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Bio-processing of agro-byproducts to animal feed

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 Critical Reviews in Biotechnology, 2012; Early Online: 1–19
© 2012 Informa Healthcare USA, Inc.
ISSN 0738-8551 print/ISSN 1549-7801 online
DOI: 10.3109/07388551.2012.659172

Review article

Bio-processing of agro-byproducts to animal feed

C.M. Ajila1, S.K. Brar1, M. Verma2, R.D. Tyagi1, S. Godbout2, and J.R. Valéro1
1
INRS-ETE, Université du Québec, 490, Rue de la Couronne, Québec, Canada G1K 9A9 and 2Institut de recherche et de
développement en agroenvironnement inc. (IRDA), 2700 rue Einstein, Québec (Québec), Canada G1P 3W8
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Abstract
Agricultural and food-industry residues constitute a major proportion (almost 30%) of worldwide agricultural
production. These wastes mainly comprise lignocellulosic materials, fruit and vegetable wastes, sugar-industry wastes
as well as animal and fisheries refuse and byproducts. Agro-residues are rich in many bioactive and nutraceutical
compounds, such as polyphenolics, carotenoids and dietary fiber among others. Agro residues are a major valuable
biomass and present potential solutions to problems of animal nutrition and the worldwide supply of protein
and calories, if appropriate technologies can be used for their valorization by nutrient enrichment. Technologies
available for protein enrichment of these wastes include solid substrate fermentation, ensiling, and high solid or
slurry processes. Technologies to be developed for the reprocessing of these wastes need to take account of the
peculiarities of individual wastes and the environment in which they are generated, reprocessed, and used. In
particular, such technologies need to deliver products that are safe, not just for animal feed use, but also from the
perspective of human feeding. This review focuses on the major current applications of solid-state fermentation in
For personal use only.

relation to the feed sector.

Keywords:  Animal feed, solid-state fermentation, agro-residues, nutrient enrichment

Introduction feeding. Therefore, modern science must emphasize for

Advances in technology and science have led to an
improved standard of living. This has resulted in an
increased demand for food supplies in traditional as well
as modern ways. Traditional ways include the utilization
of land and sea to yield more food with the help of high-
yielding crop varieties as well as the use of fertilization
and better irrigation. In addition to these traditional
ways, alertness to the problem of waste disposal and
its utilization has been regularly proposed (El Boushy,
1990). Wastes are mainly generated by agricultural and
municipal segments of population. The utilization of
wastes need urgent attention as the recycling and reduc-
tion of wastes can minimize environmental pollution
and ameliorate the present situation by creating new
products from wastes, such as animal feed. The increas-
ing costs and pressure concerned with waste disposal
stress the need for a reappraisal of the utilization of
wastes either directly (as diet ingredient) or indirectly
(nutrient enrichment by microorganisms) for livestock
research on the management of wastes, its processing
technology, and subsequent feeding value for livestock.
The rapid changes in animal farming imply scrutiny
in the studies on nutrition evaluation with respect to
the target animals and low-cost technology approach.
Studies should involve nutritional evaluation on digest-
ibility, feeding value, bio-hazards and feasibility for waste
management and utilization. Even though the potential
of agro-byproducts are very obvious, the utilization of
feed from the waste in diet formulation until now has
been negligible due to the constraints imposed by several
nutritional and technical considerations.
An improvement in animal feeding is one of the
important and basic conditions for the better manage-
ment of farming of animals. It was recognized that
poor quality of the feed is mainly responsible for the
poor animal performance. Most of the animal feed is
quite expensive for most farming practices. Therefore,
adequate and good-quality feed supplies are the most

Address for Correspondence: Satinder Kaur Brar, Institut national de la recherche scientifique, Centre Eau, Terre & Environnement/Centre
for Water, Earth and Environment, 490 de la Couronne, Québec (QC), Canada G1K 9A9. E-mail: satinder.brar@ete.inrs.ca
(Received 01 May 2011; revised 12 December 2011; accepted 17 January 2012)

1
 2  C. M. Ajila et al.
Horticulture
Fodders &
Forage
crops
Crop residue
Grains/ oil
seeds
Animal feed Digestive
enzymes
By-Products
Other feed Antibiotics
additives
Figure 1. General animal feed system practiced in live-stock farms.
Critical Reviews in Biotechnology Downloaded from informahealthcare.com by 206.162.180.2 on 03/01/12
important factors in farm management. The approach
has been part of traditional agriculture for centuries.
The production of animal feed is one of the most logical
ways of utilizing a substantial portion of the enormous
potential agro-residues. More than 70% of the expendi-
ture in farming is mainly for animal feed. Figure 1 shows
the general animal feed system practiced in livestock
farming system. The production of animal feed from
agro-residues represents one of the highest cash returns
due to the fact that the demand for animal feed is always
For personal use only.
stable and huge. The marketing is also relatively easy
and the technologies involved are not too complicated.
The technologies involved for the production of animal
feed can be handled in small-scale industries also. Thus,
the production of animal feed from agro-residues will
be one of the most sustainable technologies to find an
effective way to attain income by the agro community
and also for the better management of environmental
pollution.
Agro-wastes are of various types and can be classified
into different categories, such as crop waste and residues,
byproducts from fruit- and vegetable-processing indus-
try, sugar, starch and confectionary industry byproducts,
byproducts from grain- and legume-milling industry
and oil industry, and byproducts from distilleries and
breweries. The handling and technologies used for the
processing of agro-wastes are mainly based on their type.
This review discusses and summarizes some aspects of
animal-feed management, by employing different strate-
gies and bioprocessing of agro-products for the produc-
tion of animal feed in a cost-effective way, which in turn
leads to sustainable eco-management.
Nonconventional food resource as an
animal feed
Increasing the utilization of agro-industrial byproducts
and nonconventional feed resources is one of the most
challenging tasks concerning components of animal
industries and it is justified by serious feed deficits,
continuing low animal productivity, and inability of the
 
