A Comparison of Lipid Components of the Fresh and Dead Leaves and Pneumatophores of the Mangrove Avicennia Marina

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Phytochemistry,Vo l. 20, No . 4, pp . 659 -666 , 1981. 0031-9422/81/040659--08 $02.00/0Printed in Grea t Britain. © 1981 Pergamon Press Ltd.

A C O M P A R I S O N O F L I P I D C O M P O N E N T S O F T H E F R E SH

L E A V E S A N D P N E U M A T O P H O R E S O F TH E

M A N G R O V E A V I C E N N I A M A R I N A

A N D D E A D

G . P . WANNIGAMA,* J . K. VOLKM AN,~ F . T . GILLAN, P . D . NICHOLS an d R. B . JOHNS~

De partm ent o f Organic C hemistry, University of Melbourne, Parkville, Victoria, 3052, Au stralia

(Revised received 18 August 1980)

Key Word Iudex--Avicennia marina; Avicenniaceae; mangrove; hydrocarbons; alcohols; sterols; tr i terpenealcoho ls; monobasic ad ds ; ~t¢o-dibasic cids; ¢o-hydroxy acids.

A b s t r a c t - - T h e c o m p o n e n t h y d r o c a r b o n s , s t er o l s, a l c o h o ls , m o n o b a s i c , ~ , c~ -d ib a si c a n d c o - hy d r o x y a c i d s o f t h e f r e sh a n d

d e c a y e d le a v es a n d t h e p n e u m a t o p h o r e s o f t h e m a n g r o v e Av ic e nn ia mar ina a r e r e p o r t e d i n d e t a il . F r o m t h e q u a n t i t a t i v ec o m p a r i s o n s w h i c h c a n b e d r a w n , r e l a t iv e c h a n g e s i n t h e l i p i d c l a s se s o c c u r r i n g d u r i n g l e a f d e c a y c a n b e h i g h l i g h t e d .T h e s e b a s e - l in e d a t a a r e i m p o r t a n t t o o u r u n d e r s t a n d i n g o f i n p u t s t o m a r i n e i n t e r ti d a l s e d im e n t s . D u r i n g l e a f d e c a y t h eo n l y s i g n i fi c a nt c h a n g e s w e r e a r e d u c t i o n i n t h e t o t a l a b s o l u t e c o n c e n t r a t i o n s o f m o n o b a s i c a c i d s d u e l a r g e l y t o a d e c r e a s ei n c o n c e n t r a t i o n o f t h e C 1 8 p o l y u n s a t u r a t e d f a t ty a c i ds , a n d a n e n h a n c e m e n t o f t h e c o n c e n t r a t i o n s o f t h e l o n g - c h a i nm o n o b a s i c a c i d s , c o - h y d r o x y a c i d s a n d ~ , co - d ib a s ic a c id s . T h i s r e s i s ta n c e t o d e g r a d a t i o n s h o w n b y t h e c u t i n d e r i v e d a c i d s( or,c o-d ib as ic , c o - h y d r o x y a n d l o n g - c h a i n m o n o b a s i c a c i d s ) r e la t i v e to t h e c e l l u l a r a n d w a x d e r i v e d l i p i d s m a y a l l o w t h e s ec u t i n c o m p o n e n t s t o b e u s e d a s q u a n t i t a ti v e m a r k e r s o f A. marina i n m a n g r o v e a s s o c i a te d s e d i m e n t s.

INTRODUCTION

I n t h e c o u r s e o f t h e s t u d i e s o f l i p i d i n p u t a n d d i a g e n e s i s i n

m a n g r o v e a s s o c i a t e d s e d i m e n t s w e h a v e p r e v i o u s l yr e p o r t e d a n a n a l y s i s o f t h e d i a c i d s o f t h e m a n g r o v eAv ic e nn ia mar ina [ 1 ] a n d p r o p o s e d d i r e c t in p u t o f th e

d i a c i d s t o t h e s e d i m e n t s . A s t u d y b y S a s s e n o n m a n g r o v ep e a t s [2 ,3 ] , i n v o l v i n g a n a l y s i s o f m o n o b a s i c a c i d s o n l y ,s u g g e s te d a m a j o r i n p u t o f m a n g r o v e l i p i d s t o a s e d i m e n t,a n d p r e s e n t e d e v i d e n c e ( l e af l i t t e r a n a l y s i s ) o f s e le c t iv ed e g r a d a t i o n o f s h o r t- c h a i n f at t y a d d s . T h e m a n g r o v e sRhizophora mang le , L agunc u lar ia rac e mosa , Av ic e nn ia

ge rminans a n d A. n i t i da w e r e a n a l y s e d . S i m i l a rd e g r a d a t i v e s tu d i e s h a v e b e e n r e p o r t e d o n t h e s e a g r as sSpar t ina a l t e rn i f l o ra [ 4 ] , a n d o n l e a v e s i n f r e sh w a t e rs t r e a m s [ 5 , 6 ] .

