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Potential of the use of peanut (Arachis

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Aquaculture Research, 2008, 39, 1299^1306 doi:10.1111/j.1365-2109.2008.01995.x

Potential of the use of peanut (Arachis hypogaea) leaf

meal as a partial replacement for fish meal in diets for
Nile tilapia (Oreochromis niloticus L.)

Mario Gardun
o-Lugo1 & Miguel AŁngel Olvera-Novoa2

1
Centro de Ensen
anza, Investigacio¤n y Extensio¤n en Ganader|¤ a Tropical, Facultad de MedicinaVeterinaria y Zootecnia,
Universidad Nacional Auto¤noma de Me¤xico,Veracruz, Me¤xico
2
Centro de Investigacio¤n y de Estudios Avanzados (CINVESTAV), Unidad Me¤rida,YucataŁn, Me¤xico

Correspondence: M Gardun
o-Lugo, Centro de Ensen
anza, Investigacio¤n y Extensio¤n en Ganader|¤ a Tropical, Facultad de MedicinaVeter-
inaria y Zootecnia, Universidad Nacional Auto¤noma de Me¤xico, Km. 5.5 Carretera Federal Mart|¤ nez de la Torre-Tlapacoyan, AP 136, 93600
Mart|¤ nez de la Torre,Veracruz, Me¤xico. E-mail: tilapia1@prodigy.net.mx

Abstract (Francesco, Parisi, Me¤dale, Lupi, Kaushik & Poli

2004; FAO 2007). Indeed, its average annual growth
The rapid growth of tilapia culture has stimulated
has been 10% compared with 3% in the cattle indus-
the expansion of tilapia feed production and a search
try and1.6% in capture of aquatic species from natur-
for novel protein sources to replace ¢sh meal.Vegeta-
al environments. Aquaculture’s strong growth has
ble or plant sources are promising alternatives and
generated a consequent 30% annual growth in the
legumes are both naturally abundant and high in
production of aquatic species feeds (Francis, Makkar
protein content. A study was carried out to evaluate
& Becker 2001), and has made raw material supply a
the e¡ects of replacing the protein from ¢sh meal
continuous challenge in this industry.
with peanut (Arachis hypogaea) leaf meal (PLM) in
Attaining truly sustainable growth in the aquacul-
diets for male tilapia (Oreochromis niloticus, initial
ture industry will depend on a progressive decrease in
body weight 75.3 g). Four isonitrogenous (35% CP)
the use of marine protein and lipids in feeds for farmed
and isocaloric (18.834 kJ g
1) diets were prepared to
¢sh (Francis et al. 2001; Francesco et al. 2004). Another
include 0% (CON), 10% (PLM10), 20% (PLM20) and
industry goal is to decrease the levels of phosphorus
30% (PLM30) of PLM protein. Average ¢sh weights
and nitrogen in aquaculture farm e¥uents to lower
at the end of the 126-day experiment were not statis-
aquaculture’s environmental impact (Francesco et al.
tically di¡erent among the ¢sh fed CON, PLM10 and
2004;Yamamoto, Sugita & Furuita 2005).
PLM20 diets. The PLM30 diet produced the poorest
Animal- and vegetable-origin protein sources have
growth performance. Organic matter and protein
been tested as alternative feedstu¡s in ¢sh feed pro-
contents of ¢sh were similar in the CON, PLM10 and
duction with varying degrees of success (Ulloa Rojas
PLM20 diets. Carcass chemical composition showed
& Verreth 2003; Kaushik, Cove's, Dutto & Blanc 2004;
a decrease in body fat content as PLM replacement
Li,Wang, Hardy & Gatlin III 2004; Fasakin, Serwata &
levels increased. The high survival ratio in all dietary
Davies 2005). Because of the greater natural abun-
groups (497%) suggests that PLM can be used in
dance of vegetable products, they are of particular in-
O. niloticus feeds for long periods without a¡ecting
terest to researchers as ingredients for ¢sh feed
¢sh growth performance or health.
production (for a complete review, refer to El-Sayed
1999; Ogunji 2004). The use of raw vegetal mater-
Keywords: cichlids, plant protein, ¢sh nutrition,
ials can be limited, however, because of their anti-
legume, ¢sh feeding
nutritional components, which are grouped into
three categories: (1) those a¡ecting protein utiliza-
Introduction
tion and digestion; (2) those a¡ecting mineral utiliza-
Over the last 30 years, aquaculture has grown faster tion; and (3) anti-vitamins and toxic substances
worldwide than any other animal production sector (Francis et al. 2001).