components of the animal industries to meet national
targets, especially that of animal proteins.
Nonconventional food sources can be defined as
those feeds that have not been traditionally used in ani-
mal feeding and are not normally used in commercially
prepared rations for livestock. The nonconventional
food resources mainly consist of variety of feeds from
perennial crops. This includes feed stuff, such as palm
press fiber (oil palm byproducts), single-cell proteins,
feed materials derived from agro-industrial byproducts
of plant origin, poor-quality cellulosic roughages from
farm residues, such as slaughterhouse byproducts and
byproducts from processing of sugar, cereal grains, cit-
rus fruits and vegetables from the processing of food for
human consumption (Devendra, 1987).
The major characteristics of nonconventional food
sources are as follows:
1. They are mainly organic and can be in a solid, slurry,
or liquid form.
2. Their economic value is often less than the cost of
their collection and transformation for use, and con-
sequently, they are discharged as wastes.
3. Nonconventional food sources are generally excel-
lent sources of fermentable carbohydrates e.g. cas-
sava and sweet potato.
4. Fruit byproducts have sugars which are energetically
very beneficial.
5. Most of the nonconventional food sources are bulky,
poor-quality cellulosic roughage with a high crude
fiber (CF) and low nitrogen contents, suitable for
feeding to ruminants.
6. Some of these feeds have deleterious and toxic effects
on animals and there is not enough information
about the nature of the active principles and ways to
detoxify these effects.
7. They have considerable potential as feed materials
by the modern technological applications.
8. Further information is needed on chemical composi-
tion, nutritive value, toxic factors, and value in feed-
ing systems.
Many of the nonconventional food resources are con-
sidered as wastes since they have no further economic
value. These wastes can be utilized and converted by
animals into valuable products for human benefit as new
feed resources and in turn increase the availability of
limited feed resources. Agricultural and agro-industrial
activities produce thousands of tonnes of byproducts, e.g.
sugarcane bagasse, citrus bagasse, fruit peel, corn straw,
and corncobs. Nutritionally, the agro-industry produces
two groups of residues: fibrous residues (high and low
digestibility) and bran. Fibrous residues can be divided
into parts with high digestibility, e.g. citrus pulp, corn
gluten bran, soy husk, and brewing residues (barley),
and those with low digestibility, e.g. sugarcane bagasse,
cereal, corn, soy and cotton straw, cotton husk, soy husk,
peanut husk, and harvest remnants of forage grass seeds.
 Bio-processing of agro-byproducts to animal feed  3

Table 1. Utilization of agro-industrial and nonconventional feed resources as animal feed.

Feed source Characteristics Animals
1. Energy and protein concentrates (e.g. rice bran, coconut cake, High energy and high protein Poultry, pigs, ducks, cattle
soybean meal, poultry litter)
2. Good-quality crop residues (cassava peels, cereal-grain waste, High protein and high energy Pigs, ducks, cattle
animal waste etc)
3.  Medium-quality crop residues Medium proteins Pigs, ruminants
4.  Low-quality crop residues Low protein and high fiber Ruminants
Devendra (1997).

The nonconventional and agro-industrial byproducts Table 2. Percentage of fruit and vegetable processing byproducts.
can be divided into three categories (Table 1): Fruits / vegetables Nature of wastes Amount of waste (%)
Apple Peel, pomace, seed 25
1. Energy-rich feeds: e.g. bananas, citrus fruits, pine- Mango Peel, stones, 45
apple, sugarcane, and root crop wastes, Citrus Peel, rag, seed 50
2. Protein supplements: e.g. oilseed cakes and meals,
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Tomato Skin, core, seed 20

3. Byproducts from cereal milling and palm-oil refining. Pineapple Skin, core 33
Grape Stem, skin, seed 20
Guava Peel, core, seed 10
Quantitative and qualitative value of
Potato Peel 15
agro-byproducts as animal feed Onion Outer leaves 10
Many byproducts of agro-industry can be fed to animals Banana Peel 35
directly without any modification or can be used after Pea Shell 40
fermentation of the agro-residues. Generally, byproducts
which can be used as feedstuff should be economical, rich barley, wheat, rice, beans, oats, rye stalks, or stovers
in nutrients, and free of toxins or other substances that from corn, sorghum, cotton among others. Fruit and
may be unhealthy to animals. Recovering byproducts for crop residues include apricots, almonds, apples, avoca-
For personal use only.

use as animal feed can help the agro-industry save money
by reducing waste discharges, cut waste-management
costs, and also prevent environmental pollution. Most
of the agro-residues offer an inexpensive source of nutri-
ents as compared to traditional feeds and also support
acceptable animal performance. The increase in global
human population over the last decades has influenced
the demand for food products of animal origin. In turn,
this requires a considerable development of animal pro-
duction. The feed situation in Europe has demonstrated
that to maintain the population of animals at the present
level, the protein deficit is of the order of 25–28 million
tonnes and varies from several hundred thousands to
million tonnes in various countries (Szebiotko, 1984).
Denmark imports 30% of the total amount of utilized
protein to cover the country’s protein demand even with
its developed agriculture and high animal production.
About two-third of the imported feedstuff consists of
oilseed meals and animal-origin meals; the rest is grain
and cereal products (Szebiotko, 1984). Food industry
byproducts and crop residues are used as components
of commercial feed mixtures or added to feeds prepared
directly on the farms.
Agro-byproducts refer to byproducts derived in the
industry due to processing of main products in large
quantity. They are less fibrous, more concentrated,
highly nutritious, and less expensive as compared to
crop residues (Aguilera, 1989). The crop-waste residues
mainly consist of materials remaining after harvesting
and processing, such as straw, husk or stubble from
dos, grapes, lemons, limes, olives, oranges, and plums
among others. The vegetable-crop residue consists
mainly of vine, peel, and leaves which remain after har-
vesting and processing. The major source of the waste
in the fruit- and vegetable-processing industry are
illustrated in Table 2. Pomace is the major byproduct
of apple-processing industry which consists of crushed
flesh, stalks, peels, seeds, etc (Rahmat et al., 1995). Peel
(15–20%), coarse fibrous pulpy waste (5–10%), and ker-
nel (15–20%) are the major byproducts from the mango-
processing industry (Ajila et al., 2007). The wastes from
starch industry, such as tapioca produce tapioca rind or
peel and spent pulp. Rice husk is a byproduct obtained
during rice milling.
Food and agro-industry byproducts and agricultural
wastes used as animal feed can be divided into two main
groups:
1. Plant origin
2. Fermentation industry.
Byproducts of plant origin
Plant origin consists of milling industry byproducts, such
as bran, waste flour, wastes resulting from grain-cleaning
processes, wheat, corn and rye germs, hulls of some
seeds, e.g. pea, barley, and buckwheat. The byproducts
of the oil industry, such as solvent-extracted cakes from
soybean and oil-yielding rape, sunflower, flax, and prod-
ucts formed during refining of plant oils, lecithin and
fatty acids can also be used for the production of animal

© 2012 Informa Healthcare USA, Inc.  

