Animal Feed Science and Technology, 26 (1989) 1-28 1

Elsevier Science Publishers B.V., Amsterdam - - Printed in The Netherlands

Leucaena leucocephala i n P o u l t r y N u t r i t i o n -
A Review

J.P.F. D'MELLO and T. ACAMOVIC*

Department of Agricultural Biochemistry, The Edinburgh School of Agriculture, West Mains
Road, Edinburgh EH9 3JG (Gt. Britain)
(Received 27 June 1988; accepted for publication 29 November 1988)

ABSTRACT

D'Mello, J.P.F. and Acamovic, T., 1989. Leucaena leucocephala in poultry nutrition - - a review.
Anita. Feed Sci. Technol., 26: 1-28.

The nutritional attributes of Leucaena leucocephala and the factors limiting its use in poultry
diets have been the subject of considerable research over the past 10 years. Recent data confirm
the previous findings concerning the favourable proximate composition of Leucaena leaf meal
( LLM ), although the carotenoid content of this legume is now emerging as its major asset for the
pigmentation of egg yolks and broiler carcasses.
Toxicological evaluation continues to be a major feature of research on Leucaena. Following
recent innovative developments in analytical chemistry, there is now more reliable in/brmation
pertaining to the concentration of mimosine in LLM and seeds and to the degradation of this
amino acid to 3-hydroxy-4 (1H)-pyridone. Data on the concentrations of tannins, trypsin inhib-
itors, galactomannan gums, saponins and flavonols are also reviewed in this paper.
Detoxification of Leucaena for poultry remains an outstanding problem, since there are limi-
tations to conventional approaches such as plant breeding and the use of heat treatments. Con-
siderable amelioration of the adverse effects of LLM may be obtained by the use of dietary addi-
tives such as ferric sulphate and polyethylene glycol. However, these improvements rely upon
appropriate adjustments for the low apparent metabolisable energy (AME) value of LLM. Thus
despite recent advances, the low AME of LLM remains a challenging issue for the future.

INTRODUCTION

Leucaena leucocephala (Lam.) de Wit is a tropical legume, indigenous to

Mexico, but now widely distributed in the high-rainfall regions of Central
America, Africa, Asia and northern Australia (National Academy of Sciences,
1977 ). In economic terms, it is the most prominent member of a genus of shrubs
and trees comprising 10 recognised species. The two important types of this

*Present address: Chemistry Department, School of Agriculture, 581 King Street, Aberdeen AB9
1UD, Gt. Britain.

0377-8401/89/$03.50 © 1989 Elsevier Science Publishers B.V.

Fig. 1. Leucaena leucocephala of the Peru cultivarthriving in the rockyterrain of the Yucatan
province of Mexico.

legume are the giant cultivars growing up to 20 m in height and ideally suited
for timber production; and the Peru type of medium-sized trees (Fig. 1 ) grow-
ing up to 10 m in height, yielding prolific quantities of palatable forage and
capable of withstanding repeated defoliation. Its common name is Leucaena
but in many countries local names prevail such as ipil-ipil (Philippines), lam-
toro ( I n d o n e s i a ) a n d k o a haole (Hawaii).
Many investigations have been conducted on the nutritive value and detox-
ification of L. leucocephala since the appearance of the early reviews on this
subject (National Academy of Sciences, 1977; D'Mello and Taplin, 1978). The
surge in interest in this tropical legume stimulated the launching of a new
journal in 1980 (Leucaena Research Reports ) and the compilation of compre-
hensive bibliographies on Leucaena research (Arellano, 1979; Oakes, 1982,
1983). This is, therefore, an opportune time to review the recent literature,
with particular respect to the role of Leucaena in poultry nutrition.
As stated previously (D'Mello and Taplin, 1978), Leucaena is widely grown
in the tropics as a fodder plant for ruminants. However, there is considerable
controversy about its nutritional value for this class of animal. Thus, in Mal-
awi, it has been shown that dried Leucaena leaf meal is equivalent to cotton-
seed cake as a source of protein for fattening beef cattle in stalls (Thomas and
Addy, 1977). On the other hand, ruminants consuming Leucaena in Papua
New Guinea {Holmes, 1981) and in Australia (Blunt and Jones, 1977; Jones
and Megarrity, 1983 ), show adverse signs such as alopecia, excessive salivation
and goitre associated with poor and highly variable live weight gains. This
discrepancy in responses of ruminants has been attributed to geographical dif-
ferences in rumen microbial ecology (Jones and Megarrity, 1986).
Current views on the role of Leucaena in poultry nutrition are no less con-
troversial, although the general consensus is that dietary Leucaena depresses
broiler growth and egg production in layer hens. However, there are differences
in toxicity between batches of Leucaena and, in any event, the recent Edin-
burgh experience suggests that a marked alleviation of this toxicity can be
effected by dietary manipulation. Much of the ensuing review refers to work
carried out in Edinburgh with Leucaena imported from Malawi and South East
Asia.
In Malawi, the shrub is cut and the branches are dried on concrete floors in
the sun. Once dried, the leaves are easily harvested by threshing. This dried
leaf meal may be incorporated directly into poultry diets. However, owing to
problems associated with its bulk, the leaf meal may be pelleted prior to trans-
port and storage.
In the Edinburgh studies, 4 batches of dried Leucaena leaf meal (LLM) of
the 'Peru' cultivar were subjected to detailed analysis in the laboratory. Of
these batches, two were sun-dried in Malawi in 1977 and 1979. The 1977 sam-
ple was used in the whole leaf (WL) form, without prior grinding. The 1979
batch was produced in 2 forms: a ground leaf (GL) form, and a powder form
( GP ), derived by grinding pellets of Leucaena leaf meal. A third Malawi sam-
ple, identical to that harvested in 1979, was prepared by drying fresh leaves in
an oven at 60°C. The fourth batch was sun-dried in Thailand during 1978.

CHEMICAL COMPOSITION

Proximate and mineral composition

The proximate and mineral contents of the 4 samples of LLM are shown in
Table 1 (D'Mello and Fraser, 1981 ). The sample from Thailand had the lowest
concentrations of crude protein, phosphorus, iron and zinc and the highest
4

TABLE 1

Proximate and mineral composition (dry matter basis) of four samples of dried Leucaena leaf
meal (‘Peru’ cultivar ) ’

Constituent Source and type of Leucaena leaf meal

Malawi Thailand

Sun-dried Sun-dried Oven-dried Sun-dried

1977 1979 1979 1978

Proximate analysis (g kg- 1)

Crude protein (N X 6.25) 294.1 291.0 281.3 224.4
Fibre 73.3 89.1 88.4 123.6
Ether extract 34.0 47.7 45.3 39.8
Ash 104.1 70.0 68.4 97.8
Major minerals (g kg-‘)
Calcium 23.3 18.1 18.5 23.7
Phosphorus 2.5 2.5 2.9 1.9
Sodium 0.4 0.0 0.1 0.2
Potassium 19.9 8.0 8.1 18.0
Magnesium 4.7 5.1 5.6 4.2
Trace minerals (mg kg- ’ )
Copper 7.0 9.9 10.6 9.7
Iron 407.1 239.9 191.0 181.2
Zinc 21.0 21.2 29.9 18.1
Manganese 60.0 42.1 46.0 49.2

‘D’Mello and Fraser (1981).

level of fibre in comparison with the African samples. However, in a Nigerian

study, Ekpenyong (1986) reported crude fibre concentrations of 182.4 g kg-’
dry matter (DM) for fresh Leucaena leaves but only 93.3 g kg-’ DM for sun-
dried leaves of the same batch. A striking feature of the mineral profiles is the
relatively large variation in calcium, potassium, iron and manganese concen-
trations which reflects the wide fluctuations in ash contents. As will be dis-
cussed later, the variations in iron content may have nutritional consequences.
In another Nigerian study (Adeneye, 1979) it was observed that ash content
increased from 52 g kg-’ DM in semi-open leaves to 90 g kg-’ DM in more
mature leaves of Leucaena . These results suggest that the 2 samples of LLM
harvested in Malawi during 1979 (Table 1) may have contained a higher pro-
portion of young leafy material than the batch harvested in 1977.