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Journal Compilation r 2008 Blackwell Publishing Ltd 1299
Peanut leaf meal in tilapia feeding M Gardun
o-Lugo & M AŁ Olvera-Novoa Aquaculture Research, 2008, 39, 1299^1306

Among plant-derived raw materials, legumes have Diets

received special attention due to their high protein
Four isonitrogenous and isocaloric diets were formu-
content. This has led to their use as ¢sh meal replace-
lated to provide 35% protein and 18.834 kJ g
1 gross
ments in tilapia feeds at levels as high as 100% in the
energy (El-Sayed & Teshima 1992); energy content
case of soybean meal, or at lower levels when using
was calculated from caloric values (Jauncey & Ross
novel sources such as Leucaena leucocephala leaf meal
1982). The proximate composition of A. hypogaea and
(Jackson, Capper & Matty 1982; Santiago, Aldaba,
¢sh meal was obtained using standard methods
Laron & Reyes 1988), alfalfa meal (Medicago sativa)
(AOAC 1984) (Table 1). The control diet (CON) con-
(Yousif, Alhadhrami & Pessarakli 1994; Ali, Al-Asgah,
tained ¢sh meal (anchovy) as the sole protein source.
Al-Ogaily & Ali 2003) and protein concentrates
The remaining three diets were formulated using
(Olvera-Novoa, Campos, Sabido & Mart|¤ nez-Palacios
three replacement levels of ¢sh meal protein with
1990; Rivas 1993; Borgeson, Racz, Wilkie, White &
PLM protein: 10% (PLM10), 20% (PLM20) and 30%
Drew 2006).
(PLM30; Table 2). The other diet ingredients were
Peanut or groundnut (Arachis hypogaea) is impor-
corn starch, ¢sh oil, edible oil, a vitamin premix
tant in human nutrition due to its high protein and
(Tacon, Sta¡ord & Edwards 1983) and a mineral pre-
energy content. Annual worldwide production of pea-
mix (Tacon,Webster & Mart|¤ nez 1984), as well as car-
nuts in the shell surpasses 33 million tonnes in the 20
boxymethylcellulose (CMC) as a binding agent. These
countries with the highest production (FAO 2006).
ingredients were mixed in a Hobart mixer to form a
In addition to the seed, peanut plants produce high-
dough that was processed into pellets using a meat
protein forage that has long been used as swine and
grinder. The diets were prepared by ¢rst mixing the
ruminant feed, and its £owers are a nectar source
dry meals and premixes for 15 min, adding the oils,
in apiculture. Peanut stem and leaf production (i.e.
starch, some water and CMC and then mixing until
excluding seed production) can be as high as 6 tonnes
a uniform dough was formed that passed throughout
of dry matter per hectare (Rivas 1993).
the meat grinder to form pellets. The feeds were then
To better understand the feed potential of peanut
dried in a forced-air oven at o40 1C for 24 h and
leaf in tilapia culture, the aim of this study was to de-
stored in plastic bags at 5 1C until use. An additional
termine whether peanut (A. hypogaea) leaf meal (PLM)
set of diets was prepared as described earlier to eval-
can partially replace ¢sh meal as a protein source in
uate the digestibility, but including 0.05% chromic
grower feeds for Oreochromis niloticus over long peri-
oxide as a marker (Tables 3 and 4).
ods without toxic or adverse e¡ects on growth perfor-
mance and/or carcass chemical composition.
Experimental design
The growth trial was conducted by placing 15 juve-
Materials and methods nile male O. niloticus (75.3  7.8 g initial weight)
Peanut leaf meal in each of the 15 £oating cages (1.2 m  1.2 m 
0.6 m; 0.9 m3) made of a PVC-coated steel mesh
Leaves were collected from peanut A. hypogaea plants (2.5 cm opening). The cages were placed at equal dis-
in a mixed peanut/corn Zea mays cultivation system at tances from each other in a pond (11.0 m  10.0 m 
Santa Elena, Uxmal, in the state of Yucatan, Me¤xico. 1.2 m) with aeration and water recirculation. The
Collections were carried out when plants exhibited
an average of 10% £owering by cutting the aerial por- Table 1 Proximate composition of peanut Arachis hypogaea
tion of the plant and storing it in plastic bags. Collec- leaf meal and ¢sh meal used in the growth diets for Oreo-
tions were carried out in the morning and, later the chromis niloticus
same day, the collected material was spread out under
a shade covering to partially dry. Forty-eight hours Content (%) Arachis hypogaea meal Fish meal
later, the material was placed in a forced-air oven at
Moisture 4.53 1.06
60  5 1C for 8 h. The leaves were then separated by Crude protein 22.3 67.4
hand from the stems and placed in the oven for an- Lipids 2.09 8.29
other 24 h. After drying, the leaves were ground in a Crude fibre 18.6 0.00
hand mill, then in an electric mill, screened through Ash 9.75 15.0
NFE 42.8 8.22
a 590 mm mesh (Rivas 1993) and stored in a plastic
NFE 5100
(% moisture1% crude protein1% lipids1% ash).
container at room temperature until use.