 4  C. M. Ajila et al.
feed. Byproducts of sugar and starch industry, such as
beet pulp, molasses, deface-saturation residues, potato
pulp, potato cell juice, and other residues of seeds after
starch extraction, gluten gums are rich sources of protein
and carbohydrates. Byproducts of the fruit and vegetable
industry include products resulting from peeling veg-
etables, pomace, and fruit seeds, among others.
Plant byproducts, such as husk and pods of common
pulses with leaves and tender stems can be used as nutri-
tious cattle feed and are also good sources of digestible
protein (Ranjhan, 1993). They can be used for livestock
feeding along with the concentrate mixture (Bhatti &
Khan, 1996 ). It was reported that urea treatment of straws
significantly increased their nutritive value and digestibil-
ity (Wanapat et al., 2009; Williams et al., 1984). Similarly,
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Mahmood (1988) found that dry-matter digestibility was
maximum with 4% urea. The legume byproducts with
wheat bran were used as animal feed to meet the entire
nutritional requirements of lactating cows producing up
to 5 L of milk per day (Ranjhan, 1993). The major limiting
factor in legume byproducts is the higher level of phos-
phorus and high CF (38%), which may inhibit the feed
intake. This inhibition of feed intake can be overcome
by sprinkling a mixture of molasses and diammonium
phosphate or preferably a balanced liquid supplement
on chopped groundnut straw (Maglad et al., 1986). Wheat
bran, which is a major component in formulating feed for
For personal use only.
dairy animals, contains about 10% DCP (digestible crude
protein), 65% TDN (total digestible nutrients), 0.07% cal-
cium (Ca) and 0.35% phosphorus(P), can be fed to sick
animals without any ill effects, and produces a laxative
effect in the intestine (Verma, 1997). Bran has an amino-
acid balance superior to that of whole wheat, is high in P
but low in Ca, and is also a good source of water-soluble
vitamins, except niacin (Cheeke, 1991). Wheat middlings
are similar to bran but they are lower in fiber and higher
in flour contents, so they are higher in digestible energy
than bran. Wheat middlings contain 10–14% crude pro-
tein (CP) and 9.5% CF (Cheeke, 1991). Rice husks contain
8–11% water, 15.6–22.6% ash, 14.5–17.5% acid insoluble
ash, 2.9–3.6% CP, 39–42% CF, and 25–29% nitrogen-free
extract. Feeding of rice bran alone may result in colic pain
due to the formation of ball inside the intestine. Hence,
rice bran should always be mixed with other components
(Ranjhan, 1993). Rice bran contains 7% DCP, 65% TDN,
0.06% Ca and 1.12% P, is rich in vitamin B-complex, and it
can be used for feeding cattle, buffaloes, sheep, and goat
(Verma, 1997). Rice bran is a major feed in tropical coun-
tries and has high oil content (13%) (Cheeke, 1991). Rice
polish contains about 3% fiber, 12% fat, 12–14% CP, and
vitamin B-complex. Maize gluten feed is a rich source of
protein (45–48%) and is used in livestock feeding (Verma,
1997).
Tomato pomace contains 14.3% DCP and 65% TDN,a
and the dried and ground tomato pomace can be used
as a part of animal feed up to 35% level (Verma, 1997).
Citrus pulp contains a high amount of soluble sugars
with a digestibility of 85%, contains about 6.0% CP, and
 
can replace one-third of animal feed mixture without ill
effects on the health and utilization of nutrients in buf-
faloes (Ranjhan, 1993; Verma, 1997). Pineapple waste
together with maize grain, wheat bran, and molasses has
successfully been fed to draught animals as a source of
energy (Verma, 1997). Ranjhan (1993) conducted a study
in Philippines on 5000 feedlot beef cattle raised on fresh
as well as ensiled pineapple wastes containing peel,
pomace, and leaves, and found that a growth rate of 600–
700 gm /day was achieved in growing and finishing steers
when fed with 90% waste material and 10% concentrate
mixture (Ranjhan, 1993). Dried cashew fruit contains
about 9.5% CP and 68.5% soluble sugars and it can
replace some of the grains in livestock ration (Aguilera,
1989). Mango byproducts, such as mango-seed kernels
contain 6% DCP and 70% TDN, and 5–7% tannin and can
be used as an ingredient in livestock rations (Sanon &
Kanwe, 2010).
Byproducts of the fermentation industry
Fermentation industry byproducts mainly consist of
grain, molasses, potato distillers’ soluble, brewer’s and
wine yeast, bacteria and fungi biomass as a waste for
production of lactic and citric and other organic acids,
as well as spent grain and malt sprout in breweries.
Molasses is a viscous byproduct obtained during the pro-
cessing of sugarcane or sugarbeets into sugar. Molasses
is one of the major sources of fermentable carbohydrate
(Cheeke, 1991) and is also a readily available source of
sugar and phosphorus to the animal body. The urea
molasses diet can be used as diet with little protein sup-
plement (Ranjhan, 1993) and it was reported that 10–15%
molasses addition in concentrate mixture increases the
palatability of the concentrate mixture (Verma, 1997).
Molasses contains 20.6% water, 60.8% total sugar, 3.2%
CP, 2.2% soluble gums, 8.2% ash, and 5.0% free acids.
Earlier Pathak et al. (1976) studied the effect of liquid urea
molasses diet on reproductive performance. Sugarcane
tops are palatable and cattle can be maintained entirely
on them with a little supplement of concentrate mixture
or leguminous feeds (Verma, 1997). Sugarcane tops
with or without leaves have good feeding value and are
readily accepted by ruminants, either in fresh, dried, or
ensiled form (Bhatti & Khan, 1996). Bagasse is a good
source of cellulose but is poor in protein (1.3%), high in
lignin (16%), and has been used successfully as roughage
for ruminants in many countries (Bhatti & Khan, 1996).
Digestibility of unprocessed bagasse is low because of its
high lignin content (Bhatti & Khan, 1996).
The major wastes from the sugarcane industry are
bagasse, molasses, and sugarcane press mud. Wine-
making industries produce grape pomace as a byprod-
uct consisting of skin, seed ,and stem with an estimated
amount of 13% by weight of grapes (Torres et al., 2002).
The distillery wastes mainly consist of yeast sludges. The
spent grain is the most abundant brewery byproduct cor-
responding to approximately 85% of the total byproducts
generated (Reinold, 1997). The spent grain accounts for
 Bio-processing of agro-byproducts to animal feed  5
an average of 31% of original malt weight representing Table 3. Use of agro-byproducts as animal feed.
20kg/100L of beer produced (Reinold, 1997). Brewer’s Agro-waste Animal References
spent grain is a lignocellulosic material containing about Mango byproduct Fish Mahadevaswamy and
17% cellulose, 28% noncellulosic polysaccharides, chiefly Venkataraman (1990)
arabinoxylans, and 28% lignin. Apple byproduct Poultry Zafar et al. (2005)
In general, organic substances, such as polysaccha- Tomato waste Poultry Lira et al. (2010)
rides, proteins, sugars, lipids, acids, starch and other Juice waste Poultry Rizal et al. (2010)
nutrients, such as vitamins, minerals, fibers, among Corn liquor waste Fish Chovatiya et al. (2011)
others are present in agro-industrial wastes. The major Brewery waste Chicken Gondwe et al. (1999)
organic compounds in most of the agro-wastes is ligno- Brewery waste Cow Dhiman et al. (2003)
celluloses which have a compact, partly crystalline struc- Brewery waste Hamster Zhang et al. (1992)
ture consisting of linear and crystalline polysaccharides, Brewery waste, Pig Dung et al. (2002)
cellulose, branched noncellulosic and noncrystalline wheat bran, rye bran
heteropolysaccharides, hemicelluloses, and branched Citrus byproduct Ruminant Bampidis and Robinson
(2006)
noncrystalline lignin (Glasser et al., 2000). Cellulose
Olive byproduct Sheep and Molina-Alcaide and
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is made up of a linear polymer chain that consists of a goat Yáñez-Ruiz (2008)