Grass and metabolisable energy content

Until recently, the apparent metabolisable energy (AME) value of LLM for
poultry was not well documented, despite interest in the use of this legume as
TABLE 2

Gross and apparent metabolisable energy (AME) values of Leucaena leaf meal (MJ kg- J DM)

Source and type of Leucaena leaf meal

Malawi Thailand

Sun-dried Sun-dried Oven-dried Sun-dried

1977 1979 1979 1979

Gross energy I 19.0 19.6 19.5 19.3

Malawi, sun-dried 1979

WL 2 Gp ~

Inclusion rate (g kg- 1 diet)

400 200 400

AME values :~
Experiment 1
Classical AME 2.4 2.3 2.2
N-corrected AME 2.8 4.0 3.6
Experiment 2
Classical AME 2.7
N-corrected AME 3.3

~D'Mello and Fraser (1981).

eWL, whole leaf; GP, ground form of Leucaena leaf pellets.
:~D'Mello and Acamovic (1982a).

a feeding stuff for farm livestock. Indeed, the only values available at the time
of the previous review (D'Mello and Taplin, 1978) were those of D'Mello and
Thomas (1978) who reported classical and N-corrected AME figures of 2.74
and 2.83 MJ kg-1 DM, respectively. In view of the need for more data, two
experiments were subsequently conducted by D'Mello and Acamovic (1982a)
in order to examine the effects of type of LLM and level of dietary inclusion
on AME values (Table 2). Two forms of LLM were assayed, WL and GP. Both
types were included in semi-purified diets at 400 g kg-I, and in addition, the
GP form was incorporated at 200 g kg- 1 diet.
It is clear that the classical values shown in Table 2 agree well with those
previously published by D'Mello and Thomas (1978). The N-corrected AME
values for the GP form of LLM (Table 2) are considerably higher than the
corresponding classical values. The differences are not readily explained. At
first sight, it would appear that the GP form may have exerted profound del-
eterious effects on the N metabolism of the chicks to the extent that N reten-
tion was severely reduced. However, this view should be balanced with the
observations that the determined classical and N-corrected AME values of
diets containing GP were very similar. In any event, the relevance of N-cor-
rected AME data is a matter of much debate (Vohra, 1972). For practical pur-
poses it is suggested that the classical values for LLM be used in formulating
poultry diets until the issues concerning N-corrections are resolved.
Whichever estimates are used, it is evident that the AME value of LLM is
inordinately low despite its relatively low fibre content (Table 1). In this re-
spect, the data shown in Table 2 may be compared with a value of only 5.35
MJ kg-1 DM for the pig (Gonzalez Vargas and Wyllie, 1982) and with values
ranging from 7.1 to 10.4 for ruminants fed on Leucaena forage in Malaysia
(Devendra, 1982). D'Mello and Thomas (1978) attributed the low AME fig-
ures to poor digestibility.

Amino-acid composition

Despite the considerable interest in Leucaena as a protein source, there ap-

peared to be insufficient information on the amino-acid content of this legume
with respect to variables such as cultivar, geographical origin and nature of

TABLE3

Amino -acid composition ( g kg - 1DM ) of four samples of dried Leucaena leaf meal ( 'Peru' cultivar) 1

Amino acid Source and type of Leucaena leaf meal

Malawi Thailand

Sun-dried Sun-dried Oven-dried Sun-dried

1977 1979 1979 1978

Aspartic acid 27.7 23.1 23.6 18.9

Threonine 13.5 12.1 11.9 8.7
Serine 13.3 12.0 11.5 9.8
Glutamic acid 37.4 35.1 32.7 28.2
Glycine 23.5 13.3 13.4 10.2
Alanine 17.6 12.7 12.2 11.3
Valine 15.2 14.4 13.0 10.1
Cystine 2.1 2.0 1.6 1.6
Methionine 5.4 4.6 4.6 2.3
Isoleucine 15.3 13.7 14.1 12.4
Leucine 22.6 21.7 24.2 16.0
Tyrosine 12.1 12.5 11.7 8.1
Phenylalanine 15.8 14.8 14.2 10.7
Lysine 15.5 17.6 16.9 12.8
Histidine 6.6 5.4 5.7 4.0
Arginine 16.4 15.1 15.8 10.2
Tryptophan 3.3 3.8 3.3 2.4

1D'Mello and Fraser (1981).

post-harvest treatment at the time of the last review (D'Mello and Taplin,
1978). Since then, more data have appeared and Table 3 summarises some
recent data (D'Mello and Fraser, 1981 ). These results highlight the wide vari-
ation in essential amino-acid content between different types and sources of'
LLM derived from the same cultivar. Differences are especially notable for
arginine, lysine, phenylalanine, tyrosine, leucine, methionine, cystine, glycine
and threonine, the Malawi samples having generally higher concentrations of
these amino acids than the Thailand sample. The isoleucine content of LLM
deserves some comment here, in view of the much higher value (24 g kg- i DM )
cited by the National Academy of Sciences (1977). This discrepancy is readily
accounted for by the method of amino-acid analysis. Studies at Edinburgh
(Acamovic and D'Mello, 1981a) indicated that unless special buffer gradients
were employed in the analysis of LLM hydrolysates by ion-exchange chro-
matography, isoleucine concentrations were grossly over-estimated owing to
the simultaneous elution of isoleucine with the unusual amino-acid, mimosine.
As will be demonstrated later, mimosine is present in appreciable concentra-
tions in the leaves and seeds of Leucaena.