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1300 Journal Compilation r 2008 Blackwell Publishing Ltd, Aquaculture Research, 39, 1299^1306
Aquaculture Research, 2008, 39, 1299^1306 Peanut leaf meal in tilapia feeding M Gardun
o-Lugo & M AŁ Olvera-Novoa

Table 2 Formulation and proximate composition of diets Table 4 Formulation and proximate chemical composition
containing increasing levels of peanut Arachis hypogaea leaf of diets for Oreochromis niloticus used in the apparent digest-
meal protein in substitution of ¢sh meal protein ibility study

Diet Diet

CON PLM10 PLM20 PLM30 CON PLM10 PLM20 PLM30

Ingredient (%) Ingredient (%)

Fish meal 51.9 46.7 41.5 36.3 Fish meal 52.4 47.2 42.0 36.7
A. hypogaea leaf meal 0.00 15.7 31.4 47.1 A. hypogaea leaf meal 0.00 15.2 30.3 45.5
Fish oil 0.70 1.13 1.56 1.99 Fish oil 0.00 1.15 0.69 1.23
Soy oil 5.00 4.67 4.34 4.02 Soy oil 4.61 4.72 4.44 4.16
Corn starch 36.4 25.8 15.2 4.60 Corn starch 38.4 28.2 18.0 7.88
Mineral premix 1.50 1.50 1.50 1.50 Mineral premix 1.50 1.50 1.50 1.50
Carboxymethyl cellulose 1.50 1.50 1.50 1.50 Vitamin premix 3.00 3.00 3.00 3.00
Nutrient content (% as fed) Chrome oxide 0.05 0.05 0.05 0.05
Moisture 4.88 6.77 3.24 3.70 Nutrient content (% as fed)
Crude protein 38.3 37.0 37.9 38.9 Moisture 5.90 6.64 6.34 6.64
Crude fat 10.2 10.4 10.9 11.7 Crude protein 36.7 36.7 37.7 36.8
Crude fibre 1.19 3.63 5.81 7.16 Crude fat 6.04 7.52 10.9 10.4
Ash 11.1 11.3 12.2 12.7 Crude fibre 1.28 3.71 6.40 9.97
NFE 34.3 30.9 30.0 25.8 Ash 12.0 12.4 12.8 13.0
Gross energy (kJ g
1) 18.6 17.8 18.0 17.8 NFE 38.1 33.0 25.9 23.2
Gross energy (kJ g
1) 18.6 17.8 18.0 17.9
NFE 5100
(% moisture1% crude protein1% lipids1% ash).
NFE 5100
(% moisture1% crude protein1% lipids1% ash).

