Sal Eiol. Bwchcm Vol. 23. No. I. pp. 8S88. IWI 0038-0717 91 53.00 + 0.

00
Pnnted m Circa Bntan. All nghts mcrved CopyrIght I’ IWI Pcrgamon Pxss plc

NITROGEN RELEASE FROM THE LEAVES OF SOME

TROPICAL LEGUMES AS AFFECTED BY THEIR LIGNIN
AND POLYPHENOLIC CONTENTS

C. A. PALM and P. A. SASCHEZ

Department of Soil Science. North Carolina State University, Box 7619, Raleigh. NC 27695. U.S.A.

(Accrprrd 30 June 1990)

Summary-Leguminous plant materials used as mulches. green manures and cover crops are generally
assumed to provide a readily-available source of N to crops. However. little is known about the chemical
composition and N release patterns of the variety of legumes being used in tropical agroecosystems. N
release patterns from the leaflets of IO tropical legumes and rice straw were determined in a laboratory
expenment. Ground leaf material was allowed to dtxompose in an acid soil (pH 4.5) for 8 weeks and the
soil was analyzed periodically for extractable NH;-N and NO;-N. N release in the soil plus plant
material were compared IO that of the soil without plant material added and related IO the N. lignin and
polyphenolic concentrations of the leaflets. Three patterns of net N mineralization emerged during the
S-weeks. One pattern exhibited by the control ~011. rice straw and leaves of two of the leguminous plants
was a low. positive net mineralization. Another pattern showed much higher rates of mincrulization than
the control soil and the third pattern showed initial net immobilization followed by low but positive net
mincrahrutton rates. The amount of N mineralized during the 8 weeks as compared to the control soil
ranged from +46 IO -20% of the N added in plant matcnal. Net mincralizltion was not correlated to
% N or % lignin in the leaf material but was found to he negatively correlated to the polyphcnolic
conccntrntcon. r = -0.63. or the polyplicnolic-to-N ratio. r = -0.75. Mineralization in excess of the
control soil was found only for materials with a polyphenolic-to-N ratio ~0.5. Mechanisms to explain
the low mincralir;ttion by materials high in polyphcnolics include the formation of slahlc polymers
bctwccn polyphcnolics and amino groups. and nitrosotion. a chemical reaction of nitrite (NOI ) with
p<>lyphcnollcs. Our results show that Icguminous plant material with ;I high polyphcnollc content or
polyphcnollc-to-N ratio may not be a rc;ltlily-av;lil;thlc source of N.