I n t h i s p a p e r a m o r e d e t a i l e d l i p i d a n a l y s i s o f t h e l i v i n ga n d d e a d l e a v e s a n d t h e p n e u m a t o p b o r e s o f Av ic e nn ia

mar ina c o l le c t ed f r o m P o r t F r a n k l i n , V i c t o r i a i s r e p o r t e d .L i p i d c l a s s e s a n a l y s e d i n c l u d e h y d r o c a r b o n s , s t e r o l s , f a t t ya l c o h o l s , m o n o b a s i c a d d s , ~ ¢ o - d i b a s i c a c i d s a n d ~ o-h y d r o x y a d d s . T h e f a t t y a c id s a n d a l c o h o l s w e re f u r th e rs u b d i v i d e d i n t o b o u n d a n d s o l v e n t - e x tr a c t a b le f r ac t io n s ,a s p r i o r w o r k [ 7 ] h a s s u g g e s t e d t h e i n t e r p r e t a t i v eu s e fu l n es s o f t h is p r o c e d u r e . T h e m o n o b a s i c a c i d d a t ar e p o r t e d i n c l u d e s t h e m o l e c u l a r s t r u c t u r e s o f t h ec o m p o n e n t f a tt y a d d s , s i n ce s u c h d a t a a r e e s s en t i al fo rb o t h b i o c h e m i c a l a n d g e o c h e m i c a l i n t e r p r e t a t i o n o f t h ec o m p o n e n t s p r e s e n t a n d o f t h e c h a n g e s o b s e rv e d d u r i n g

* On leave from the D epartm ent of Chemistry, University ofPeradeniya, Peradeniya, Sri Lanka.

t Present address: Org anic Geochemistry Un it , School ofChemistry, Bristol University, BS8 ITS, U.K.

To w hom reprint requests should be addressed.

m a n g r o v e b i o d e g r a d a t i o n a n d e a r l y d ia g e n e s is in r e c e n tm a r i n e s e d i m e n t s [ 7 ] . S u c h k n o w l e d g e i s p e r t i n e n t t om o d e l s o f l i g n i t e c o a l d e p o s i t i o n .

RESULTS AND DISCUSSION

T h e c o m p o n e n t h y d r o c a r b o n s o f t h e fr e sh a n d d ead

l e av e s a n d t h e p n e u m a t o p h o r e s o f Av ic e nn ia mar ina are

p r e s e n t e d in T a b l e 1. T h e h y d r o c a r b o n d i s t r i b u t i o n s i nt h e l e a f w a x a n d i n t h e l e a v e s af t e r w a x r e m o v a l a r e a l s op r e s e n t e d . T h e n - a l k a n e d i s t r i b u t i o n o f t h e f r es h l ea v e s iss i m i l a r t o t h o s e r e p o r t e d f o r t h e w a x e s o f m o s t h i g h e rp l a n t s [ 8 - 1 0 ] . F e a t u r e s s u c h a s th e a b u n d a n c e o f

b r a n c h e d a l k a n e s a n d t h e p r e s e n ce o f lo n g - c h a i n a l k e n e sd o h o w e v e r m a k e t h e h y d r o c a r b o n p r o f il e o f A . m a r i n ad i s t in c t i v e . T h e p a u c i t y o f p r e v i o u s a l k a n e r e p o r t s w h i c h

i n c lu d e d a t a o n t h e s e c o m p o n e n t s m a y b e t h e r e s u l t o f t h ei n su f fi ci e nt r e so l u t i o n o f t h e p a c k e d G C c o l u m n sp r e v i o u s l y e m p l o y e d f o r s u c h a n a l y s e s . H o w e v e r , o n ee x a m p l e i s t h e n o t i n g o f b r a n c h e d a l k a n e s i n t h ee p i c u t i c u l a r w a x o f Andropogon haUi i [ 1 1 ] . T h eh y d r o c a r b o n d i s t r i b u t i o n i n t h e d e a d l e av e s i s s i m i l a r t ot h a t f o u n d i n t h e f r es h l e a v es e v e n t o t h e p r e s e n c e o fb r a n c h e d a l k a n e s a n d t r a ce s o f l o n g - c h a in l e n g t h a l k en e s .T h e l e a f w a x c o n t r a s t s w i t h t h e w h o l e l e a f a n a l y s i s o n l y i nt h e a b s e n c e o f s q u a l e n e .