Carotenoid concentrations

An important attribute of Leucaena is its relatively generous content of car-

otenoids. This class of compounds includes the carotenes, which can be con-
verted with varying efficiency by animals to vitamin A, and the xanthophylls,
which have no vitamin activity but which can be used by poultry as a source of
pigments. Egg yolk and broiler carcass colour are becoming increasingly im-
portant criteria of quality, and in many countries, there is a consumer prefer-
ence for egg yolks and chicken carcasses of a particular yellow colour. The
pigments deposited in eggs and carcasses cannot be synthesised by the fowl
and must, therefore, be supplied in the diet if an acceptable product is to be
obtained. Natural sources of the carotenoids are maize and forage crops, and
in addition, synthetic compounds are available such as carophyll yellow (fl-
apo-8' -carotenoic acid ethylester). As indicated previously (D'Mello and Tap-
lin, 1978) LLM is well endowed with fl-carotene and xanthophylls with con-
centrations ranging from 227 to 248 mg fl-carotene kg - i DM and 741-766 mg
total xanthophylls k g - ~ DM for different cultivars of Leucaena. Xanthophyll
levels in dehydrated alfalfa leaf meal are 400-550 mg kg- 1 DM (Scott et al.,
1969). Wood et al. (1983) reported even higher carotenoid levels in freshly
harvested LLM from Malawi. For example, rapid sun drying yielded LLM with
carotene and xanthophyll levels of 484 and 932 mg kg- 1 DM, respectively, but
substantial losses occurred during oven-drying at 60 and 145 ° C. Losses of car-
otenes during storage of LLM were of the order of 19-40 mg k g - 1 month 1,
and of xanthophylls in the range of 29-53 mg k g - 1m o n t h - ~. Carotenoids were
more stable in sun-dried LLM than in oven-dried samples. Pelleting or addi-
tion of ethoxyquin (an antioxidant) did not decelerate these carotenoid losses
during storage or processing (Wood et al., 1983).

TOXIC FACTORS IN LEUCAENA

It is now widely acknowledged that diets containing LLM invariably depress

performance of poultry (D'Mello and Taplin, 1978; D'Mello and Acamovic,

TABLE4

Daily weight gain (WG, g per chick), daily food intake (FI, g DM per chick) and efficiency of
food conversion (EFC, g gain g- ~DM consumed ) of chicks fed on a control diet or diets containing
Leueaena leaf meal (LLM) 1

Diet WG 2 FI 2 EFC 2

Control 54.8 88.4 0.620

L L M '*diets
50 GL 53.4 90.0 0.594
100 GL 47.4 86.6 0.547

~D'Mello et al. (1987).

~Period of observation: 22-31 days of age.
:~GL: ground leaf included at 50 or 100 g kg- 1diet.

TABLE5

Antinutritional components of Leucaena leucocephala

Component Part of plant Concentration 1

(g k g - ' DM)

Mimosine Leaf 10-120

Seed 33-145
3-Hydroxy-4 (1H)-pyridone Leaf 5.1-8.2
Seed Not detected
Tannins Leaf 13-44
Seed 7.1
Trypsin inhibitors Leaf Weak activity
Seed Strong activity
Galactomannan gums Leaf 46
Seed 320
Saponins Leaf 2-11
Seed 2-11
Flavonols Leaf 30-60
Seed Not detected
Haemagglutinins Leaf Not tested
Seed Low activity

~See text for sources of data.

 9

TABLE 6

Mimosine and tannin contents (g kg- 1DM ) of four samples of Leucaena leaf meal ('Peru' cultivar )

Source and type of Leucaena leaf meal

Malawi Thailand

Sun-dried Sun-dried Oven-dried Sun-dried

1977 1979 1979 1978

Mimosine 10.2 25.5 14.7 14.1

Tannin
AOAC method 2 21.7 33.6 24.3 25.5
Price and Butler method :~ 13.3 43.6 11.6 15.6

'D'Mello and Fraser ( 1981 ).

2As gallotannic acid.
:~As catechin.

1982a; ter Meulen et al., 1984). The toxic nature of LLM appears to be unaf-
fected by age of poultry and is not always mediated via depressions in food
intake. Thus D'Mello et al. (1987) have shown that broilers at 22 days of age,
while able to maintain satisfactory food intakes on a diet containing 100 g
LLM kg -1, still exhibit severe depressions in growth and efficiency of food
utilisation (Table 4). The deleterious effects of Leucaena have been attributed
to the presence of antinutritional or toxic factors (D'Mello and Taplin, 1978;
ter Meulen et al., 1984).
In common with many crop plants, particularly legumes, Leucaena contains
a wide variety of compounds which are capable of inducing adverse effects in
animals.
The toxic components of Leucaena are presented in Tables 5 and 6.

M imosine and 3-hydroxy-4(1H)-pyridone

It has been suggested frequently that the relatively poor nutritive value of
Leucaena arises principally from its content of the toxic amino acid, mimosine
and its immediate degradation product, 3-hydroxy-4 (1H)-pyridone (DHP)
(Ross and Springhall, 1963; Librojo and Hathcock, 1974; D'Mello and Taplin,
1978). For many years, studies on the metabolism and role of these compounds
have been thwarted by the lack of suitable and specific methods for their de-
termination. The traditional FeCl:~ colorimetric assay for mimosine (Matsu-
moto and Sherman, 1951 ) is influenced by factors such as pH, the presence of
other phenolic compounds such as tannins and DHP, and variability in recov-
eries caused by the use of charcoal in the decolourisation procedure (Hegarty
et al., 1964; Megarrity, 1978). These disadvantages highlighted the need for
10

alternative and specific methods of analysis. Acamovic and D'Mello (1981a)

developed a specific ion-exchange chromatographic technique for the deter-
mination of mimosine. The advent of this method permitted studies of the
mimosine content of Leucaena and of the fate of ingested mimosine in the fowl
(D'Mello and Acamovic, 1982b ). However, the ion-exchange chromatographic
method was slow and, more importantly, could not be used for the determi-
nation of DHP. These disadvantages were overcome when Acamovic et al.
(1982) subsequently developed a high performance liquid chromatographic
method which allowed the simultaneous estimation of mimosine and D H P
within a few minutes. Chromatographic procedures for these compounds have
also been developed by Tangendjaja and Wills (1980) and by Lowry et al.
(1985).
Mimosine concentrations (Table 5) are generally higher in the seed (Aca-
movic et al., 1982; Szyszka and ter Meulen, 1984; Chandrasekharan and Dam-
othiran, 1985) than in the leaf (Table 6; Acamovic and D'Mello, 1981a; Hut-
ton, 1985). The toxicity of mimosine for poultry has not been established beyond
reasonable doubt. Springhall (1965) demonstrated that the adult cockerel is
capable of metabolising a single oral dose of mimosine without adverse con-
sequences. In relatively long-term studies Tangendjaja and Sarmanu (1986)
showed that pure mimosine did not affect the onset of sexual maturity in laying
hens. However, our unpublished investigations show that the young chick is
more sensitive to pure mimosine than the older bird. Both growth rate and
food intake were severely reduced in chicks given 3.3 g mimosine kg -1 diet.
Previous conclusions regarding the toxicity of mimosine were based on studies
involving feeding with LLM or with a mixture of LLM and Leucaena seeds.
The interpretation of such investigations is fraught with difficulties. Thus ter
Meulen et al. ( 1984 ) observed that a dietary mimosine concentration of 4.94 g
kg- 1 severely retarded growth and food intakes of chicks when a mixture of
LLM and Leucaena seeds, in the ratio of 36:64, was included at 200 g k g -
diet. While these adverse effects may be attributed to the intake of mimosine,
it should be noted that mimosine from LLM is but poorly absorbed (D'Mello
and Acamovic, 1982b ) and that Leucaena seeds are well endowed with trypsin
inhibitors and galactomannan gums (Table 5 ) which also could have contrib-
uted to the toxicity of the mixture used by ter Meulen et al. (1984).
Most, if not all, of the D H P in LLM probably arises from the post-harvest
enzymatic degradation of mimosine (Table 5; T. Acamovic, unpublished data;
Hegarty et al., 1964; Lyon, 1985; Wee and Wang, 1987). Although D H P has
potent goitrogenic properties in mammals (Christie et al., 1979; Jones and
Megarrity, 1983) its effects in poultry remain obscure. It is likely that D H P
plays a minor role in the toxicity of LLM for poultry since D H P intakes are
low and mimosine is relatively stable, even within the gastrointestinal tract of
the bird (D'Mello and Acamovic, 1982b). However, it has been reported that
the performance of broilers and ducks was reduced when mimosine in LLM
 11

had been enzymatically degraded to DHP prior to dietary incorporation (Tan-

gendjaja and Lowry, 1984). Although these authors postulated that DHP might
act as an appetite depressant, it is possible that enzyme treatment may have
induced other deleterious changes in the composition and nutrient availability
of LLM.