Table 3 Proximate composition of peanut Arachis hypo- remained within the ranges suitable for tilapia:
gaea leaf meal and ¢sh meal used in diets for the apparent O2 (mg L
1) 07:00 hours, 4.1  0.74; O2 15:00 hours,
digestibility analysis 5.7  0.6; water temperature ( 1C) 29.1  1.7; total
hardness (mg L
1) 106  20.0; phosphorus P2O4
Content (%) A. hypogaea meal Fish meal
(mg L
1) 1  0.41; ammonium NH4 (mg L
1)
Moisture 5.22 5.36 0.2  0.08; nitrite NO2 (mg L
1) 0.1  0.29; nitrate
Crude protein 23.1 66.7 NO3 (mg L
1) 0.5  0.36; transparency (cm)
Lipids 1.85 10.3 29.4  9.30; suspended solids (mg L
1) 17.1  20.1;
Crude fibre 17.4 0.00
and turbidity (NTU) 33.2  20.8.
Ash 9.62 17.4
NFE 42.8 0.23
The apparent digestibility trial was performed
at CINVESTAVusing a closed water-recirculating sys-
NFE 5100
(% moisture1% crude protein1% lipids1% ash).
tem consisting of eight aquaria (60 cm  40 cm 
60 cm; 72 L volume) with sloped bottoms and an ac-
experimental diets were randomly assigned to the cessory for continuous faeces collection. This system
cages by triplicate. Fish were fed to apparent satiation also included a biological ¢lter and a 2000 W
three times daily, 7 days a week. An additional cage titanium heater to maintain the temperature (28 1C).
with the same dimensions was placed in the same Five juvenile male O. niloticus (74.5  9.2 g) from
pond and stocked with individually marked tilapia CINVESTAV stocks were randomly placed in each
from the same lot (71.0  6.5 g initial weight). These aquarium. Fish were fed the experimental diets for
¢sh were not fed during the experimental period to 15 days, till enough faeces was collected to run the
determine the in£uence of natural food availability digestibility analysis. Faeces collection began on
on weight gain. day 5, after four days of adaptation to the feed and
During the 126-day experimental period, the handling.
¢sh were weighed every 2 weeks. Water quality
was checked periodically to determine when water
changes were needed to ensure the proper water
Analytical methods
quality for normal growth (Boyd 1990; Gardun
o-
Lugo, Granados AŁlvarez, Olvera-Novoa & Mun
oz- Standard methods (AOAC 1984) were used to deter-
Co¤rdova 2003). All water quality parameters mine moisture content, crude protein (N  6.25),

