatony of t^e phloem,

i^ustral. Jour. diol . oc i . l i t 506-5lb.

Eames, A. J. and Mac iJaniels, L.U, (1947), tm. introdueticm to

pltfit anatoBQr, 2nd ed. p. 427.

lisehrioh, ttf. (1963). Jienidhungen ^wisehsn deiu nuftreten von

Callose und der i^elnstruktur des primaren phloeot bei

CMttTbiU f i f l i fo l ia . Planta 591 243-261.

S l l i o t t , J.H. (1935). iieasonal changes in the development of

phloem in i>yoanore, (ncer pseudoplatanus i«^;. Proe.

iiMds Phi l . Li t . aoe. Sci . Section 3i 55-67.

aiifara, C.J. (2944). Organogenesis j|i]||m£. tAdiv. Hawaii, Res.

Piibl. a i t 834*

fticl«man, S.N. (1»«8). Si«ve elem«it of limaatiens so l tan i i

XX. 0«v«lipMntftl asyeots. Aan. i o t . 89i lOS-llS.

•h]

Brvin, £•!:• ttid fivtrt, K.I, (1967), Aiptets of • ! • • • eleaMt

ontog«ny and struetur* In IJUHiM rft«ft4lfolii» ^ t . a«x.

1281 138-144.

_ _ and ^ « . « . . « _ « _ « (1970). Observations on • ! • • •

•lemMits In thv porflzinlal monoeotylsdans. Am, J. Bot.

571 21B-225.

Ssan, K. (1936). Ontogony and structure of eoll«idiyBa and

of vasottlar t issues In Celery pe t io le s . Hilgardia 101

431-476.

.. ._,... , . . .^ (1938). Ontogeny and structure of the phloea of

tobacco, dilgardia l i t 343-424.

(1939). Jevelopffi«ttt and structure of the phloeo

t i s sues , iiot. Hev. 5t 373-432.

(1943). Origin and development of primary vascular

t issues in seed plants. Bot. Hev. 9t 125-206.

(1945). Vascularization of the vegetative shoots

of BtlliMtt^ff w<i t ambneus, AS. J. Bot. 321 lB-29.

. . . ^ (1947). A study of soma sieve-tube inclusions. An.

J. Bot. 34I 224-233.

(1948). Phloem structure in the grapevine, and i t s

seasonal changes. Hilgardia 181 217-296.

___ (1950). Development and structure of the phloem

t i ssue . I I . Bot. Bev. 16t 67-114.

..^ (19S4). Primary vascular differentiation In plants.

Bot. Hev. Cambridge Phil. Soe. 29t 46-86.

... (1963). Ultrastruetare of differentiated eells ia

higher plants. Am. J. Bot. 60t 496-506.

(1964a), Aap«ets of ultras true tore of pbloan.

InI "Iha fornatloi of vood in forest troas". ad. M.H,

Zinmarmann. Aoad, Prass, Nav York. pp. 51-63.

_ _ _ (ld64b>. atruetura and davalopmant of tha bark

In dleotyladons. Ins "Iha formaticoi of wood in forast

traes". ad. H.H. ZlimnerBumn. ^cad Prass. Haw York,

pp. 37-60.

(1965a). "Plant anatomy". 2nd ad, Jcrtin «f i l^

and HODBf New York.

. , . .^ , . .__ (ic>65b). linatocgr and cytology of Yit ls phloam.

Hilgardia 371 17-72.

(1970). On the phloem of MAips^ luuiica ^* Ann.

ijot. 341 505-515.

(1972). Changes in the nucleus and tiie endo­

plasmic retioulum during differentiation of sieTe-elenent

in lUaSiA andlSA * ^ nn. dot. 36i 703-710.

end Cheadle, V.I. (1959). Size of pores and

their contents in sieve elwamits of dicotyledons, Proc.

Natl. Aoad. HeU (U.S.) 45t 166-162.

end — ^ . — . - « _ « . (1961). An aTaluation of

studies on ultrastruetura of sieve p lates . Natl. Acad.

Sc i . Proc. 47i 1716-1726.

snd (1962a). in ayaluation of

studies on ultrastruetura of toaoplast in s ieve alesients.

Rati. Aeed. Sei . Proc. 481 1-8.