Tannins

As shown in Tables 5 and 6, tannin concentrations are higher in LLM

(D'Mello and Fraser, 1981 ) than in the seed (T. Acamovic, unpublished data).
The precursors and hydrolysate products of the tannin polymers, namely ca-
techin, epicatechin, cyanidin and procyanidin B-2 have all been found in LLM
(D. Kufidis, personal communication). Tannins reduce digestibility by bind-
ing to dietary proteins within the alimentary canal and to proteins in digestive
secretions (Hewitt and Ford, 1982 ). The poor N retention of birds given LLM
and the markedly low AME value of LLM for poultry may, at least in part, be
attributed to the tannin components (D'Mello and Thomas, 1978; D'Mello and
Acamovic, 1982a).

Trypsin inhibitors

Limited work (Acamovic and D'Mello, 1984) suggests that the seed displays
considerably higher trypsin inhibitor activity than the leaf (Table 5 ). Trypsin
inhibitor activity in Leucaena seeds is of a similar magnitude to that found in
other legume grains (Valdebouze et al, 1980; Kadam et al., 1987). Since trypsin
inhibitors can severely reduce protein utilisation (D'Mello et al., 1983 ), their
presence in the seed of Leucaena is likely to be an important factor contribut-
ing to its toxicity (ter Meulen et al., 1984).

Galactomannan gums

Polymeric carbohydrates such as galactomannan gums occur widely in leg-

umes including Leucaena (Table 5;~Lyon and Kohler, 1981; Arora and Joshi,
1984). The reduced food intakes'hnd growth rates of broilers fed on guar seed
( Cyamopsis tetragonoloba) have been ascribed to its content of galactomannan
gums (Verma and McNab, 1982). These gums increase the rate of excretion of
bile acids (Gee et al., 1983) and, in addition, possess weak haemagglutinating
properties (Lesniak and Liu, 1981 ). Galactomannans may play an important
role in the toxicity of Leucaena seeds.
12

Saponins

Although initial observations suggested the absence of significant quantities

of saponins in LLM (Tangendjaja et al., 1984), our unpublished data indicate
the presence of up to 11 g kg -1 DM in Leucaena leaf and seed. At these con-
centrations, however, saponins are unlikely to contribute significantly to the
toxicity of this legume.

Flavonols

Lowry et al. (1984) found flavonols in Leucaena leaf at concentrations of up

to 60 g kg-1 DM (Table 5). The major fiavonol compounds identified in LLM
are glycosides of quercetin and myricetin.

Haemagglutinins

The haemagglutinins (or lectins) present in many legume grains are capable
of inducing acute antinutritional effects in animals (Grant et al., 1985). How-
ever, there appears to be only weak haemagglutinating activity in Leucaena
seeds (Table 5) possibly arising from the galactomannan components (Les-
niak and Liu, 1981; Lyon and Kohler, 1981 ).

DETOXIFICATION OF LEUCAENA

The presence of such a wide range of deleterious compounds in Leucaena

poses a challenging problem of detoxification. Attempts to improve the nutri-
tional value of Leucaena may be classified into 3 main categories.

Genetic improvement

This is the most obvious method of detoxification. It is well recognised that

substantial variations occur in the mimosine content of different cultivars of
Leucaena (Krishnamurthy and Mune Gowda, 1983; Hutton, 1985). Consid-
erable differences also occur between different batches of the same cultivar
(Table 6). While there can be little doubt that genetic methods have a role to
play in legume detoxification, it is important to recognise potential problems
associated with such methods. Firstly, there is good evidence that many of the
toxic factors present in legumes confer insect and fungal resistance to the plant
and the removal of these substances could result in agronomic disadvantages.
Thus, insect infestation of Leucaena by Psyllidae is now widespread in the
tropics (Beardsley, 1986) and attempts to develop lines low in mimosine and
tannins are likely to compound this problem (Lowry et al., 1986). Secondly,
genetic improvement is relatively slow, being an essentially long-term exercise.
 13

Finally, the breeding of cultivars low in an individual toxic component may

not automatically improve nutritional quality. Many enzyme inhibitors, such
as the protease inhibitors, contain relatively large quantities of the amino acid,
cysteine. It is possible, therefore, that a reduction in protease inhibitor content
by genetic manipulation might be accompanied by a deleterious change in
amino-acid composition. Furthermore, the exact identity of the main toxic
component in Leucaena must be known with a greater degree of certainty if
plant breeding techniques are to have any chance of success. Thus, although
mimosine is frequently implicated as the main cause of the poor nutritional
value of LLM for poultry, the experimental evidence for such a conclusion is,
at best, tenuous.

Effect of treatment during processing

Since a number of toxic factors in Leucaena are sensitive to heat (D'Mello,

1982), the application of heat during processing of the leaf should provide a
rapid and effective means of inactivating these toxic compounds. Experience
with grain legumes indicates that the destruction of heat labile inhibitors dur-
ing cooking is governed by factors such as temperature, duration of heating,
particle size and moisture conditions. It is widely acknowledged that moist
heating procedures are generally more effective than dry heating methods, and
there may be further beneficial effects by soaking prior to cooking (D'Mello et
al., 1983; Kadam et al., 1987).
While nutritional benefits can accrue from heat treatment, this method is
not without its limitations. For example, excessive temperatures during heat
processing can reduce the availability of amino acids (Carpenter and Booth,
1973). The correct application of heat is, therefore, crucial to the preparation
of legume feeding stuffs. Furthermore, although heating can reduce mimosine
concentrations in LLM (Table 6) it is worthwhile recalling that this disap-
pearance results from conversion of mimosine to DHP, a heat-stable goitrogen.
Other post-harvest treatments suggested recently are ensilage (Lyon, 1985 ),
wilting and leaching (Szyszka et al., 1983) of Leucaena. All these treatments
undoubtedly reduce mimosine concentrations. However, since it is more than
likely that D H P concentrations are proportionately increased, the value of
such treatments remains questionable, at least until animal studies are con-
ducted to verify any improvements.