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Journal Compilation r 2008 Blackwell Publishing Ltd, Aquaculture Research, 39, 1299^1306 1301
Peanut leaf meal in tilapia feeding M Gardun
o-Lugo & M AŁ Olvera-Novoa
lipids, crude ¢bre and ash for the protein meals,
experimental diets and ¢sh (before and after the
growth trial). Chromic oxide content was evaluated
by digestion with nitric and perchloric acids (Furuka-
wa & Tsukahara 1966). Gross energy was calculated
using the kilojoules values of 23.03 for protein
content, 38.10 for lipids and 17.16 for carbohydrate
(Jauncey & Ross 1982; New 1987).
Statistical methods
Growth performance, feed e⁄ciency and the feed and
protein conversion ratios were compared using an
analysis of variance (ANOVA) for a completely random
design with four treatments and three replicates
(P 5 0.05). Percentage and ratio values were Arcsin
transformed before statistical analysis. Post hoc com-
parison of means was performed using theTukey test
(Steel & Torrie 1986) using the SAS statistical package
(1985). The same design and signi¢cance level were
used for the digestibility trial, but with only two repli-
cates. Results for the ¢sh in the treatment with no
feed (without feed group, WFG) were not included in
the statistical analysis due to the clear disadvantage
of these ¢sh in comparison with those in the other
treatments as can be seen in Fig. 1.
Results
Survival, growth performance and feed e⁄ciency
results are shown in Table 5. Survival was high,
with no di¡erences among the treatments (P40.05),
300
g among the CON, PLM10 and PLM20 diets, but all
CON PLM10
250
PLM20 PLM30
WFG
200
150
100
50
0
1 2 3 4 5 6 7 8
Biweekly weights
Figure 1 Changes in body weight of Oreochromis niloti-
cus fed with three diets including Arachis hypogaea leaf
meal: PLM10, PLM20 and PLM30; one control with sole
protein from ¢sh (CON) and one unfed (WFG). PLM, pea-
nut leaf meal; PLM10, PLM protein:10%; PLM20, PLM pro-
tein: 20%; PLM30, PLM protein: 30%; WFG, without feed
group; CON, control diet.
1302 Journal Compilation r 2008 Blackwell Publishing Ltd, Aquaculture Research, 39, 1299^1306
Aquaculture Research, 2008, 39, 1299^1306
indicating that peanut leaf contains no toxic sub-
stances that might a¡ect the animals negatively. The
highest ¢nal weight occurred in conjunction with
the CON diet, although these were not signi¢cantly
di¡erent from the PLM10 and PLM20 diets, which
did not di¡er (P40.05) from each other. Fish fed the
PLM30 diet had ¢nal weights that were signi¢cantly
lower than those for ¢sh fed the CON diet, but not dif-
ferent from those fed the PLM10 and PLM20 diets. A
similar pattern was observed in the per cent weight
gain (PWG). Speci¢c growth rate (SGR), however,
was signi¢cantly higher (Po0.05) in the ¢sh fed the
CON diet than those fed the PLM diets, which had sig-
ni¢cantly lower values as PLM percentage increased.
The feed and protein utilization e⁄ciency results
showed a tendency towards a lower feed intake that
was likely due to a reduction in palatability as the PLM
replacement level in the diets increased. Body weight
also decreased as PLM inclusion increased, with ¢sh
on the CON diet having a higher (although not statisti-
cally di¡erent) body weight than the ¢sh on the PLM10
and PLM20 diets, and had a signi¢cantly higher ¢nal
weight than those on the PLM30 diet. The feed conver-
sion ratio (FCR) was signi¢cantly lower for the CON
treatment than the PLM treatments, and the FCR for
the PLM30 treatment was signi¢cantly higher than all
the other treatments. Carcass nitrogen deposition
(CND) and apparent nitrogen utilization (ANU) fol-
lowed the same pattern. The protein e⁄ciency ratio
(PER) was not signi¢cantly di¡erent between the CON
and PLM10 diets, but PER for the PLM30 diet was
signi¢cantly (Po0.05) lower than the others.
No signi¢cant di¡erences occurred in AOMD
three were signi¢cantly higher (Po0.05) than that
for the PLM30 diet (Table 5). This pattern also held
true for apparent protein digestibility (APD).
Fish body composition showed no link between
body moisture content and diet composition (Table 6);
the most signi¢cant di¡erence was observed between
the CON and PLM30 treatments. Crude protein levels
were not signi¢cantly di¡erent among the treat-
ments. Body crude lipid values, in contrast, exhibited
an inverse relationship with the PLM inclusion level,
with the highest content in ¢sh fed the CON diet,
followed by those fed the PLM10 and PLM20 diets
and the lowest in those fed the PLM30 diet. Ash con-
tent was similar among the PLM20, CON and PLM30
treatments (in descending order of content) and
the lowest in the PLM10 treatment, although the lat-
ter was only statistically di¡erent from the PLM20
treatment.
r 2008 The Authors
Aquaculture Research, 2008, 39, 1299^1306 Peanut leaf meal in tilapia feeding M Gardun
o-Lugo & M AŁ Olvera-Novoa