series of hydroglucose units in glucan chains with 1–4
glycosidic linkages. Hemicellulose consists of various
polysaccharides, such as xylose, galactose, mannose,
and arabinose. Hemicellulose is composed of linear and
branched heteropolymers of L-arabinose, D-galactose,
D-glucose, D-mannose, and D-xylose. Methyl or ace-
tyl groups are attached to the carbon chain to various
degrees. Lignin is made by an oxidative coupling of three
major C6 to C3 phenylpropanoid units, namely sinapyl
alcohol, coniferyl alcohol, and p-coumaryl alcohol.
These are arranged in a random, irregular three-dimen-
For personal use only.
Soyhull Cow Ipharraguerre and Clark
(2003)
feed with respect to their composition, digestion, degra-
dation, ruminal fermentation, and their impact on ani-
mal performance and product quality (Molina-Alcaide
&Yáñez-Ruiz, 2008). The soyhulls were also used as ani-
mal feed and the effects of feeding on ruminal fermenta-
tion, nutrient digestion, and utilization and performance
of dairy cows (Ipharraguerre & Clark, 2003) were also

sional network that provides strength and structure, and investigated.

are consequently highly resistant to enzymatic degrada-
tion (Vermerris, 2008).
Recently, scientific research has focused on many
aspects of agro-byproducts as animal feed, in particular
optimizing their nutritive value, characterization and
quantification of phenolic compounds and fatty acids,
their potential effects on the diet, animal performance,
and their product quality. Suitability of the byproducts as
animal feed is evaluated based on many factors, such as
types and proportion of byproducts generated, variability
in moisture and nutrient content, storage of the mate-
rial, potential for the presence of physical and micro-
bial contaminants and toxins, handling characteristics,
among others (Crickenberger, 1996). The suitability also
depends on the target animal, processing and handling
of byproducts, volume of material, costs versus benefits,
effects on feed consumption, safety concerns, etc.
Brewery waste is generally used as animal feed due to
its high protein content and fiber. It is a rich feed ingredi-
ent for ruminants, with other inexpensive sources such
as urea (Huige, 1994). The use of agricultural byproducts,
among other agro byproducts, as animal feed has been
studied for a range of animals including pigs, poultry,
fish, and cattle, as given in Table 3.
Citrus byproducts, such as fresh citrus pulp, citrus
silage, dried citrus pulp, citrus meal and fines, citrus
molasses, citrus peel and citrus-activated sludge have
been used as alternative feeds for ruminants in differ-
ent growth stages (Bampidis & Robinson, 2006). Olive
byproducts have been also evaluated as ruminant animal
The effluents from biogas production using mango-
processing wastes have been utilized for the production
of fresh-water fishes, such as carp, rohu among others
(Mahadevaswamy & Venkataraman, 1990). The nutri-
tional characteristics of some of the crop residues and
agro-industrial byproducts are summarized in Table 4.
Current limitation for utilization of agro-
byproducts as animal feed
There are many factors, such as protein concentration
and its biological value, digestibility, level of energy, fats
and carbohydrates, quantitative and qualitative compo-
sition of amino acids, vitamin and mineral content; and
the amount of fiber and the presence of hazardous or
toxic substances which determine the quality of byprod-
ucts as animal feed.
The nonconventional feed resources, such as agro-
residues and byproducts are underutilized or less
utilized due to various reasons. The production of agro-
residues is scattered and seasonal and in some cases,
the quality is low, especially for processing. The costs
of agro byproducts are noncompetitive and cheap for
the farmers. Processing of agro-byproducts is difficult,
costly, and problematic. Lack of managerial skills and
capital resources for the purchase and operation of suit-
able technology and also lack of the marketability of the
end-product also prevents the better utilization of agro-
processing byproducts. Generally, traditional agricul-
tural farmers have neither the resources and know-how

© 2012 Informa Healthcare USA, Inc.  

 6  C. M. Ajila et al.

Table 4. Nutritional characteristics of some of crop residues and agro-industrial byproducts.

Feed source Moisture (%) Crude protein (%) Crude fiber (%) Organic matter digestibility (%)
Rice bran 9.3–11.4 11.4–17.4 10.4–20.0 62.0
Rice hull 6.7–9.7 1.5–2.8 14.3–41.4 37.0
Bagasse 3.9–4.7 2.9–6.9 10.3–39.3 49.0
Brewer grain 9.8–10.8 24.0–27.4 15.9–17.1 60.0
Cocoa-pod husk 89.6 6.0 31.5 45.0
Coconut cake 10.0 18.0 12.0 78.0
Molasses 24.5 1.6 – 108
Palm-kernel cake 5.7 14.2 20.2 66.8
Palm-press fiber 13.8 4.0 36.4 30.8
Pineapple waste 6.8 4.9 20.8 76.0
Wheat bran 9.8 15.5 42.8 –

nor the quantities of residues to take individual action. Table 5. Antinutritional factors in agro-industrial byproducts
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Limited knowledge on the composition of the residues, feed.