IN’FKOI)UCI‘ION release patterns (Mclillo t’f ol., lY82; Melillo and

The use of leguminous plants as a source of N for
crops is particularly important in many parts of the
humid tropics where N fertilizers are not often econ-
omically feasible due to poor market and infrastruc-
ture development. In areas where use of N fertilizers
is possible, large losses of applied N due to leaching
from heavy and frequent rainstorns result in roduccd
yield and economic return. Leguminous plant ma-
tcrial used as mulches, green manures and cover crops
provides an economical source of N (Broughton.
lY77; Kang cr trl.. 1981; Wade and Sanchez. 1983).
The ctiicicncy of N use might also be improved by
sclccting and mixing plant materials that decompose
and release N in synchrony with plant demand (Vallis
and Jones, IY73; Swift, 1085).
The importance of the quality of organic additions
to N dynamics in natural and managed ecosystems
has long been recognized (Harmsen and van
Schreven, 1955; Swift c’! ul., 1979; lMclillo er (I/.. 1982:
Vitousck and Matson. 1985). There is gcncral consen-
sus that net N mineralization occurs if the N concen-
tration is above 2% and immobilization occurs below
that concentration. It has also been shown that it is
the form of N (lritani and Arnold. 1960: Berg and
Staaf. 1981; Sivapalun 6.1 (11.. 1985) and C (Tenncy
and Waksman. 1929) and the C-to-N ratio present in
the material that is important. The % lignin or
lignin-to-N ratio is often an cKcctivc index for N
Abcr, lY84).
With high N content, leguminous plant material
should release N quickly. Work by Crowther and
Mirchandani (1931). Wceraratna (lY7Y) and Ladd ct
ul. (1981) contirm this; however, Vallis and Jones
(1973) found this did not hold for Drsnrodi~nr infor-
lurlr and suggested that polyphenolic content may
affect N release patterns. In general, polyphenolics
are reactive compounds and can form stable poly-
mers with many forms of N (IMartin and Haidcr.
1980; Stevenson. 1986).
The tropics house a diversity of legumes, from
ground covers to trees. that could be used as green
manures or mulch. Little is known about the scc-
ondary compounds of tropical legumes but it is quite
possible that those growing on acid soils have high
polyphcnolic contents. N dynamics could therefore
bc atTcctcd if polyphcnolics do, in fact, alter N
rcleasc. Criteria that predict the release of N from
leguminous plant material based on their chemical
composition would facilitate the selection and man-
agement of such materials.
Our purpose was to dctcrminc the range of N,
lignin and polyphcnolic concentrations in the leaves
of several tropical legumes and to determine the
relationship bctwecn these factors and N rclcasc. A
laboratory experiment using the leaflets of tropical
legumes commonly used in agricultural production
systems was designed to address these objectives.

83
83 C. A. PALM and P. A. SANCHEZ

WATERIALS ASD METHODS

.4nal_nis of leaf material

Plant material was analyzed for 9.b N, O/Olignin and
O/bsoluble polyphenolics. Total N was determined by
microKjeldahl, lignin by the acid detergent fiber
method (Van Soest and Wine. 1968). and soluble
phenolics by a revised Folin-Denis method (King
and Heath, 1967). Polyphenolics were extracted from
the leaves in 50% aqueous methanol for I h in a water
bath (80-C) before the extract was analyzed. Total
soluble polyphenolics are expressed as % tannic acid
equivalents.

Lidxvator~ e.\-periment
Leaflets from ten leguminous trees and shrubs and
mature rice straw (Ory:n satha) were used as treat-
ments in the laboratory incubation (Table I). The
plant materials were collected from agroforestry trials
at the Yurimaguas Experiment Station in the Peru-
vian Amazon. All plants, except Leuceana leuco-
cephtrln (Lamb) de Witt. were growing on acid,
upland soils (Typic Palcudults, pH 4.5): L. kuco-
cqd~aiu was growing on a more fcrtilc alluvial soil
(Tropofluvcnt. pH 6.0). The acid, upland soils arc
rcprcscntativc of cn 30”/0 of the soils found in the
lowland humid tropics (Sanchez. 1989). All of the
spccics arc used in agroforcstry systems in the humid
tropics, four spccics are native to South or Central
America and the othsrs arc exotics (Table I). A
mixture of young and mature Icavcs wcrc hand
picked from scvcral trees of each spccics. only Icallcts
wcrc used in the cxpcrimcnt to avoid confounding N
dynamics by including lignilicd rachiscs. The fresh
Icallcts wcrc air dried and thsn ground (< I mm).
Soil used for the cxpcrimcnt was taken from the
top IOcm of the Typic Palcudult. The soil had a
loamy sand tcxturc. I% organic matter content. pH
value of 4.5, and etti.ctive cation cxchangc capacity of
3.3 cmol dm -‘. Fresh soil was sieved (<2 mm) to
remove roots and organic debris. Extractable
NH,’ ‘N and NO;. N at the beginning of the incu-
bation were 4.6 and 15.9 ilg N g-’ soil, respectively.
For each treatment, 88 mg of ground leaves were
mixed with fresh soil equivalent to 500 g oven dry
weight. to approximate a mulch rate of 3.3 I dried
plant material ha-‘. Deionized water was added
to attain a moisture content of IlOmg g-‘. field
capacity for the soil being CN I50 mg g _I. Soil without
added plant material served as a control and soil
with mature rice straw as a non-leguminous com-
parison. The soil and plant mixtures were scaled in
plastic bags and kept in the dark at room tcmpcraturc
of cu 26 C. Each trcatmcnt was replicated three
times.
Subsamples (IO g) were taken from each bag and
analyzed for cxchangcablc NH,+-N and NO;-N
immcdiatcly after addition of plant material and then
again at I. 2. 3, 4 and 8 weeks. Soil was extracted by
shaking for 30 min in 30ml of 0.5 M K$O,, cen-
trifugcd. and aliquots analyzed for NH,’ and NO<.
Ammonium was analyzed by the indophcnol blue
method (Dorich and Nelson. 1982) and nitrate by the
method of Cataldo et al. (1975). Net mineralization
was considered the diffcrcnce in exchangcablc NH,’
and NO; between two samplings.
 N release from tropical legumes 85