Table 5 Growth performance and feed utilization e⁄ciency in male Oreochromis niloticus fed diets containing peanut,
Arachis hypogaea, leaf meal (PLM)

Mean values CON PLM10 PLM20 PLM30 SE

Survival (%) 100a 100a 97.6a 100a 2.06

Initial weight (g) 73.8a 76.4a 74.6a 73.3a 4.11
Final weight (g) 282.3a 235.1ab 220.6ab 176.8b 28.2
Weight gain (%)w 282.4a 205.9b 195.1b 140.6b 25.3
Specific growth rate (% day
1)z 1.06a 0.88b 0.86bc 0.70c 0.06
Individual feed intake (g day
1)‰ 3.50a 3.12ab 2.98ab 2.60b 0.29
Feed conversion ratioz 2.13c 2.52b 2.58b 3.18a 0.18
Protein efficiency ratiok 1.23a 1.08ab 1.03b 0.81c 0.72
Carcass nitrogen deposition (mg day
1) 49.7a 32.9b 34.7b 21.3c 0.08
Apparent nitrogen utilization (%)ww 22.7a 17.9b 18.8b 14.6c 1.11
Apparent organic matter digestibility (%) 60.3a 59.3a 52.9a 17.6b 8.12
Apparent protein digestibility (%) 75.6ab 80.3a 83.0a 71.6b 2.91

Di¡erent superscript letters in the same row indicate signi¢cantly statistical di¡erences (Po0.05).
wPWG (%) 5 100[(¢nal weight
initial weight)/initial weight].
zSGR (% day
1) 5 100(ln ¢nal weight
ln initial weight)/time (days).
‰IFI (g day
1) 5 (s biweekly feed intake)/time (days).
zFCR 5 IFI/DWG.
kPER 5 DWG/protein intake.
CND (g day
1) 5 1000[(protein percentage of ¢nal weight)
(protein percentage of initial weight)]/100/time (days)/6.25.
wwANU (%) 5 100(NDB/N intake)
SE, standard error of the mean.

Table 6 Carcass proximate composition of Oreochromis protein without a signi¢cant e¡ect on the growth
niloticus fed diets with Arachis hypogaea leaf meal replacing performance. This di¡ers from the results of Yousif
¢sh meal et al. (1994), who reported a decline in ¢sh growth
with the replacement of ¢sh meal with sun-dried
Diet
alfalfa (M. sativa) leaf meal at replacement levels as
Average values (%) Initial CON PLM10 PLM20 PLM30 SE low as 5% over 129 days. They attributed this to anti-
c
nutritional factors such as trypsin inhibitors in the
Moisture 80.3 70.8 74.0ab 71.9bc 76.7a 0.97
Crude protein 13.1 17.3a 15.3a 16.9a 14.9a 0.60
alfalfa meal. On testing alfalfa leaf protein concen-
Crude fat 2.32 5.40a 4.70b 4.30b 2.30c 0.18 trates for 9 weeks, however, Olvera-Novoa et al.
Ash 6.19 5.40ab 4.30b 5.60a 5.00ab 0.28 (1990) reported no mortality in Oreochromis mossam-
Di¡erent superscript letters in the same row indicate statistical bicus (Peters), indicating that the protein concentrate
di¡erence (Po0.05). process eliminates or reduces the anti-nutritional or
SE, standard error of the mean. toxic substances in alfalfa. In this study, ¢sh meal
could be substituted with up to 35% cytoplasmic pro-
tein concentrate, although obtaining the concentrate
The ¢sh in the cages without feed had an average was complex and extremely expensive, far more so
¢nal weight of 62.8  7.1g, implying that they lost an than the process used in the present study to dry the
average of 8.2 g compared with their 71.0  6.5 g peanut leaves.
mean initial weight. Four of the 15 originally stocked The present results coincide with those of Fasakin,
¢sh died during the trial, likely due to handling dur- Balogun and Fasuru (1999), who reported no signi¢-
ing the weighing. The results for that treatment were cant e¡ect on Nile tilapia growth or feed utilization
not included in the statistical analysis because of with up to 20% replacement of sun-dried water lentil
their clear disadvantage in growth performance. (Spirodela polyrrhiza) meal for ¢sh meal. The authors
further supported their results with economic ana-
lyses indicating that replacement up to 30% would
Discussion
not a¡ect the economics of the culture operation. Un-
The results indicate that peanut (A. hypogaea) leaf fortunately, economic analyses could not be carried
meal can replace ¢sh meal at levels up to 20% crude out in the present study to support the replacement