such as proximate component intake and nutritive val- Agro-byproducts Antinutritional factor
ues, is also an important factor for the development of Rape seed and mustard Thioglucoside, goitrin, isothiocyanate
utilization technology for agro-byproducts. wastes
There are many factors which limit the use of feed Castor seed meal Ricin, hemagglutinin
from agro-byproducts (Ravindran & Blair, 1992) and it Cotton seed cake Gossypol
can be divided into two main categories as nutritional Linseed meal Cyanogens, anti B-6
and technological aspects, and are described below: Peanut meal Aflatoxin, goitrogen, protease
inhibitors, saponisn
Nutritional aspects:
Guar meal Protease inhibitors
–– Variability in nutritional level and quantity. Beet pulp Saponins
–– Presence of naturally occurring antinutritional and Sesame meal Mineral binders
For personal use only.

toxic factors. Soybean meal Hemagglutinins, goitrin, protease

–– Presence of pathogenic microorganisms
–– Need for nutrient supplementation.
–– Technological aspects:
–– Seasonal and unbelievable supply.
–– Bulkiness, wetness, and /or powdery structure.
–– Requirement of further processing.
–– Lack of proper research and development efforts.
A wide variety of antinutritional factors, including
various toxic compounds, which are deleterious to ani-
mal health and performance are found in agro-byprod-
ucts and are shown in Table 5 (Bhatti & Khan, 1996). The
ruminants in general, have the ability to tolerate much
higher concentrations of toxic factors as compared to
the nonruminants. A number of technologies and meth-
ods have been developed to detoxify or minimize the
effect of these toxins or antinutritional factors in animal
feeds and improved their nutritive value (Bhatti & Khan,
1996).
Developmental strategies for animal feed
from agro-residues
The quantity of utilization of byproducts as animal feed
depends on the abundance of the resources and the
technical equipment used for its preparation, preserva-
tion, and improvement. Recently, in many countries,
there has been a tendency toward maximum utilization
of all products which are suitable for feed purposes.
Further, the elaboration of suitable methods of collection,
inhibitors, saponins
Bhatti and Khan (1996).
transportation, and processing helps to reduce the costs
and improve the nutritive value.
The following strategies are considered important
in the objective to increase productivity from the ani-
mal resources through more efficient utilization of
agro-byproducts:
1. All year-round feeding systems.
2. More intensive use especially in stall-feeding
systems.
3. Development of complete rations.
4. Strategic supplementation.
5. Wider use of agro-byproducts.
6. Nutrient enrichment of agro-residues.
Agro-byproducts should be expanded because of the
need for more intensification in the future. A change
from the traditional to more intensive systems of pro-
duction is likely to result from different factors such as,
scarcity of grazing land, need for more control over ani-
mals, and higher returns from the stall-feeding system.
Various techniques are used for the better utilization of
agro-byproducts as animal feed, which includes:
1. Removal of substances or microflora harmful to ani-
mal health.
2. Increasing storage-life and subsequent use as feed
components or directly as fodder.

 