T h e c o m p o n e n t h y d r o c a r b o n s i n th e p n e u m a t o p h o r e sa r e c h a r ac t e r i z ed b y a p r e d o m i n a n c e o f s h o r t e r c h a i n n -a l k a n es . L o n g - c h a i n n - a lk a n e s , n -a l k e ne s a n d b r a n c h e da l k a n e s w e r e n o t d e t e c t e d i n c o n t r a s t t o t h e l e a v e s .S q u a l e n e w a s p r e s e n t , b u t a t a r e d u c e d r e l a t i v e l e v elc o m p a r e d t o t h e l e a f s a m p l e s . T h e s e d i f f er e n ce s c a n b er a t i o n a l i z e d i n t e r m s o f t h e d i f f e r e n t s t r u c t u r e a n df u n c t i o n o f t h e s e t w o p l a n t o r g a n s .

T h e s t e ro l c o m p o s i t i o n o f t h e f re s h a n d d e a d l e a ve s o fA. mar ina ( T a b l e 2 ) a r e e x t r e m e l y s i m i l a r . S i t o s t e r o l ( t h e

659





Lipids of mangrove leaves 661

Table 3. Fatty alcohol composition of Anicennia marina

Percentage composition of total alcohols

Fresh leaves Dead leaves

Homologue TSE

12:o 0.25

14:o -

15:o 0.17

16:0 0.13

18:0 1.88

18:l -

20:o 0.88

21:o tr

22:o 0.84

23:0 0.45

24:0 4.45

25:0 0.33

26:0 7.06

27~0 1.84

28:0 8.50

29:0 0.31

30:o 5.08

Phytol 55.84

Others 8.90

Wax

0.66

tr

2.0

tr

18.7

2.4

37.6

1.7

21.2

1.3

14.4

(TSE - wax)

0.7

tr

1.3

tr

1.1

1.4

17.5

0.71

49.5

0.51

20.9

nad

tr

Bound

0.25

tr

tr

0.13

0.67

tr

3.41

0.42

5.45

0.70

11.81

1.61

11.11

1.52

25.14

0.10

20.96

12.97

3.76

TSE

0.32

0.60

tr

0.27

0.40

1.80

tr

1.77

0.16

6.28

0.99

9.10

0.75

12.75

0.20

3.00

33.76

27.67

Bound

tr

0.30

0.58

0.13

3.96

1.40

4.66

1.25

8.89

0.69

6.53

1.21

9.53

tr

2.93

33.46

24.29

Pneumatophores

7.9

7.3

12.7

17.3

30.8

5.9

9.2

2.6

6.3

-

tr

Total absolute

concn (kg/g

dry wt.) 2270 150 1500 100

tr = Trace, ~0.1%.nad = Not accurately determined.

TSE = Total solvent extractables.

differs from the leaves where the ratio of sitosterol to

stigmasterol is 3 to 1.

The distribution of alcohols in the leaves of A. marina

(Table 3) is typical of most higher plants [8], ranging

from 18 :0 to 30:0 and maximiz ing a t 28:0 . Phyto l was

detected in significant quantity in both the fresh and dead

leaf samples, but was not detected in the wax fraction of

the fresh leaves. The alcohol fraction of the solvent extract

of the fresh leaves contained a significantly higher relative

and absolute amou nt of phytol than was found in the deadleaves indicating that rapid degradation of the phytol and

the chlorin pigment occurs du ring leaf decay (equivalent

chlorophyll-a ratio of fresh leaves/dead leaves is 35). The

concentration of bound alcohols in the fresh leaves was

appreciably lower than the level of solvent extractable

alcohols (Table 4 ), and was not observed to increase on

leaf decay. The distribution of alcohols within the

pneum atophore sample was unusu al, with li tt le even over

odd predominance occurring. Phytol w as a minor

com ponent only of this fraction after sapon ification

reflecting the low concen tration of chlorop hyll in the

pneumatophores.