.«..—.—.— a ^ ^^-'^f^62l>). Mitodiondrla ia tha ^cc N o . ••¥

phlo«i of J2l8iyem8ii lotu...Oid] ]g4t 79-86

7-) to

•nd (1965). Cytologic studies on

phloem. UhiT. Calif. Publ. 3ot. 36i 253-344.

, and aifford, Jr. E.M. (1963).

Comparative structure and possible trends of speelalisa*

tion of the phloem. M&, J. Bot. 40t 9-19.

^ _ _ - _ - ^ ana Klsley, E,^. (1962). i)evelop-

ment of s ieve-plate pores, dot, 3as. 1231 233->243.

and Cranshaw, J. (1968). aidoplasnie reticulum

in the s ieve element of Cuourbita. J. Ultrastruct. Res.

831 1-14,

Evert, H.F, ( i960) . Phloen structure in Fvrna cocmunis L, and

i t s seasonal changes. Univ. Calif. Publ. Bot. 3Zt 127-194,

. ,____ (1962). 3one aspects of phloem development in

IXXia. ASmlSSEA* UB. J, 3ot. 4Qf p. 659.

_ _ _ _ _ _ (1963a). Ontogeny and structure of the secondary

phloem in EzESIi m\m* Am. J. ^ot. 501 8-37,

_......„^ (19€3b), Hie caajblum and seasonal develc^mttit of

the phloem In fvrus i alrUf, iim. J, Bot. 501 149-159.

, Davis, J.O., Tucker, CM. and Al f ier i , I . J . (1970).

On the oeeurrwiee of nuclei in mature s ieve elemmits.

Elanta 951 281-896.

. . . . _ _ and i>eshpaade, B.P. (1969). Blectron microscope

investigation of sieve-element ontogeny and structure

in i£Lffil!: amerieana^ Protoplasna 681 403-432.

Mid (M71). Plast ids in s ieve

eloMnts and companion c e l l s of Ti l ia a—i i«» i|i| . Planta

96i 97-100.

7:1

and Eiehhorn, S.E. (1971). Lateral • ! • • • -

araa pw* In voodx dieotyladont. Ca&ad. J. Bot, 491

1509.1515.

t ««,-—.«— w d C1973). P.protain

dittribtttlon In matura slaY* alaaenta of Cnaurbita

Planta 1091 193.210.

t MurmanlB, L. and iiachs, I . J . (1966). Mother

vlev of th« ultrastrnctura of Cuenrbj-ta phloem. Jam, Bot.

SOt 563.585.

and -alfierl, F.J. (1965). Ontogeny and struetura

of ooatfarous slave c e l l * . Am. J. Bot. 52 s 1058-1066.

Jahn, rt, (1»67). "Plant jinatoay". Pergamon Press, Oxford.

and Shchorl, y. (1968). Ihe organlxatlon of the

seooadary oonduoting t issues in soine species of the

Chenopodlaeeae. Phytooorphology 17: 147-154.

*falj£, U. (1962). iieitrage zxir ultrablatologle der wurzelspltze

bel iiillliB £t&ft« iTOtopIasroa (Mien) 55; 237-254.

't'Plscher, 4. (1886). iieue beitrage zur kenntnls der slebrohren.

i^r. Verb. Kon. ;:>aehs. Jes, ^Iss. Leipzig. Nath-Phys.

38It 29X-336.

Foster, 4.S. (1934). Ihe use of tannic a d d and Iron chloride

for staining e e l l va i l s of aerlstenatle t lsst ies . Stain

Teehnol. 9t 91-92.

Frey-vyssllng, A. and Mnller, H.a, (1957). 3ubBlcroseopl«

differentiat ion of plaSKOdesaata and s ieve plates In

gWlirtflli* J. latrastruet . Bes. i i 38-48.

Oandet, J. (I960). Ontogeny of fo l iar selerelds la

a l t t i l i * An. J. Bot. 471 525-532.

80

Qhoui«, A.K.M. (1973). Zrantfutlon t i ssu* la th« ! • « • • • of

yaxM fraeata L, I a C«Ilul« 70(1) t 159.162. B«IgluB.

^__^.^_.^_^ ^14 a«»hini, ;»• (ld76)« Itie aaotmt of aet iv t t—.

of • • r t l e a l eonduetlon In the eondueting phlooa of aoM

•••rgraan and doolduous tropical troat. Bull. loxray Bot.

Club. 1031 252.254.

and (1977). Coll length variation of

phloom fibres within the bark of aoTic evergreen and

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* Oriflaal not seen.