3. Improving digestibility and nutrient availability.
4. Improving the nutritional value by using some con-
stituents as a medium in the biosynthesis of protein
and of other biologically active substances required
in animal feeding.
Mycotoxins are one of the major toxic compounds
found in many agro-byproducts and animal feed.
Mycotoxins are fungal secondary metabolites that affect
both human and animal health. Aflatoxins are carci-
nogenic compounds produced by certain strains of the
Aspergillus genus and are often found as a contaminant
of ruminant diets. There are five major classes of myco-
toxins, which are of particular concern in the animal
feed industry, such as aflatoxins, tricothecenes, zearale-
Critical Reviews in Biotechnology Downloaded from informahealthcare.com by 206.162.180.2 on 03/01/12
nones, fumonisins, and ochratoxins. The ideal solution
for minimization of health risk that aflatoxins pose is the
prevention of feed contamination (Saima et al., 2009). A
potential strategy to reduce mycotoxicoses in animals is
to use microorganisms for biotransformation, and thus
transform the mycotoxin into another nontoxic metabo-
lite (Schatzmayr et al., 2006). Many species of bacte-
ria and fungi, such as Flavobacterium aurantiacum,
Corynebacterium rubrum, Candida lipolitica, Aspergillus
niger, Trichoderma viride, Armillariella tabescens,
Neurospora spp., Rhizopus spp. and Mucor spp., among
others. have been shown to enzymatically degrade myco-
For personal use only.
toxins (Bata & Lasztity, 1999). Many strains of lactic acid
bacteria (LAB) have been shown to bind aflatoxin in a
strain-specific manner (Peltonen et al., 2001).
Mycotoxins can contaminate agricultural products in
the field during storage and processing as animal feed.
The use of pre-harvest control strategies, such as resis-
tance varieties, field management, use of biological and
chemical agents, harvest management, and postharvest
applications, including improving drying and storage
conditions, together with the use of natural and chemi-
cal agents, and irradiation have clearly been shown to
be important in the prevention of mycotoxigenic mould
growth and mycotoxin formation (Milicevic et al., 2010).
Natural plant extracts and spice oils of eugenol, cinna-
mon, oregano, onions, lemongrass, turmeric, mint, and
chemical compounds (fungicide, herbicide, and surfac-
tant) are reported to prevent both mycotoxigenic mould
growth and mycotoxin formation during postharvest
season (Kabak et al., 2006). Precise antifungal proper-
ties of LAB are still not clear, but may involve microbial
competition, as well as extracellular metabolites which
are heat-stable and of low molecular weight.
There is no single step detoxification method reported
which will be suitable for all foods and animal feeds. The
effectiveness of a detoxification method for mycotoxins
depends on the nature of the food, environmental condi-
tions (moisture content, temperature) as well as the type
of mycotoxin, its concentration, and the extent of binding
between mycotoxin and constituents. Recently, there has
been an increasing interest in the use of bacteria, yeast,
and fungi to help minimize the toxic effect of mycotoxins
© 2012 Informa Healthcare USA, Inc.  
Bio-processing of agro-byproducts to animal feed  7
(Kabak et al., 2006). The most recent approach to the
problem has been the use of mycotoxin-binding agents in
the diet that sequesters the mycotoxin in the gastrointes-
tinal tract thus reducing their bioavailability. Chemicals,
such as aluminosilicate, zeolite, and bentonite have
shown good potential for use in the animal feed to help
overcome aflatoxicosis (Kabak et al., 2006).
SSF as a bio-processing method
Recently, there has been an increasing trend toward the
utilization of the solid-state fermentation (SSF) tech-
nique to produce several bulk chemicals and enzymes
(Robinson & Nigam, 2003). This methodology has often
been found to produce a more stable product, with less
energy requirements, in smaller fermenters and smaller
volumes of polluting effluents, as compared to sub-
merged fermentation (SmF) systems. SSF is defined as
any fermentation process carried out on a solid material
in the absence of free-flowing liquid (Pandey, 1992). A
wide range of solid materials can be employed as sup-
ports in SSF and they are usually classified into two
categories: noninert and inert materials (Krishna, 2005).
In the first category, a divided and humidified solid (e.g.
cereal grain, flour, bran, sawdust) behaves as support
and nutrient source, while in the second, a nutritionally
inert solid (e.g. synthetic foam), acting exclusively as a
support, is soaked in a nutrient solution. It is noteworthy
that there is growing interest in utilizing agro-industrial
wastes as noninert SSF supports, making the whole pro-
cess much more economical (Pandey et al., 2000). The
use of SSF has been proposed not only as a method for
food preparation or amelioration, but also as a suitable
technology for obtaining useful additives for food and
feeds or reagents usually employed in food manufacture
(Robinson & Nigam, 2003) (Figure 2).
For the production of animal feed
Solid fermentation of agro-industrial residues has been
proposed as a suitable pretreatment that can lead its use
as animal feed. SSF can upgrade the nutritive value of
agricultural byproducts and represent potential solution
to feeding animals in developing countries (Robinson
Fungi
Agro-industrial
Yeast
by-products Solid state fermentation
Bacteria
Feed additives
Protein enriched
animal feed
Enzymes, Vitamins,
colorants, antibiotics
Figure 2. Food and feed applications of agro-industrial waste by
solid-state fermentation.
 8  C. M. Ajila et al.
& Nigam, 2003). The advantage of using SSF to achieve
both goals is the low-tech fermentation system requir-
ing the possibility of performance on farms. The use of
SSF for protein enrichment of lignocellulosic residues
has been a focus of attention due its direct applicability
of the fermented product for ruminant feed purposes.
Generally, agro-byproducts have low protein content
(2–6%) and fungal growth can improves it to 10–15%
(Shojaosadati et al., 1999). However, the degradability of
dietary protein in the rumen from fermented lignocellu-
losic byproduct was higher compared with unfermented
byproducts (Iconomou et al., 1998). Besides protein, an
increase of total lipid and fatty acids was observed after
solid-state fungal fermentation of agricultural residues
(Abu et al., 2000). SSF has been reported to increase
Critical Reviews in Biotechnology Downloaded from informahealthcare.com by 206.162.180.2 on 03/01/12
the protein content in cereal grain and potato residues.
Gélinas and Barrete (2007) obtained an increase of 7–8%
of protein content of potato processing waste by SSF
by using yeasts. Fungal growth showed decrease in the
neutral detergent fiber , acid detergent fiber , and hemi-
cellulosic contents of lignocellulosic fibers, increase
in the bioavailability of nutrients, and decrease in the
antinutritional factors, such as phytic acid, polyphenols,
and tannins (Manmdebvu et al., 1999; Elyas et al., 2002).
Thus, two nutritionally important contributions can be
obtained from the fermentative process: depolymerizing
enzyme and protein enrichment and high degradability
For personal use only.
of the feed material.
Nutrient-enriched animal feed from byproducts of
plant origin
Apple pomace, a byproduct from the apple-processing
industry, is not a suitable feed since it is deficient in
digestible protein from an animal-nutrition point of view
(Rumsey, 1978). Growth of yeast on the apple pomace
increases protein and vitamin contents (Hang, 1988).
However, the low level of fermentable sugars limits pro-
tein enrichment of the pomace by yeasts; a major portion
of the pomace comprises lignocelluloses. It was reported
that coculture of Candida utiIis and A. niger on dried and
pectin-extracted apple pomace increased the protein
content to 20% and 17%, respectively (Bhalla & Joshi,
1994).
Total CP content of 7.5% (w/w) increased during the
solid-substrate fermentation of apple using Candida
utilis and Kloeckera apiculafa after 72 h for each yeast.
The concentration of essential amino acids in the modi-
fied apple pomace was more than twice in the control,
enhancing its nutritive value as a stock-feed supplement
(Rahmat et al., 1995). The fermentation of pomace as
described can be used to reprocess this waste material
into a useful value-added product for the agricultural
sector. The higher yield of protein from the protein
enrichment of apple pomace by C. utilis and A. niger
combination probably resulted from the enzymatic
hydrolysis of the lignocellulosic component of the pom-
ace by A. niger releasing hexoses and pentoses which can
 
be efficiently metabolized by C. utilis. Saccharomyces
cerevisiae, however, uses only hexoses (Norris, 1981)
and consequently is less efficient than Candida spp.
Sequential fermentation of apple pomace by C. utilis and
Pleurotus ostreatus resulted in a significant CP and min-
eral-level enrichment, which could permit the utilization
of this residue as protein supplement in cattle feeding
(Villas-Bõas et al., 2003).
Zheng and Shetty (1998) used apple pomace to pro-
duce a food rich in proteins by employing the fungus
Rhizopus oligosporus. High moisture content results in
an increase in mycelial production, consequently inter-
fering in the transfer of oxygen. The protein enrichment
of apple pomace by Gongronella butleri through solid-
state cultivation of apple pomace has been investigated
and the resultant protein-rich diet increased 44% of the
body mass in tilapia fry, which demonstrates that bio-
transformed apple pomace may represent an excellent
food supplement (Vendruscolo et al., 2009). Apple pom-
ace has been used as the raw material to produce bio-
feed with enriched enzymes, such as pectinase, cellulase,
xylanase, proteinase, and amylase (Zheng & Shetty, 2000;
Zhong-Tao et al., 2009). The limited use of palm-kernel
cake in fish feeds, due to its high fiber and low protein
content, could be overcome by processing this material
under solid-state culture conditions with some fungal
strains (Trichoderma longiobrachiatum). This treat-
ment led to a significant increase in protein level, and a
decrease in cellulose and hemicellulose (Iluyemi et al.,
2006). Bisaria et al. (1997) used the fungus Pleurotus
sajor-caju in the bioconversion of rice straw and wheat
straw. The authors observed that supplementation of the
solid residue with urea and ammonium nitrate increased
the bioconversion and the protein level increased from
2.87 to 6.3% (w/w) with rice straw and from 3.1 to 7.5%
(w/w) with wheat straw.
The fermentation of cassava root by Saccharomyces
cerevisiae enhanced the nutritive value with an increase
in protein and fat contents and also reduced hydro-
cyanic acid contents (Boonnop et al., 2009). Different
vegetable processing wastes, such as Chinese cabbage
juice (Choi & Park, 2003), waste brine generated from
kimchi production (Choi & Park, 1999), deproteinized
leaf juices (Chanda & Chakrabatri, 1996), corn silage
juice (Hang et al., 2003) has been used as a nutrient
source for yeast growth. Single-cell protein production
by SSF of orange peel has been reported and the protein
can be used as a component in ruminant animal feed
(Nicolini et al., 1993). Potato pulp is the industrial waste
produced during starch extraction from potatoes and
its discharge has become an environmental-pollution
problem and is a decisive cost factor due to stricter
environmental legislation and associated expenses
in the potato-starch industry. Recently, a model was
developed to assess potato pulp for use as a new poultry
feed, which will help starch factories to decrease waste
discharge (Wang et al., 2010). Animal feed produced by
SSF of wastes from potato pulp has been assessed for