Six compou nds identified by characteristic massspectral fragmentation patterns and GC retention data as

triterpene alcohols were found in both the fresh and dead

* Double bond positions (w) are numbered from the methyl

end of the fatty acid, all subsequent double bonds are methylene

interrupted.

leaves (Table 5) and in the pneumatopho res of A. marina

(these compoun ds were major components in the wax

fraction of the leaves). The triterpene alcohols identified

were: urs-12-ene-38-01, olean-12-ene-3p-01, lup-

20(29)ene-3 /3-ol and friedoolean-38-01. The major

component (RR, 1.168 ) has not yet been fully identified.

The absolute amou nts of this class of compou nd in both

the fresh and dead leaf samples were similar, apart from a

large decrease in the concentration of friedoolean-3B-01.

This constancy in absolute amou nts of the triterpenealcohols between the two samples was also observed for

the hydrocarbons, sterols and fatty alcohols indicating

that these classes of compoun ds are not degraded rapidly

during leaf decay. T his may reflect their l imited value as

microbial nutrients.

The monobasic acid composition of the fresh leaves of

A . marina (Table 6) ranges from C,, to C,, with 16:0 ,

1 8 : lw9*, 18 :2 ~6 a n d 1 8 :3~3 present as the predominant

acids. These data are similar to those reported previously

fo r A . nitada an d A. germinans [2,3] but include the

molecular structures of the component mono carboxylic

acids unlike the previous work. The significance of

knowing the isomer positions in the monoenoic acids hasbeen documented previously [14,15 ], and the monoenoic

acids detected in the fresh leaves were: 1.5:1, 16:lw7 ,

1 6 : 1 ~9 , tram 16: 1~13, 17:108, 18: lw7, 18:109, 19: l

and 20: 1~9.

The bound monob asic acids (released after saponi-

fication of the solvent extracted leaf residue) of the fresh



66 2 G. P. WANNIGAMA et trl.

Table 4. Absolute amounts of lipid classes of Aricennia mu r in u

Concentration klg/g (dry wt.)

Lipid class Fresh leaves Dead leaves Ratio dead,‘fresh

Hydrocarbons 3070 3200 1.04

Sterols 1900 1640 0.86

Alcohols TSE 2270 1500 0.66

TSE less phytol 1000 990 0.99

Phytol 1270 510 0.40

Bound alcohols 150 100 0.06

Triterpene alcohols 1090 1130 1.04

Monobasic acids TSE 18420 7730 0.42

Saturated short-chain 5190 4040 0.78

Unsaturated 12660 3370 0.27

Long-chain 570 320 0.56

Bound 930 2520 2.71



Long-chain I60 610 3.81

Total 19350 10250 0.53



Long-chain 730 930 1.27

r,cu-Dibasic acids 1600 2870 1.79

c+Hydroxy acids 1190 1550 1.30

TSE = total solvent extractable.

leaves contained lower relative am ounts of 18 :2 ~6 a n d

18:3w 3. This decrease is largely compensated for by a

corresponding increase in the relative am ount of high

M W mon obasic acids (22:0 to 30:O ). The ratios ofsolvent

extractable and bound long chain acids in dead to fresh

leaves (Table 4) suggest some interconversion from solvent

extractable to bound is occurring for this l ipid class. A

similar change is observed for the short chain (C < 22)

fatty acids. T he absence of long-chain acids in the solvent

extracted fraction of the fresh leaves after the wax has been

removed demonstrates their non-involvement in the

membrane lipids.

The major difference between the monob asic acid

composition of the fresh and dead leaves is the marked

decrease in the relative and absolute amo unt of 18:2w 6

and 18:3w 3. The rela tive abundance of 14:O and 16:0

increased dramatically. During leaf decay under oxic

conditions all unsaturated acids present would be

expected to be degrad ed after cellular lysis: the rate being

dependent on the degree of unsaturation.

The presence of a numb er of iso and ant&so branched

acids, 17:lwlO and ~6, cyclopropyl 19:0 along with

higher relative amou nts of 14:0, 16 :0, and 18: lo>7 in the

dead leaves sam ple is indicative of a significant

contribution to the component fatty acids by sedimentary

microorganisms involved in leaf degradation. The

bacterial origin of these acids has been reportedpreviously [ 15 1, which suggests that significant bacterial

colonization of the dead A. mu r i ru z leaves has occurred.