Albkia saman (Jacq.) F.V.M. (syn. Samanea saman),

in which net mineralization was higher than the
control during the first 2 weeks and then again at 4-8
weeks.
Total net materialization after 8-weeks differed
significantly from the 24.Opg N g-’ of the control
soil in 6 treatments (Table 1): G. sepium. Er_vrhrina sp.
and A. saman with higher concentrations and the two
Desmodium species and C. cajun with lower concen-
trations. G. sepium had the highest net mineraliz-
WEEKS ation. 54.Opg N g-’ soil. and D. o~ali/olium the
Fig. I. Examples of the three N release patterns shown by lowest, 15.9pg N g-’ soil.
the plant materials as compared to the control (0). soil with The net immobilization or mineralization during
no amendment: lnga sp. (0) showed no difference from the the first week is reflected in the 8-week cumulative net
control throughout the incubation, Desmodium ocu~ijolium mineralization. This relationship is shown by a corre-
(m) showed net immobilization during the first week fol- lation coefficient of 0.81 (P = 0.0001) between the
lowed by low but positive net mineralization. and G. sepium amount mineralized in the first week and the net
(+) showed immediate net mineralization higher than the cumulative mineralization for the 8 weeks. The corre-
control soil.
lation increases to 0.95 (P = 0.0001) when the
amount mineralized during the first 2 weeks is com-
pared to the 8-week total. The correlation does not
improve after that time. suggesting a I or 2 week
RESL’LTS
incubation may bc long enough to provide infor-
Characrerisrics of plant material mation on the relative availability of N from the
leaves.
N concentrations for the legumes ranged from a
high of 3.94% in L. leucocephala to a low of 2.37%
in Desmodium walifolium Wall, compared to I. 13% Relarion of nel minerali:ation In chemical characrer -
N in rice straw (Table I). Lignin contents ranged isrics of Iewes
from 17.9 to 52% in Inga sp. and L. leucoccphala. The amount of N mincralizcd. as compared to the
rcspcctivcly. Rice straw was at the lower end of the control. ranged from 46% of the N added in plant
range. Soluble polyphcnolics ranged from 3.61 to material, in the case for G. .scpium. to -20% for I).
1.02% for the Icgumcs compared to a low value of odi/o~ium (Table I). N mincralizcd during the first
0.69% for rice straw. There wcrc no significant week and after 8 weeks was not significantly corre-
correlations among % N. % lignin or % polyphen- lated to the N concentration of the leaves (Table 3)*
olics in the leaves. or to their lignin concentration. The highest corrc-
lation for N mineralized was with O/u polyphcnolic
Purrerns of nel N minerali:o~ion content, r = -0.59 and r = -0.63 for the first week
Net mineralization in the soil without amendment and 8-week totals, respcctivcly. The lignin-to-N ratio
was low throughout the 8 weeks but was always was only slightly correlated to the I(-week total while
positive (Pattern 1). The plant materials showed three the polyphrnolic-to-N ratio was significantly corre-
patterns of net N mineralization (Table I. Fig. I). lated for the two periods, r = -0.69 and r = -0.75.
Mineralization from lnga edulis Mart.. Ingu sp., respectively.
and rice, never ditTered significantly from the control The relationship between N mineralized and poly-
at any period during the 8-week incubation. Pattern phenolic-to-N ratio is best described by a log func-
2. exhibited by Cnssiu rericulata Willd.. Cajanus tion, MIN = -21.78*ln(PP/N) + 18.77 (r’ = 0.76).
cujan (L.) Millsp.. L. leucocephula. and the two where MlN is the cumulative 8-week mineralization
Desmorlium species, showed negative net mineraliz- and PP/N is the polyphenolic-to-N ratio (Fig. 2). Net
ation (immobilization) during the first week, followed mineralization in excess of the control was found only
by low but positive net mineralization during the for materials with a polyphenolic-to-N ratio ~0.5.
remainder of the experiment. In these species net For those materials N mineralization was linearly
mineralization generally occurred at rates equal to or related to the N content, r = 0.997; for the remaining
below that of the control. Pattern 3 is that shown by materials the correlation between net mineralization
Gliricidia sepium (Jacq.) Walp.. Eryrhrina sp., and and N content was only 0.03.