its toxicity, nutritional quality and its effect on growth
performance of poultry farming.
Nutrient-enriched animal feed from fermentation-
industry byproducts
Bagasse has been commonly used for the production of
protein-enriched animal feed by SSF, employing yeasts
and fungi. Nigam et al. (1987) and Nigam (1990) inves-
tigated SSF of bagasse for animal-feed production using
basidiomycetes. Zadrazil and Puniya (1995) differenti-
ated bagasse into four fractions of particle size with a view
to enhancing its nutritive value as animal feed and found
that the difference in particle size affects degradation by
white-rot fungi and also in vitro rumen digestion. SSF of
bagasse has been carried out using a strain of P. sajor-caju
Critical Reviews in Biotechnology Downloaded from informahealthcare.com by 206.162.180.2 on 03/01/12
in a closed system, with the aim of optimization of the gas-
eous atmosphere and development of a cost-effective and
simple technology for animal-feed production (Puniya et
al., 1996). There was 21% CP increase and 45% depletion
of lignocelluloses in bagasse after 12 weeks’ cultivation of
a fungal strain of Streptomyces (Iyo & Antai, 1991). A pat-
ent was obtained on the application of bagasse, softened
with alkali treatment, for feedstuff, fertilizer, and sweet-
ener by cultivating Enterococcus faecium in SSF (Iritani et
al., 1995). Chaudhary et al. (1994) also reported feedstuff
production from bagasse using two strains of Pleurotus sp.
The use of distillery grape stalks from the winery indus-
For personal use only.
try in the form of single-cell protein, as ruminant feed or
feeding component has also been proposed (Nicolini et
al., 1993) by using SSF. After biological lignin removal,
the cellulose was better accessible to rumen microorgan-
isms due to its protein value and low lignin content.
Production of carotenoid-enriched animal feed by
secondary fermentation of co-products of grain-based
ethanol industry has been reported. Whole stillage is a
coproduct of grain-based biofuel and is dried to produce
dried distiller’s grain. Recently, carotenoid enrichment of
dried distiller’s grain has been carried out by submerged
fermentation of red yeasts. Mono and mixed culture of
Phaffia rhodozyma and Sporobolomyces roseus were used
to produce astaxanthin and β-carotene. The secondary
fermentation presents an innovative strategy to utilize
whole stillage and enhance market-growth potential of
distiller’s grain (Ananda et al., 2009).
Generally, the byproducts from plant origin are rich
in nutrients. There are many reports on the utilization
of these agro-byproducts as animal feed and nutrient
enrichment can be performed by SSF and bio-processing
technologies. Most of the reports showed that nutritional
and nutraceutical properties of the agro-byproducts as
solid substrates were improved by fermentation. Thus,
scope exists for microbial upgradation of low-quality
agro-wastes and development of healthy animal-feed
supplements. The production of animal feed from agro-
residues represents one of the economically beneficial
methods due to the continuous and huge demand for
animal feed. The marketing is also relatively easy and
the technologies involved are not too complicated. The
© 2012 Informa Healthcare USA, Inc.  
Bio-processing of agro-byproducts to animal feed  9
technologies involved can be handled in small-scale
industries also. The production of animal feed from agro-
residues will be one of the most sustainable technologies
for better economic resources of local agro-communities
and also for the better management of environmental
pollution.
Production of enzymes as additives in feed
Animal-feed additives are products used in animal
nutrition for the purpose of improving the quality of
feed and food from animal origin, or to improve the ani-
mals’ performance and health, e.g. providing enhanced
digestibility of the feed materials. Feed additives may
not be brought into the market unless authorized fol-
lowing a scientific evaluation demonstrating that the
additive has no harmful effects on human and animal
health and the environment (EUROPA, 2008). Most of
the agricultural products contain antinutrients which
limit the nutritional value of the feed stuff. The reduc-
tion in nutrient utilization due to the antinutrients can
contribute to environmental pollution due to excessive
secretion of unabsorbed nutrients, especially nitrogen
and phosphorus and also increase the production cost
because of insufficient feed utilization (Woyengo et al.,
2010). Depolymerizing enzymes are used as feed addi-
tives to increase the digestibility and nutritional value of
feeds (Llewellyn et al., 2010; Nortey et al., 2007; Sands &
Kay, 2007). The use of agro-byproducts as a substrate for
SSF for the production of various feed-grade enzymes,
such as xylanase, amylase, cellulase, pectinase, chitin-
ase, liginases, and phytases is more important because
in addition to its process-engineering advantages, the
enzymes may be produced and used in feeds in the situ-
ation where the substrate is also used as enriched feed
with enzymes and proteins, leading to enhanced digest-
ibility and nutritive value (Ugwuanyi, 2008).
Enzymes have been used for decades to improve the
utilization of swine and poultry diets. Recent studies
have examined the use of exogenous enzyme products
in high-forage diets fed to growing cattle (Wang et al.,
1999; ZoBell et al., 2000). The enzymes such as phytase,
amylase, b-glucanase ,and xylanase are added to the
cereal-based diets of monogastrics to increase the utili-
zation of dietary P, starch, β-glucans, and arabinoxylans.
It was reported that supplementing dairy cow and feedlot
cattle diets with fiber-degrading enzymes has significant
potential to improve feed utilization and animal perfor-
mance (Beauchemin & Rode, 1996). The improvements in
animal performance due to the use of enzyme additives
can be attributed mainly to improvement in ruminant
fiber digestion (Ghorai et al., 2009) resulting in increased
digestible energy intake. Spraying enzymes onto feeds
prior to feeding provides increased management flexibil-
ity and bypasses any negative interactions that the ensil-
ing process may have on silage enzyme performance.
Treating feeds with enzymes may improve digestibility
via a number of different mechanisms including, direct
 10  C. M. Ajila et al.
Table 6. Production of enzymes using agro by-products for animal nutrition use.
Agro-residue Microorganisms
Sugarcane bagasse Penicillium echinulatum
Sugarcane bagasse, wheat straw, Phanerochate chrysosporium
corn cob, rice husk, peanut shell
Wheat straw Aspergillus sp.
Rice straw Trichoderma reesei
Barley Talaromyces emersonii
Corn silage Thermiascus aurantiacus
Corn stalk Fusarium oxysporum
Citrus waste Eupencillium javanicum
hydrolysis, improvements in palatability, changes in gut
viscosity, complementary actions with ruminal enzymes,
and changes in the site of digestion. Fungal direct-fed
Critical Reviews in Biotechnology Downloaded from informahealthcare.com by 206.162.180.2 on 03/01/12
microbials (DFM) have been popular additions to rumi-
nant diets for many years. In some applications, DFM
may be mixed with feed or administered in the drinking
water (Ghorai et al., 2009).
The use of exogenous enzymes and other additives
to improve the nutrient availability of feed has found
important feed application in animal farming. A num-
ber of studies and reviews on SSF production and use of
depolymerizing enzymes using agro-residues in animal
nutrition have been recently published (Cao et al., 2010;
Ugwuanyi, 2008). Table 6 summarizes the literature avail-
able on byproducts that can be used in enzyme produc-
For personal use only.
tion related with animal nutrition.
α-Amylases
α-Amylases (endo-l, 4-a-D-glucan glucanohydrolases EC
3.2.1.1) are extracellular enzymes that randomly cleave
the 1, 4- linkages between adjacent glucose units in the
linear amylose chain, ultimately generating glucose,
maltose, and maltotriose units. Microbial α-amylases
are mainly secreted as primary metabolites and their
production is reported to be growth associated (Francis
et al., 2003).
Amylase is a class of enzymes which can catalyze
hydrolysis of starch. It mainly contains α-amylase,
β-amylase, glucoamylase, and branch amylase.
Normally, α-amylase is used in feedstuff, which can
catalyze hydrolysis of starch into oligosaccharide, disac-
charide, dextrin, glucose, and fructose, and then improve
the digestibility of starch. Amylase as animal feed has
supplemented the deficiency of endo-enzymes during
weaning period, improved the utilization of grain starch,
and reduced the cost of feed. Avizyme 1505 is an enzyme
preparation containing endo-1,4-β-xylanase produced
by a genetically modified strain of Trichoderma reesei,
subtilisin produced by a genetically modified strain of
Bacillus subtilis and α-amylase produced by a genetically
modified strain of Bacillus amyloliquefaciens. Avizyme
1505 is marketed in a dry powder form containing a
minimum guaranteed of 1500 xylanase, 20,000 subtilisin,
and 2000 amylase U/g. Avizyme can be used as zootech-
nical additive for laying hens with animal feed, under
the functional group of digestibility enhancer. The EFSA
 