r 2008 The Authors

Journal Compilation r 2008 Blackwell Publishing Ltd, Aquaculture Research, 39, 1299^1306 1303
Peanut leaf meal in tilapia feeding M Gardun
o-Lugo & M AŁ Olvera-Novoa
of ¢sh meal with PLM at the recommended levels
because the cost of peanut leaf processing on a
commercial scale is not known.
The high survival in the present study indicates
that PLM can be used in tilapia diets for long periods.
This contrasts with Wee and Wang (1987), who re-
ported the appearance of ocular cataracts in O. niloti-
cus fed diets with 25% or 50% Leucaena leaf meal
after 4^10 weeks of feeding.
In another study to replace the ¢sh meal with Leu-
caena leaf meal in O. niloticus diets, ocular cataracts
occurred between the 12th week and the end of the
trial (24 weeks) and the ¢sh lost 10% of their initial
weight due to the presence of anti-nutritional factors
like mimosine (Santiago et al. 1988). Mart|¤ nez-
Palacios, GalvaŁn-Cruz, Olvera-Novoa and ChaŁvez-
Mart|¤ nez (1988) also reported high mortality in
O. niloticus ¢ngerlings fed diets containing Jack bean
(Canavalia ensifomis) seed meal associated with the
non-thermolabile toxic compound L -canavaline.
Use of moringa (Moringa leifera) meal in tilapia
diets can cause adverse e¡ects, but these can be re-
duced or eliminated by extracting the leaf with
methanol for 24 h to remove the anti-nutritional fac-
tors (Afuang, Siddhuraju & Becker 2003). In that
study, the anti-nutritional factors a¡ected the meta-
bolism and reduced feed intake because feed palat-
ability was reduced when untreated moringa meal
was used. Palatability was also adversely a¡ected in
the present study as the PLM content increased.
Heat treatment of peanut meal at 60 1C during de-
hydration e¡ectively inactivated its trypsin inhibitor
content (Rivas 1993), as well as any other thermola-
bile substances that might a¡ect nutrient utilization
or might be toxic (Francis et al. 2001), and is much
faster and cheaper than other methods (e.g. metha-
nol extraction). Again, the high survival observed in
this study supports these conclusions and shows that
PLM can be used in tilapia feed for long periods with-
out causing any apparent pathologies that could
a¡ect ¢sh health, even at the 30% replacement level.
The present study showed that tilapia has a high
capacity to digest the protein from the PLM, with
values comparable to those for ¢sh meal. The APD
increased 8% in the PLM10 and PLM20 diets com-
pared to the CON diet, but even the PLM30 diet had
an APD comparable with the CON diet. This has also
been the case in other studies, indicating that tilapia
can e⁄ciently use that feedstu¡ as a protein source.
Mart|¤ nez-Palacios et al. (1988) also reported high
APD when using C. ensiformis meal as a substitute
for ¢sh meal in tilapia diets; the same result was
1304 Journal Compilation r 2008 Blackwell Publishing Ltd, Aquaculture Research, 39, 1299^1306
Aquaculture Research, 2008, 39, 1299^1306
obtained by Olvera, Mart|¤ nez, GalvaŁn and Chavez
(1988) when Sesbania grandi£ora seed meal was in-
cluded in tilapia diets, but only at the 10% replace-
ment level. These results can be explained by the
high capacity of tilapia to digest protein from plant
dietary sources even when the digestibility for the or-
ganic matter was low, as was the case in this study.
This ability is attributed to the very low stomach pH
observed in tilapia, which allows them to digest and
extract the cellular content even without breaking