Dietary manipulation

The commercial use of dietary additives to improve the nutritional value of

feeding stuffs is a comparatively recent innovation and much of the work in
this field is still at an experimental stage. Additives are generally used with the
14

aim of minimising the effects of the heat-stable toxic components of food crops,
although they may be effective against some heat-labile inhibitors as well.
The inactivation of tannins in legume and cereal grains has been attempted
by the use of polyvinylpyrrolidone and polyethyleneglycol (PEG). It is be-
lieved that tannins bind more strongly to these additives than they do to pro-
teins. Hewitt and Ford (1982) concluded that dietary supplementation with
PEG of molecular weight 4000 was a simple and cost-effective means of im-
proving the nutritional value of high tannin grains without sacrificing the ag-
ronomic advantages of high tannin cultivars.
As discussed previously (D'Mello and Taplin, 1978), attempts to improve
the nutritional value of LLM by dietary supplementation have consisted of the
use of ferrous sulphate or of structural analogues of mimosine. The efficacy of
ferrous sulphate is well recognised (Ross and Springhall, 1963; Gloria et al.,
1966), although it has been claimed that pre-treatment of LLM with ferrous
sulphate in solution prior to dietary incorporation is an important pre-requi-
site (Ross and Springhall, 1963 ). Following the observations of Lin et al (1964)
that dietary supplements of L-phenylalanine and L-tyrosine could partially or
completely reduce the toxicity of mimosine in rats On the basis of structural
similarities of the three amino acids, Labadan (1969) attempted to improve
the nutritional value of LLM for chickens by this method. Tyrosine and other
putative analogues of mimosine were totally ineffective.
Since the digestibility of LLM is relatively poor, the possibility exists that
some amelioration of its adverse effects may be achieved by increasing nutrient
concentrations of diets containing this legume (D'Mello and Thomas, 1978).
Such an effect has, indeed, been observed. Thus Gloria et al. {1966) reported
a reduction in LLM toxicity on increasing dietary concentrations of protein,
energy and methionine.

The Edinburgh studies

An integrated approach has been adopted in the detoxification of LLM im-

ported from Malawi. The 'Peru' cultivar was introduced into Malawi on the
basis of its relatively low mimosine content. Indeed, one batch harvested in
1977 contained only 10.2 g mimosine kg -1 DM (Table 6) and, as such, ap-
peared to be a suitable product for studying the effects of a low mimosine cul-
tivar on growth of broiler chicks (D'Mello and Acamovic, 1982b). The WL
form of this batch of LLM was incorporated at a rate of 150 g kg- 1 diet, the
inclusion being made at the expense of maize and soya bean meal in a control
diet, Owing to the strikingly low AME value of LLM (Table 2), and in order
to maintain uniform dietary energy concentrations, all diets containing the
leaf meal were supplemented with maize oil. This study also provided the op-
portunity to investigate the effects of a number of additives on the nutritive
value of LLM. Thus an enzyme mixture consisting of cellulase, acid and neu-
 15

tral proteases, and a-amylase was employed in an effort to enhance digestibil-

ity. Another dietary treatment involved the addition of ferrous sulphate
(FeSOt-7H20) in dry form, without the pre-treatment suggested by Ross and
Springhall (1963). The final additive tested in this experiment was aluminium
sulphate (A12(SO4)3"16H20) following the report by Tsai and Ling (1973)
that A1 ions formed stronger complexes with mimosine than ferrous ions. The
additives were tested singly and in certain combinations.
As shown in Table 7, the 1977 low-mimosine batch of LLM promoted highly
satisfactory growth performance of broiler chicks. Thus, there were no appre-
ciable differences in weight gain or efficiency of food utilisation between any
of the groups given LLM and the control birds fed on the maize-soya bean
meal diet. However, food intake was reduced in groups fed on the LLM diets
supplemented with FeSO4 alone or in combination with A12 (SO4):~. In the ab-
sence of any growth depressing effects of the 1977 batch of LLM, the effec-
tiveness of the various additives employed in the investigation remained un-
resolved. It was anticipated that the addition of enzymes might have enhanced
growth through improvements in digestibility of LLM. However, in retrospect,
it appears that the transit time of food in the gut was probably too brief to
allow any beneficial response from the relatively slow-acting enzymes. AI-
though the metal ions employed in the above-mentioned experiment (Table
7) had little effect on growth performance of chicks fed on the low-mimosine
batch of LLM, mimosine output in the excreta was greatly enhanced by the
dietary addition of either FeSO4 or A12 (SO 4 )3- As predicted by Tsai and Ling

TABLE7

Daily weight gain (WG, g per chick), daily food intake (FI, g DM per chick), efficiency of food
conversion (EFC, g gain g-1 DM consumed) and mimosine output: intake ratios ( M O / M I , g
mimosine voided g-I mimosine consumed) of chicks fed on a control diet or diets containing
Leucaena leaf meal (LLM)

Diet WG 2 FI e EFC 2 MO/MV

Control 25.8 37.7 0.682

LLM diets
Basal 150WL :~ 25.4 36.6 0.693 0.781
Basal 150WL+enzymes4(5 g k g - 1diet) 25.6 36.7 0.696
Basal 150 WL+FeSO4"7H~O (5g kg 1 diet) 24.1 35.6 0.677 0.881
Basal 150WL + A12 (SO4):~. 16H~O (11.7 g kg- ~diet ) 24.1 36.4 0.661 1.003
Basal 150WL + FeSO4 + A12 (SO4)3 23.6 34.6 0.681
Basal 150WL + FeS04 + A12 (S04):~ + enzymes 24.8 36.9 0.668

~D'Mello and Acamovic ( 1982b ).

~WG, FI and EFC measured during the period 7-20 days of age; MO/MI determined between 15
and 16 days of age.
:~WL: whole leaf (1977 batch) included at the rate of 150 kg kg ~diet; mimosine content of LLM:
10.2gkg ~DM.
'~Enzyme mixture contained cellulase, acid and neutral proteases and o~-amylase.
16

(1973), the A1 salt was much more effective than FeSO4,the former inducing
total excretion of ingested mimosine. The increases in mimosine excretion were
achieved despite the use of the dry forms of these salts, and without any pre-
treatment of the LLM prior to dietary incorporation. Furthermore, it is evident
that the fowl has an inherent propensity to excrete large proportions of in-
gested mimosine, unaided by metal ions (Table 7). The increased excretion of
mimosine on supplementation with metal ions is generally attributed to che-
lation of mimosine by these minerals. However, it is also likely that these salts
alter dietary pH and ionic balance to such an extent that microbial breakdown
of mimosine in the gut is curtailed, thus enhancing mimosine excretion.
It is now apparent that the 1977 low-mimosine WL form of LLM was a
rather unique sample. In addition to its relatively low content of mimosine,
this batch had the lowest tannin (Table 6) and fibre contents and the highest
iron concentrations (Table 1 ) of all LLM samples analysed in our laboratory.
Subsequent samples of the same cultivar proved to have much higher concen-
trations of mimosine (Table 6) tannins and fibre (Table 1). The reasons for
the increase in deleterious constituents are not known, but may be related to
seasonal factors, to soil conditions, and to variations in the proportions of the
leaf and stem components.
The higher mimosine contents of the 1979 batches (Table 6) allowed further
investigations of the ability of the bird to dispose of dietary mimosine. The
results of such a study were reported by D'Mello and Acamovic (1982b). In
that investigation, different dietary concentrations of the 1977 and 1979 batches
were used for young chicks, thus ensuring varying intakes of mimosine. As in
the earlier experiment reviewed in this paper, the LLM diets were supple-
mented with maize oil to ensure uniform dietary AME concentrations. Table
8 shows that the GP form appreciably reduced growth rate and food intake
when increased from 50 to 100 g kg -1 diet. As before (Table 7) substantial
proportions of ingested mimosine appeared in the excreta, although it is not
clear why the GP form should have elicited a higher excretion of mimosine
than the GL form (Table 8). The relationship between mimosine output in
the excreta and mimosine intake is more clearly seen in Fig. 2 which incorpo-
rates data from the unsupplemented LLM groups in the two experiments re-
viewed in this paper (Tables 7 and 8). It is apparent from this relationship
that, irrespective of the type of LLM, a substantial proportion of ingested mi-
mosine is voided in the excreta. Furthermore, the excretion of mimosine is
essentially linear over the range of intakes tested (Fig. 2; D'Mello and Aca-
movic, 1982b).
The relatively high excretion rates of ingested mimosine recorded in the
investigations just described contrast with the low excretion rates of orally
administered mimosine observed by Springhall (1965) in surgically modified
cockerels. This discrepancy may have arisen through the use of crystalline
mimosine by Springhall (1965). It is possible that the absorption of mimosine
 17