The longer chain monoenoic acids 22: 1 and 24: 1 not

found in the fresh leaf sample and believed to be mainly

the w9-isomers were detected in the dead leaves of A.

ma r in a . The occurrence of these acids in sediments

(thought to be of yeast origin) and M ~~rohn ctrri~~ has been

reported previously [ 16,1 71 indicating that an exogenous

origin for these acids in the dead leaves of A. rn a r in u is

probable.

The absolute amou nts of the long-chain saturated fatty

acids in the dead leaves is increased when com pared with

the fresh leaves (T able 4 ). These acids are probably less

viable microbiological substrates than the short chain

saturated and unsaturated fatty acids as noted previously

[2,3]. The presence of a num ber of C,, and Cl2

polyunsaturated fatty acids not detected in the fresh

leaves, but present in the dead leaves sample indicates that

an alternative source for these acids is l ikely. Marine

dia toms a re r ich in 20:4~16 and 20:5w 3 [18] , however ,

colonization by other microscopic algae e.g., certain

Chrysoph yta [19] could also account for these acids. The

fatty acid distribution of the pneumatopho res is similar to

the fresh mangrove leaves although the relative

concentration of 18:30.1 3 n particular is reduced. Sm all

quantities of 22: 1 and 24: 1 were also detected.

The r,w-dibasic acid composition of A. nzariru isolatedby sapon ification of the leaf residues after solvent

extraction (Table 7) shows di 16:0, di 18:0 and di 18: 1 as

the major components with di 18:2 present at a lower

relative level. The absolute amo unt of the r,w-dibasic

acids in the fresh leaves is 8”, of that of the monobasic

fatty acids. The dead leaves sample showed a similar

diacid distribution to that of the fresh leaves of A. rrrcviuuwith some changes in the minor co nstituents observed.

The relative amo unt of di IX::! was diminished and the

higher M W components increased. Th e absolute am ounts

of the diacids in the dead leaves were 1.8 times that

observed in the fresh leaves and 28 I’ ,>of that of the

monob asic acids in the dead leaves. It was also noted that

the absolute amo unt of long-chain monoacids in the dead



Lipids of mangrove leaves

Table 5 . Triterpene alcohol concentration of the fresh and dead leaves of Auicennia marina

6 6 3

Co mp o u n d

Percentage composition MS fragmentation m/e (rel. int.)RR,* Fresh leaves Dead leaves M ’ (of TMSi ether)

Unidentified A

Lup-20(29)ene-

38-01 (lupeol)

Olean-12-en-3P-

01 (/I-amyrin)

Urs-12-en-3B-01

(a-amyrin)

Unidentified B

Friedoolean-

3/?-01 friede-

linol)

0 .9 0 4

0 .9 3 7

1 .0 0 0

1 .1 1 0

1 .1 6 8

1 .3 9 1

1.6 2 .7

3 .6 4 .9

11.9 12 .1 498(1 .3) 218(100) , 203(35) , 190(19) , 189(17) ,

204(12) , 109 (7) , 121(7) , 135(7) , 147(7) ,175(7) , 161(4) , 257(3) , 279 (3) , 483(l ) .

17 .6 20 .5 4 9 8 (3 )

57.6 57 .1 4 9 8 (1 9 )

218(100) , 18 9(22) , 190(16) , 203(16) ,

220(16) , 135 (12) , 121(10) , lC9(8) ,

147(7) , 161(6) , 175 (5) , 279(5) , 257(l ) ,

408(l ) .

189(100) , 109(73) , 190(64) , 135(58) ,203(51) , 121 (50) , 218(32) , 147(29) ,

175(28) , 156(27) , 369(21) , 231(16) ,

279(15) , 393(10) , 257(9) , 407(8) ,

299(7) , 483(6) , 245(5) , 325(4) .

N o m a s s spectral data.

498(17) 189(100) , 205(90) , 190(86) , 218(80) ,203(75) , 109(74 ) , 147(26) , 119(22) ,

161(20) , 174(20) , 279(20) , 229(18) ,

293(8) , 257(6) .