the cellulose cell wall (Balarin & Hatton 1979; Bowen
1981; Getachew 1987; Ekpo & Bender 1989; Horn &
Messer 1992).
Organic matter digestibility can be a¡ected by
intestinal motility, and higher ¢bre content in tilapia
diets decreases feed residence time in the gastroin-
testinal tract (Anderson, Jackson, Matty & Caper
1984). This was clear in the present study, where
higher PLM replacement levels resulted in higher
faeces production that directly a¡ected the feed di-
gestibility, as can be seen in diet PLM30, where the
digestibility in ¢sh was 17.6%.
The lower weight gain observed with the higher
PLM replacement levels may be due to amino acid de-
¢ciencies (especially lysine, cysteine methionine, iso-
leucine and threonine) (Rivas 1993), as the diet
pro¢les were not those recommended for O. niloticus
(NRC 1993; Jauncey 2000). Future research on A. hy-
pogaea as a ¢sh meal substitute could complement
these de¢ciencies by including other alternate pro-
tein sources, or by adding the de¢cient amino acids,
although use of the latter in tilapia diets is apparently
contradictory, given the organism’s digestive physiol-
ogy (Teshima & Kanazawua 1988). For instance, cow-
pea (Vigna unguiculata) is a promising source of
isoleucine, treonine and lysine (Olvera-Novoa, Per-
eira-Pacheco, Olivera-Castillo, Pe¤rez-Flores, Navarro
& SaŁmano 1997) that can be mixed with peanut meal
to balance the amino acid pro¢le.
Body composition changed as PLM replacement
levels increased. The moisture content increased with
the PLM30 diet, but the water content was not higher
than those observed in the ¢sh at the beginning of the
study. The ¢nal protein, lipid and ash contents were
similar to the levels in previous reports (Anderson
et al. 1984; Wee & Wang 1987). These same authors
stated that protein, fat and lipid levels declined as
plant protein replacement levels increased, mainly
due to lower digestibility and consequent lower nutri-
ent availability in diets with high plant protein levels.
It is unlikely that intake of natural food had any
signi¢cant e¡ect on growth, given the weight loss in
r 2008 The Authors
Aquaculture Research, 2008, 39, 1299^1306 Peanut leaf meal in tilapia feeding M Gardun
o-Lugo & M AŁ Olvera-Novoa
the ¢sh that were not fed, although it may have had a
positive e¡ect on the organism’s health (Santiago et al.
1988).
In conclusion, PLM can be used as a protein source
as a replacement for ¢sh meal in tilapia feeds without
signi¢cant e¡ects on ¢sh performance. Up to 20% of
this feedstu¡ had no signi¢cant negative e¡ects on
growth, but an economical evaluation should be un-
dertaken to de¢ne the impact of this substitution on
production costs.
Acknowledgments
The authors thank the Faculty of Veterinary Medi-
cine and Zootechny, Universidad Auto¤noma de
YucataŁn, for ¢nancially supporting this study; the
Faculty of Veterinary Medicine and Zootechny,
Universidad Nacional Auto¤noma de Me¤xico, for ¢-
nancial support and access to its installations and
laboratories; and the Centro de Investigacio¤n y de
Estudios Avanzados, Unidad Me¤rida, for access to
its laboratories for the digestibility analysis and for
providing tilapias. Special thanks are due to Leticia
Olivera Castillo and Wilbert Che Leo¤n for their
invaluable laboratory assistance, and the peanut
growers of Santa Elena, Uxmal, Yucatan, for their
help in donating and collecting raw material.
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