Table 2. Correlulion coefficicnls rclaring cumulilllve ilmounts of

_
(NH; + NO, )-N IO chemical char;rclcr!dlcs ot pl.mt m.twuI

% L1gnm % Phcnohc
Cumulative % %
NH; + NO, % N Lignin Phcnolics % N % N

Week I 0.31 -0.1 I -0.59 -0.3 -0.69

(0.32) (0.74) (0.06) (0.41) (0.02)
Week 6 0.42 -0.34 -0.61 -0.53 -0.75
(0.20) (0.11) (0.04) (0.09) (001)
Numbers in parentheses are the probubihty level of the correlal~on
86 C. A. PALM and P. A. SANCHEZ

for material with high ?,O N. such as L. Ieucocephalu

-0.0 01 10 1 d
%POLYPHENOLIC-to-%NITFlOGEN RATIO
Fig. 2. Relattonship between (NH; + SO, t N levels at the
R-week incubation period and the polyphenolic-to-S ratio
of the plant materul.
DISCLSSIOS
The importance of polyphenolics in decomposition
and mineralization processes has been frequently
dcbatcd (Swift ef ~1.. iY79). The issue has been
confused by the complexity and diversity of com-
pounds referred to as polyphenolics and by the
ditfcrcnt methods used to mcasurc them (Swain.
1979: IMartin and Martin. 19X!: Mnlc and Waterman,
IOX7a.b). Extraction proccdurcs are only 30-95%
ctlicicnt ;III~ the co~npou~~ds cxtrnctcd include hy-
drolyzablc tannins and condcnscd tannins, as well as
non-tannin polyphcnolics (Swain. 1979).
The interaction of polyphcnolics with N and the
fate of the N dcpcnd on the type of polyphcnolics
proscnt in the plant material. Some polyphcnolics
form complex structures by II-bonding with basic-N-
containing groups. others form stable cross linkages
with amino groups making the material resistant to
decomposition. In addition, phcnolics are readily
oxidized to quinoncs cithcr by autooxidation at ncu-
tral ptl conditions or by enzymes at a range of pH
values. The quinoncs react with other phenolics.
amino acids and amino sugars to form stable poly-
mers (Swain, 1979; Martin and Haider, 1980). The
stable polymers formed bctwcen phenolics and N-
containing compounds have characteristics similar
to. and are considered precursors to, the fulvic and
humic acids found in soil organic matter (Martin and
Haidcr. 1980; Stevenson. 1982). Another interaction
between phcnolics and N that may be especially
important in acid soils is referred to as nitrosation, a
chemical reaction of NO; with phcnolics forming
organic N compounds. Nelson and Bremncr (1969)
reported that NO; was fixed by organic matter and
Icd to gaseous losses of N in boils with neutral pH but
the formation of stable organic compounds in acid
soils. Although NO; concentrations arc generally
low in soils, NO, is formed during the process of
nitrification. and nitrosalion could account for the
apparent immobilization of inorganic N (Azhar et al..
1986a. b).
Our results indicate that polyphenolic content may
play a more important role in influencing mincraliz-
ation patterns for lcpuminous lcavcs than oio N or
lipnin-to-N ratio. This is in contrast to mineralization
patterns of non-leguminous plant material (Melillo
1’1(II.. 1982). The large amount of N immobilized and
the cxtcndcd period of net immobilization observed
and C. cajan was unexpected. but both had high
polyphenolic concentration. Millar er ul. (1936) re-
ported that net N immobilization by legumes such as