TABLE8

Daily weight gain (WG, g per chick), daily food intake (FI, g DM per chick), efficiency of food
conversion (EFC, g gain g-~ DM consumed) and mimosine output: intake ratios (MO/MI, g
mimosine voided g ~ mimosine consumed) of chicks fed on a control diet or diets containing
Leucaena leaf meal (LLM)

Diet WG 2 FI 2 EFC 2 MO/MI 2

Control 24.4 36.0 0.679

LLM diets
50 WL :~ 25.5 36.9 0.692 0.769
100 WL 25.9 37.3 0.693 0.820
50 GP ~ 26.5 37.9 0.697 0.924
100 GP 22.9 33.4 0.684 0.916
100 GL '~ 24.0 34.5 0.691 0.708

1D'Mello and Acamovic (1982b).

~See h)otnote to Table 7.
:~WL: whole leaf (1977 batch) included at 50 or 100 g kg-1diet; mimosine content of WL, 10.2 g
kg 1DM.
4Gp: powder produced by grinding pelleted Leucaena leaf {1979 batch); rate of incorporation: 50
or 100 g kg- 1diet; mimosine content of GP, 25.5 g kg- 1 DM.
'~GL: ground leaf (1979 batch) included at 100 g kg-~diet; mimosine content of GL, 25.5 g kg-
DM.

4O0
o

.E 300

~ ~ 2o0
E~
E~
6 100
~ y= 0"826x-0'661
r = + 0.956
i I I I I
100 200 300 400 500
Duily mimosine intuke (rag replicate -1)

Fig. 2. Daily mimosine output in excreta, between 15 and 16 days of age, of young chicks consuming
different quantities of mimosine derived from Leucaena leaf meal (D'Mello and Acamovic, 1982b).

f r o m L L M is l o w e r t h a n t h a t o f p u r e m i m o s i n e b e c a u s e o f the relatively poor

digestibility of LLM (D'Mello and Thomas, 1978).
The GP form of LLM depressed performance despite the ability of chicks to
d i s p o s e o f o v e r 0.90 o f i n g e s t e d m i m o s i n e ( T a b l e 8 ) . F u r t h e r experiments were,
therefore, conducted to evaluate methods of detoxification. In one such exper-
 18

.'•_•100
13

°v 9O

80
o

g
70

c~

5c I |
0 20 ;4
Ferric sulphate additions (g kg -1 diet)

Fig. 3. Effect of ferric sulphate and polyethylene glycol (PEG) on relative growth rates of young
chicks fed on diets containing Leucaena leaf meal at 150 g kg -1 diet (PEG-free diets, &; diets
containing 20 g PEG kg-1, O; Acamovic and D'Mello, 1981b). Mimosine content of Leucaena,
25.5 g kg-1 DM.

13 I i! raw LLM
f r
autoclaved LLM
i raw LLM ferric sulphate
+
l i !
i I
I r. . . . . .
raw LLM+ferric sulphate + PEG
c 9C i I
.o
o
I J
\\\k\\'
. ." ".." ". LkXXX\\- autoclaved LLM +ferric
I I I . . ",o. . "o.. ' \ \ \ \ \ \ ' sulphate + PEG
m I I . . . . . xxxxx\'
o i j aI ..::..::. ,,,,,\,,\.
ii I x, \N\ \ \\ \\ '\ \ '
~,\\\\\,
I ,,\\\\\,
II I "\\\\\"
._~ I I ~\\\\\"
I ~\\\\\,
IE

"F_

,\\-.,\\,~
~\\\\\x
,\\\\\',
,\\\\\',

Treatment/additives

Fig. 4. Effects of ferric sulphate and polyethyleneglycol (PEG) additions on the relative nutri-
tional value of diets containing raw or autoclaved Leucaena leaf meal (LLM) at 150 g kg-1 diet.
Relative nutritional value assessed by comparing growth rates of LLM-fed chicks with those of
chicks fed on a soya bean control diet (D'Mello, 1982).
 19

iment (Acamovic and D'Mello, 1981b), ferric sulphate (Fe2(SO4):~'9 H20)

was tested as a dietary additive on the basis that mimosine forms a more stable
complex with the ferric than the ferrous ion (Tsai and Ling, 1973 ). Since LLM
also contains tannins (Table 6), PEG was tested as a further supplement.
Graded supplements of ferric sulphate in the absence and presence of PEG
were added to diets containing 150 g LLM (type GP) kg-1. Diets were made
isoenergetic by the addition of maize oil to all LLM diets. As shown in Fig. 3,
growth of chicks was improved substantially by dietary supplements of the
ferric salt. Further improvements in growth were achieved by combined addi-
tions of ferric sulphate and PEG, the latter supplement being totally ineffec-
tive in the absence of iron. The combined supplements restored growth of chicks
given LLM to 0.9 of that attained by birds fed on a conventional maize-soya
bean control diet.
The possibility of denaturing heat-labile inhibitors by autoclaving mois-
tened LLM and drying at low temperature prior to dietary incorporation was
investigated in another experiment. The results are presented in Fig. 4 which
also illustrates in general terms some of the advances which have recently been
made in the detoxification of the GP form of LLM. Autoclaving undoubtedly
improves the nutritional properties of LLM, which is consistent with recent
observations on the occurrence of trypsin inhibitors in this product (Acamovic
and D'Mello, 1984). However, the beneficial effects of cooking disappear when
the LLM diet is supplemented with ferric sulphate and PEG (Fig. 4). The
proposition which may, therefore, be considered is that LLM contains trypsin
inhibitors, primarily in the form of heat-labile phenolics, and that once these
are complexed with dietary additives, heat treatment is unnecessary. Auto-
claving on its own is not as effective as ferric sulphate and PEG additions.