7 .7 2 .7 SO O (2 .5 )69(100) , 109 (70) , 123(68) , 125(62) ,

205(42) , 273(37) , 163(33) , 137(32) ,

159(30) , 190(26) , 179(25) , 149(23) ,

246(21) , 231(18) , 302(16) , 217(15) ,

426(15) , 341(11) , 287(6) , 259(5) ,

41 l (5) .

Total absoluteconcn (fig/gdry wt.) 1 0 9 0

* RR, on SE 30, p-Amyrin = 1 .0 .

1 1 3 0

l e aves was increased by a factor of 1.3 compared with the

fresh leaves (T able 4), this factor being the same as that

observed for the total o-hydroxy acids. The high absolute

concentrations of a,w-dibasic acids in A. marina is unusu alsince these acids are generally considered to be mino r

constituents of leaf cutin [20]. The pneumatophores

contained the same diacid components as the A. marina

leaves although di 18: 2 was not detected. The dibasic acid

concentration in the pneumatophores was calculated,

however, to be 1.7 times the monobasic acid

concen tration, significantly different from that found in

the leaves.

The w-hydroxy acid profile of the fresh and dead leaves

of A. marina (Table 8) shows both identical chain-length

distribution and similar relative concen trations of

individual components when compared with the a,w-di-

basic fatty acids of A. marina (Table 7) . These data suggestthe direc t biosynthesis of the a ,w-dibasic ac ids from the o-hydroxy acids in the mang rove A. marina. This hypothesis

has been repor ted previously [21] , as has the oxidativeconversion of monobasic ac ids to o-hydroxy acids [22] .

The tota l absolute amounts of monob asic ac ids in thedead leaves o f A . marina are 53 % of the absolute amounts

found in the f resh leaves, wh ereas the absolute amounts ofboth the so-dibasic and w-hydroxy acids have increased(Table 4) . This observed increase in the absolute level of

diac ids in the dead leaves indicates preferentia ldegradation of the ce llular contents and resultant

enrichm ent of the cutin (obtained by saponification of theleaf residues a f ter solvent extrac tion) and w ax der ivedcomponents .

The chain length distr ibution of the a ,o-dibasic ac idsand w-hydroxy acids in the pneum atophores of A. marina

(Tables 7 and 8) are very similar , as expected. The

presence of o-hydroxy 12:0 and 14:0 in the

pneum atophores but not in the leaf suggests somedifferences in the chain- length specif ic i t ies of the enzymesinvolved. These shor ter chain-length o-hydroxy acids arenot fur ther oxidized to cc ,w-dibasic ac ids. We suggest tha t

the o-hydroxy acid dehydroge nase is specif ic for the C,,and C, , chain lengths in the pneum atophores of A.

marina.

The data p resente d on the lipid distribution in thedecayed leaves re la t ive to the f resh leaves of A. marina

(Table 4 ) show the only signif icant changes to be areduction in the tota l amount of monoba sic ac ids present







666 G . P. W A N N I G A M A t u l .

SIL 47C NP (47 “/ ‘, cyanopropyl silicone) 43m x 0.2mm i.d.

column [25]. He was used as carrier gas (linear flow: 20cm/sec).

The SE30 column was programmed from 160” to 280” at 2.5” per

mm and the SIL47CNP column from loo” to 240” at 3” per min.

FID detectors were used with injector and manifold temperatures

at 280” and 300” respectively. Monobasic Me esters were

tentatively identified by co-chromatography with authentic

standards and by RR, [26,27] and ECL measurements [28,29].

All lipid components were quantitated by calibrated GC response

and are subject to errors of + 10 T0

Sterol identifications were based on RR, co-injection with

authentic standards and 0~0, oxidation [2.5]. The latter

technique allowed differentiation of the degree of unsaturation of

individual sterols. This is especially important in the separation

of 28isofucosterol and Sa-stigmastanol which had the same R,

on the GC columns employed. The major sterols were confirmed

by GCMS.

Acknowl&ements-The authors thank the ARGC for financial

support. J.K.V. and F.T.G. acknowledge C.P.G. Awards andP.D.N. a University of Melbourne Postgraduate Award.

Professor G. Eglinton FRS, is thanked for access to the NERC

financed C-GC-MS (GR3/2951 and GR3/3419).

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Documents

A Comparison of Lipid Components of the Fresh and Dead Leaves and Pneumatophores of the Mangrove Avicennia Marina