alfalfa and clovers was small and the effect persisted
for < 1 weeks. They attributed the immobilization to
an abundance of soluble organic-C relative to N.
Vallis and Jones (1973) were among the first to show
a significant and prolonged net immobilization of N
by a legume. They suggested that the polyphcnolics
20 in the leaves bound to proteins forming complexes
resistant to decomposition.
Differences in the type and amount of soluble
polyphcnolics present in the leaves could explain the
initial net release or immobilization of N seen in our
study. The patterns might also be explained by
differences in relative amounts of soluble carbo-
hydrates and soluble N in the leaves. However. the
significant correlation between net mineralization
and polyphcnolic concentration and the lack of corrc-
Iation with N and lipnin demonstrate the primary
importance of polyphcnolics in dctcrmininp the min-
cralizatlon patterns obtained. Pattern 3 cxhibitcd by
G. .x’~~u~PI.Eryrlrrirrtr sp. and rl. S(IIII(I~. in which there
was immcdiatc net mineralization of N. supgcsts the
prcscncc of both microbially-availahlc C and N
source in the Icavcs. Thcsc throc spccics also shared
low polyphcnolic contents. The only other material
used with such a low polyphcnolic content was rice
straw, but it had ;I low N content and exhibited low
net mincralir.ation.
The initial irlimohili~;Ition of N cxhihitcd by L.
lc~coce~~h~lrr. C. cujlur. C. rc.!ic.ultrttr and the two
Dcwrtotfiurtt spccics (I’attorn 2). all with rclutivcly high
N and high polyphcnolic hut low lignin conccn-
(rations, suygcsts the prcscnco of available C in the
leaves that is not mntcchcd by a sullicicnt amount of
available N. Polyphcnolics could bind to organic N
(e.g., amino acids and proteins) in the leaves making
the N unavailable, or could bind to the soluble forms
of organic N released from the Icavcs, forming rcsist-
ant complexes in the soil. In either case, thsre would
be insufficient available N from the plant material to
match the available C and net immobilization of soil
N would be expected. Sivapalan ef (11. (1985) found
lower net N mineralization from tea leaves that had
high soluble N and high polyphcnolic content in
comparison with leaves with high soluble N content
but low polyphcnolics, suggesting that the poly-
phcnolics made the soluble N unavailable. Lignin
might also bind organic N making it unavailable,
but the lack of correlation between mineralization
and lignin content and the gcncrally low lignin con-
tents of these materials make this a less probable
explanation.
The initial, rapid immobilization could also be
cxplaincd by nitrosation. the reaction discussed car-
licr. Soluble polyphcnolics that leached from the
lcavcs into the soil could bind to NOi formed during
nitrification and produce an apparent immobilization
of mineral N. This reaction readily occurs in acid soil
conditions. Nitrosation would bc implicated if NH,’
was formed at rates similar to the control but NO,
was found at dcplctcd concentrations. Thcsc rclation-
ships were not obvious for those plants with high
polyphenolic content (Table 3).
 N release from tropical legumes 87