LEUCAENA AS A SOURCE OF PIGMENTING XANTHOPHYLLS

Although the beneficial effects of dietary LLM on egg yolk colour have been
recognised for a number of years (D'Mello and Taplin, 1978) quantitative data
on the pigmenting potency of LLM relative to other natural and synthetic
sources were not available until 1981. In a study described by Taplin et al.
(1981), two levels of LLM and of grass meal (GM) were added to a low pig-
ment (LP) basal diet. The LLM additions were 8.8 and 17.6 g k g - 1diet, which
provided 7.5 and 14.9 mg xanthophylls k g - 1 DM. Inclusion rates of GM were
14.4 and 29.1 g kg -1 diet, supplying 7.6 and 12.2 mg xanthophylls kg -1 DM.
The pigmenting efficiency of these diets was compared with that of another
diet containing a synthetic compound, carophyll yellow (CY). The latter diet
was prepared by adding 0.1 g CY kg-1 LP basal diet. These diets were given to
laying pullets for a period of 4 weeks and egg yolk colour was measured chem-
ically as concentrations of xanthophylls and fl-carotene equivalents (BCE).
Yolk colour was also assessed visually using a 15-blade Roche Colour Fan.
20

7C

6C

>~ 5C

~ 4o
o

8
m
2C

kLl
1C

o 15 31o
Days after introduction of experimental diets

Fig. 5. Changes in egg yolk pigmentation over a 28-day feeding period. Hens were offered one of
the followingdiets: low pigment (LP), O; LP+8.8 g Leucaena leaf meal (LLM) kg-] diet, A;
LP + 17.6 g LLM kg- 1diet, • ; LP + 14.4 g grass meal (GM) kg- ~diet, []; LP + 29.1 g GM kg- ]
diet, . ; LP -t-0.1 g carophyll yellowkg- ] diet, O. Egg yolk pigmentation expressed as pg fl-caro-
tene equivalents (BCE) g- 1yolk (Taplin et al., 1981).

Dietary deprivation of xanthophylls, caused by the L P diet, induced a rapid

decline in egg B C E concentrations, stable values of less than 5/~g g-1 yolk
being attained within a period of 15 days (Fig. 5). Supplementation of the L P
basal diet with LLM, improved yolk B C E concentrations in proportion to the
xanthophyll content of the additions. However, neither supplement of L L M
was effective in raising yolk B C E concentrations to levels recorded at the be-
ginning of the experiment. The addition of the lowest level of G M enhanced
yolk B C E concentrations, the improvement matching that induced by the
highest level of L L M addition. Nevertheless, yolk B C E concentrations in hens
given G M remained below the initial values. However, at the higher rate of G M
inclusion, yolk BCE concentrations rose steadily above initial values and the
advantage over those groups given the two L L M supplements or the lower G M
supplement was sustained throughout the experiment. The addition of CY elic-
ited a sharp increase in yolk B C E content, and although these levels declined
markedly towards the end of the experiment, the yolks were still substantially
more pigmented than those from the other experimental group (Fig. 5).
 21

'FABLE 9

Terminal visual egg yolk scores of hens fed on a low pigment ( L P ) basal diet supplemented with
Leucaena leaf meal ( L L M ) , grass meal ( G M ) or carophyll yellow (CY)

Diet Visual yolk score

(Roche 15-blade Fan)

Mean value Range

L P only 1.7 1-2

L P + 8.8 g L L M kg -1 4.2 4-5
L P + 17.6 g L L M kg -1 5.2 5-6
L P + 14.4 g G M k g - 1 5.0 3-6
L P + 2 9 . 1 g G M kg - I 7.1 5-8
L P + 0 . 1 g CY kg -~ 11.3 11 12

'Taplin et al. (1981).

The differences in pigmenting efficiency of LLM and GM were also detected

visually as shown by the Roche Fan scores determined at the end of the exper-
iment (Table 9).
A quantitative assessment of the xanthophyll availability in the two meals
was obtained by measuring xanthophyll concentrations in egg yolks collected
during the final week of the experiment. These concentrations were plotted
against xanthophyll intakes of hens in the LP, LLM and GM groups (Fig. 6).
Regression lines constrained to meet at the same intercept were fitted to the
data for the LLM and GM groups, the data from the LP group contributing to
both regression lines. There was a statistically significant difference between
the slopes of the lines shown in Fig. 6. The equations shown in this figure
predict that the xanthophyll content of egg yolks during the final week would
be 12.2 and 23.3 pg g- 1, respectively, for hens fed on LLM and GM and con-
suming 0.8 mg xanthophylls daily. Since the average weight of yolk in the ex-
periment was 16.3 g, corresponding xanthophyll output in yolk would be of the
order of 0.198 and 0.380 mg day- 1, yielding deposition efficiency values of 0.25
and 0.47, respectively, for the LLM and GM diets. On the basis of these results,
Taplin et al. (1981) concluded that although LLM had approximately twice
the xanthophyll content of GM, the availability of these pigments in the for-
mer product was only about half that found in GM.
The relatively low xanthophyll availability in LLM has been attributed to
its comparatively high tannin content (Taplin et al., 1981). There is ample
evidence that dietary tannic acid and tannins interfere with pigment metabo-
lism in hens (Hughes, 1973) resulting in declining Roche Fan scores and an
increase in the incidence of mottling in egg yolks (Armanious et al., 1973).
In a separate study, Berry and D'Mello (1981) confirmed the low xantho-
22

50I
~ 40

30

6u

~- 20

LIJ

I I I I
0.4 0,8 1.2 1,6
Daily xanthophylL intake ( r a g h e n -1)

Fig. 6. The relationship between daily xanthophyll intake and egg yolk xanthophyll content. The
regression equations, y = 0.881 + 14.098x and y = 0.881 + 28.036x (combined r = 0.89) were derived
for groups fed Leucaena leaf meal (/x, • ) and grass meal (C], • ) , respectively. See legend to Fig.
4 for details of diets (Taplin et al., 1981 ).

Daily xanthophyll intake (mg chick -1)

Fig. 7. The relationship between daily xanthophyll intake and Roche Fan scores of broiler chicks.
Diets offered were: low pigment, I ; yellow maize, • ; Leucaena leaf meal, • (D'Mello et al., 1987).

phyll availability in LLM but were able to demonstrate enhanced yolk pig-
mentation in hens fed on LLM diets supplemented with coconut oil. It is known
that transmission of pigments from the diet to the yolk is influenced by level
and fatty-acid composition of dietary lipids (Abu-Serewa, 1976).
 23

The utilisation of LLM xanthophylls by broiler chicks has received scant

attention. With increasing commercial interest in broiler pigmentation in the
U.K., a preliminary experiment has recently been completed (D'Mello et al.,
1987), using an oven-dried sample of 'Peru' LLM imported from the Philip-
pines. The pigmenting potency of this LLM sample was compared with that
of yellow maize by relating xanthophyll intake of broilers to Roche Fan scores
of their shanks and feet (Fig. 7). It is apparent that although LLM elicits
higher scores than maize, the efficiency of utilisation of dietary xanthophyll is
similar for both ingredients since all data lie on a single smooth curve. These
results present LLM in a more favourable light than the studies of Berry and
D'Mello ( 1981 ) and Taplin et al. ( 1981 ). It is clear, however, that within the
normal rate of dietary inclusion, LLM is unlikely to promote satisfactory broiler
carcass colour on its own. With wheat-based diets, it is inevitable, therefore,
that LLM would be used in conjunction with other xanthophyll sources. Under
these conditions it would be prudent to restrict LLM inclusion rates to 50 g
k g - 1 diet since growth performance remains unaffected (Table 4 ) and there is
no clear benefit, in terms of Roche Fan scores, from higher inclusion rates (Fig.
7). Furthermore, any mimosine residues in the carcass would also be reduced
by restricting dietary inclusion rates of LLM.

SAFETY EVALUATION

The observations of ter Meulen et al. (1984) that mimosine accumulates in

body organs of broilers fed on 'Leucaena meal' raises the important issue of
safety evaluation. Although these authors recorded unacceptably high varia-
tions within groups of broilers given graded dietary concentrations of the meal,
the conclusion that mimosine accumulates in visceral and other organs is ines-
capable. It is worth noting, however, that the 'Leucaena meal' used by ter Meu-
len et al. (1984) consisted of a mixture of seeds and leaves in the proportion
64:36. Since the seeds are probably much more digestible and contain more
mimosine than the leaves (Table 5 ), it is conceivable that this amino acid was
absorbed rapidly and efficiently under these conditions, leading to accumula-
tion in various organs.

CONCLUSION

The nutrient specifications of LLM and Leucaena seeds are now well defined
as a consequence of sustained research efforts over the past decade. The out-
standing issue concerning the role of LLM in poultry diets is its low AME
content which ranges from 2.2 to 2.7 MJ k g - 1 DM. Recent results indicate that
the lignin content of LLM can be as high as 140 g kg-1 in the Peru cultivar
24

(Arora and Joshi, 1986). It is salutary to note that the digestibility in vitro of
paper by rumen micro-organisms declines from 0.99 to 0.65 when lignin con-
tent increases from 30 to 90 g kg-1 DM (van Soest and McQueen, 1973). In
poultry, this reduction in digestibility is likely to be more pronounced, since
microbial digestion is minimal, resulting in inordinately low AME values for
LLM (D'Mello and Acamovic, 1982a). Thus LLM is intrinsically of low nu-
tritional value, a property which it shares with other leaf meals such as those
derived from cassava (Manihot esculenta; Ravindran et al., 1983) and alfalfa
(Medicago sativa; Scott et al., 1969).
Recent advances in analytical chemistry have facilitated the publication of
a comprehensive set of data on the concentrations of mimosine, DHP, tannins,
trypsin inhibitors, galactomannan gums, saponins and flavonols in Leucaena
leaf meal and seeds. The toxicity of mimosine is frequently invoked as the
mechanism whereby LLM reduces performance of poultry. However, the ar-
guments for such a mode of action are not based on direct evidence and should
be reconsidered in the light of recent observations. Since LLM is poorly di-
gested (D'Mello and Thomas, 1978), it might be expected that mimosine ab-
sorption from LLM diets would be extremely low. This is, indeed, the case.
Thus, D'Mello and Acamovic (1982b) observed that up to 0.92 of ingested
mimosine was excreted by chicks fed on LLM diets. Furthermore, our unpub-
lished studies have demonstrated the total absence of mimosine in blood of
chicks given LLM. It appears unlikely, therefore, that mimosine is the agent
primarily responsible for the adverse effects of LLM in poultry diets. The pos-
sibility should be considered that these deleterious effects arise very largely
from the low availability of nutrients in LLM. It is noteworthy that some amel-
ioration of LLM toxicity in poultry can be achieved by increasing dietary con-
centrations of protein, energy and methionine (Gloria et al., 1966). Mimosine
might play an important role in the toxicity of Leucaena seeds in poultry diets.
There is some evidence that mimosine is absorbed from the seed and deposited
in the tissues of the bird (ter Meulen et al., 1984). However, even in the case
of the seed, it must be recognised that other antinutritional factors such as
protease inhibitors and galactomannan gums may contribute to its toxicity
(Table 5 ). Consequently, until their relative contributions have been resolved,
current estimates of 'acceptable' intakes of mimosine (Szyszka and ter Meu-
len, 1984) should be viewed with caution.
Considerable alleviation of the deleterious effects of LLM may be achieved
by dietary supplementation with ferric sulphate. The mode of action of ferric
sulphate deserves comment in view of the proposition that mimosine may exert
only a minor role in determining the nutritional value of LLM for poultry.
Ferric sulphate may act by reducing dietary pH or by enhancing palatability
and food intake. Although ferric sulphate increases mimosine excretion in birds
given LLM, this effect may be the result of reduced microbial activity in the
alimentary canal rather than a consequence of a complexing interaction be-
tween iron and mimosine (Ross and Springhall, 1963). It is noteworthy that
 25

ferric sulphate supplementation is not accompanied by improvements in pro-

tein utilisation or in the AME content of LLM (our unpublished data). Fur-
thermore, the growth-promoting property of ferric sulphate is contingent upon
appropriate dietary adjustments to accommodate the low AME value of LLM.
Consequently, the scope for any significant role for LLM and, indeed, for other
leaf meals in poultry diets appears to be a diminishing prospect despite recent
advances in detoxification. More attention should, therefore, be given to the
exploitation of novel legume grains such as the winged bean (Psophocarpus
tetragonolobus) and the jack bean (Canavalia ensiformis) as feeding stuffs for
poultry in the tropics (D'Mello et al., 1983, 1985). However, the carotenoid
content of LLM is an important asset in spite of the low availability of the
xanthophyll fractions. Commercial exploitation of LLM for the pigmentation
of egg yolks and broiler carcasses may now be contemplated, provided tissues
and products are monitored for residues of mimosine and its derivatives.

REFERENCES

Abu-Serewa, S., 1976. Effect and source of fat in the hen's diet on the deposition of oxycarotenoids
in egg yolks. Aust. J. Exp. Agric. Anim. Husb., 16: 204-208.
Acamovic, T. and D'Mello, J.P.F., 1981a. Determination of mimosine by ion-exchange chroma-
tography. J.Chromatogr., 206: 416-420.
Aeamovic, T. and D'Mello, J.P.F., 1981b. The effect of iron (III) supplemented Leucaena diets
on the growth of young chicks. Leuc. Res. Rep., 2: 60-61.
Acamovic, T. and D'Mello, J.P.F., 1984. Trypsin inhibition by Leucaena leaf meal, Leucaena seeds
and mimosine. Leuc. Res. Rep., 5: 74-75.
Acamovic, T., D'Mello, J.P.F. and Fraser, K.W., 1982. Determination of mimosine and 3-hydroxy-
4 (1H) -pyridone in Leucaena, avian excreta and serum using reversed-phase high-performance
liquid chromatography. J. Chromatogr., 236: 169-179.
Adeneye, J.A., 1979. A note on the nutrient and mineral composition of Leucaena leucocephala
in Western Nigeria. Anita. Feed Sci. Technol., 4: 221-225.
AreUano, J.A., 1979. Bibliografia sobre Leucaena leucocephala. Instituto Nacional de Investiga-
clones Agricolas, Merida, Yucatan, Mexico.
Armanious, M.W., Britton, W.M. and Fuller, H.L., 1973. Effect of methionine and choline on
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26

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 27

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