Table 3. Cumulatwc concm:rauons’ of extractable NH;-N term. A combination of the two types of legumes may
and Ir;O, -N m the sod for the dlffcrmt treatmmts at provide the best agroecosystem management.
8 veeks
NH;-N NO;-?4 Acknonledgemenfs-We thank 4. Salazar for providing the
TEatmmt cpg N g-1 soli) diverse plant material and W. H. Schlesinger and W.
COfll~Ol 6.89 (0.48) 37.64 ( I .98) Johnson for providing facilities for polyphenolic and lignin
G. *qYprum 30 lO(1 08) 44.54 (0.97) analyses. respectively. Comments on earlier drafts by S.
&rt.rhrmo sp 16.50(1.31) 4.4 50 (2.26) Buol. C. B. Davey. G. Hoyt. R. H. Miller. M. Reynolds.
A. sal?lan 18.51 (0.48) 43.26 (0.68)
W. H. Schlesinger. D. Worsham. and two anonymous
I. rrlulu 3.97 (0.92) 38.63 (3.17)
reviewers were helpful and much appreciated. This work
0. SUIII’U 4.8J(O.40) 39.10 (0.06)
Inru sp 3. I6 (0.79) 39.22 (0.55) was part of the North Carolina State University TROP-
L. luururephalu 5 58 (0.54) 41.21 (1 ‘l9) SOILS Program. supported by US Agency for International
c. rrriluluru 3.70 (0.50) 41 OO(l 97) Development (USAID).
ca. w/on I 89(0.:4) 37.17(1.74)
Drsmdum ~~~~rclrr 088(0.31) 37.11(1.1?)
D. o~hf~drum ORO(O.lll 36 2: (0.46)
REFERESCES
‘Mean (SD) of three rC,?h‘XleS.
Azhar El Sayed. Van Cleemput 0. and Verstruete W.
(1986a) Nitrification mediated nitrogen immobilization in
soils. Plonr and Soil 94, 4Ol-tOV.
Pattern I in which little net mineralization or Azhar El Sayed. Verhe R.. Proot M.. Sandra P. and
immobilization occurred. as for both Itrgu species, Verstraete W. (1986b) Bindlng of nitrite-N on polyphe-
could reflect their higher lignin content. and therefore nols during nitrification. Plant rrntl Soil 94, 369-382.
supposedly lower soluble-C. of these materials. The Berg B. and Staff H. (1981) Leaching. accumulation and
two In,qcl species had 50% higher lignin contents than release of nitrogen in decomposing forest htter. In Tu~I~,~-
the other spccics. Lignin content may influcncc the rricrl Nitroxc*n C~ck0.r (F. E. Clark and T. Rosswall. Eds).
pp. 163-178. Ecological Bulletin 33 (Stockholm).
long-term N rclcasc as is indicated by a weak corre-
Broughton W. J. (1977) Erect of various covers on soil
lation found for the S-week cumulative mincraliz-
fertility under flerxvr hrcuiliens~s and on growth of the
ation and the lignin-to-N ratio. tree. /(pro-Eco.ql.vlrn~s 3. I47 170.
The patterns of N mineralization obtained in this Cafaldo D. A.. Ilaroon M.. Schr;lder L. E. and Young V. L.
study arc complex and involve ;I variety of factors. (1975) Rapid colorlmctric dctcmunations of nitrate I”
The lignin-10-N ratio. which has hccn shown to plant tissue by titration of salicyclic ;~cld. <.~~,,r,frrtrri~tr/rr,,rs
control dccornposition and mincrali~ation in non- LI .%lil .scrcncX, urtd I’llrnl .-lncrl,~.vr.~
6, 7 I no.
Icguminous kxvcs (Mclillo (‘I trl.. 1982). was not ;I Dorlch R. A. and Nelson D. W. (IOX?) I)lrect coIorImctric
controlling factor in lhcsc leguminous Icallcts. at least measurement of ammonium m potassium chloride CX-
tracts in soil. SOI/ .%ic*trcv*Sf~.rc,ry I)/’ ~ln~c~r~~rJf~~rrtcrl 47.
in the short term. The lignin-to-N ratios of the
x33 X36.
Icguminous IC;IVUS used in this study, howcvcr. were
Ilarmscn G. W. and v;m Schrcvcn 0. A. (1955)Mmcr;lll/-
below that of the plant nxttorials used in the study by ation of organic nilrogcn in soil. A&mc~c*s Cr ,Igrr~n0nr~’
Mclillo C’Itrl. (I%?). Our study suggests that polyphe- 7, 299 -39x.
nolics content could control the short-term N rele;lsc Iritani W. M. and Arnold C. Y. (1960) Nitrogen release 01
and availability from legumes. Leguminous plant vegetable crop residues during incubation as rel;lred IO
materials have previously been considered as readily their chemical composition. Soil Science 89. 74 82.
available sources of N. This assumption may be Kang B. T.. Wilson G. F. and Sipkrns L. (19X1) Allcycrop-
incorrsct if ths plant material has a high poiyphenolic ping maize (Zrcr ntcr):r L.) and leucaens (Lrucorno Ieucrr-
cephulu Lam.) in southern Nigeria. Pl~rrtr und Soi/ 63.
content, or ;I polyphcnolic-to-N ratio >O.S. This type
165-179.
ofshort-term incubation could be useful for screening
King J. G. C. and tlrath G. W. (1967) The chemical analyhis
Irgumcs. Another cxperimcntal study using plant of small samples of leaf material and the rclationshlp
materials, including non-legumes, with a broader between the disappearance and composition of leaves.
range in N, lignin and polyphenolic content would Pedohiolcq+ 7. 191-197.
provide addltionul information on 1hc overall import- Ladd J. N.. Oades J. M. and Amato M. (1981) Dlstrihution
ance of polyphenolics and their interaction with and recovery of nitrogen from legume residues decompos-
lignins in controlling N release from plant materials. ing in soils sown to wheat in the tield. Soil Biohqy &
Also. it must bc pointed out that only leaflets were Biochenri.srry 13. 251-256.
Martin J. P. and Haider K. (1980) Microbial degradation
used in this study, N release patterns from the entire
and stabilization of “C-labeled lignins. phenols. and
lcavcs may be different and should be investigated.
phenolic polymers in relation to soil humus formation. In
The diverse N release patterns observed from the L&yin Biode~rudution: Microbiology. Chenkfry ud P o -
Ic;lRets of IO tropical legumes and the possible role of rmriul Applicurions. Vol. 2 (T. K. Kirk, T. Higuchi and
polyphcnolics on N availability have several manage- H. M. Chang. Eds). pp. 78-100. CRC Press. Wesr Palm
mcnt implications. Lcgumcs with low polyphcnolic Beach.
content used as green manure or mulch may release Martin J. S. and Martin M. M. (19X2) Tannin assays in
N rapidly and provide sutficient N for plant growth, ecological studies: lack of correlation between phenolics.
but the N may be readily lost by lcaching if not proanthocyanidins and protein- precipitating constituents
in mature foliage of six oak species. Oecofogio 51,
relenscd in synchrony with plant demands. Legumes
2052 I I.
with high polyphenolic content may rcac1 with or-
Melillo J. M. and Aber J. D. (1983) Nutrient immobilization
ganic N compounds do not provide enough N for in decaying litter: an example of carbon-nutrtent inter-
plant growth in the short term, but they may contrib- actions. In Trendy in Ecoloxicul Research fkr rhe 1980’s
utc to the build-up of organic N in the soil and (J. H. Cooley and F. 8. Golley. Eds). pp. 193-215.
provide a low but continual supply of N in the long Plenum. New York.
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