Enzymes Reference
Cellulase, xylanase Camassola and Dillon (2010)
Cellulase, xylanase El-Nassar et al., (1997)
Cellulase Abo-State et al., (2010)
Hemicellulose protease Eun et al., (2006)
Endo-β-glucanase McCarthy et al., (2005)
Xylanase Colombatto et al., (2004)
Endoglucanase β-glucosidase Panagiotou et al., (2003)
Xylanase, pectinase Tao et al., (2011)
(European Food Safety Authority) panel on Additives and
Products or Substances used in Animal Feed (FEEDAP)
analyzed the safety and efficacy of Avizyme 1505 (endo-
1,4-β-xylanase, subtilisin, α-amylase) as a feed additive
for laying hens and concluded that Avizyme 1505 is safe
for laying hens at the recommended dose (EFSA, 2011).
Dietary supplementation with Aspergillus oryzae
α-amylase preparation improved dairy cattle perfor-
mance and increased the milk production (Tricarico et
al., 2005; 2008). Dietary supplementation with A. oryzae
α-amylase preparation also improved finishing beef
cattle performance by improving the average daily gain
occurred during the initial 28 days on feed (Tricarico et
al., 2007).
Agro-industrial wastes are widely used as substrates
for amylase production. A mixture of wheat bran and
groundnut-oil cake, supplemented with potassium dihy-
drogen phosphate and 1% soluble starch was found to
be an excellent source of α-amylase production using
Bacillus amyloliquefaciens ATCC 23842 (Gangadharan et
al, 2006) out of 14 different agro-residues. Banana wastes
(i.e. peel, stalk) have also been used as a solid substrate
for α-amylase production by Bacillus subtilis (Kokab et
al., 2003; Krishna & Chandrasekaran, 1996) and A. oryzae
(Ragunathan & Swaminathan, 2005). Francis et al. (2003)
used spent grains in SSF for the production of α-amylase
by A. oryzae and found that supplementation of fermen-
tation media with Tween-80 or calcium ions enhanced
enzyme activity. It was reported that solid-substrate fer-
mentation material (crude enzyme, in situ enzyme) may
be considered a cheap biocatalytic material for animal-
feed rations and for alcoholic production from starchy
materials (Bogar et al., 2002). Ramachandran et al. (2004)
reported the use of different oil cakes, such as coconut,
sesame, groundnut, palm kernel, and olive as SSF sub-
strates for α-amylase production by A. oryzae under SSF
conditions. Some agro-industrial residues that have been
used for the production of microbial amylases include
rice bran (Akpan et al., 1999) wheat bran, (Pandey, 1993),
copra waste (Pandey et al., 1995), tea waste (Selvakumar
et al., 1998), cassava flour, oil-palm waste, apple pom-
ace, banana waste (Nigam & Singh, 1994), and cowpea
waste (Kareem et al., 2009). A comparative study of the
ability of B. licheniformis and B. subtilis to produce these
enzymes on potato peel and wheat bran under SSF was
also reported (Shukla & Kar, 2006).

Pectinases
Pectinases are enzymes that hydrolyze pectic substances,
which occur as structural polysaccharides in the middle
lamella and primary-cell walls of higher plants (Fogarty
& Ward, 1972). These enzymes include polygalacturo-
nase, pectin esterase, pectin lyase, and pectate lyase
on the basis of their mode of action. They are added to
livestock feed to help animals digest their feed better.
Pectinase eliminates the antinutritional factors of pectin
and cellulose and releases the nutrients, enhances the
digestibility of the animal in a wide range, and promotes
the viscosity of plant tissue, and improves animals on the
digestibility of feed.
Polygalacturonase production by Bacillus sp.
MG-cp-2 increased significantly during SSF using cheap
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agro-residues, such as wheat bran, sunflower seed cake,
rice bran, and orange peel (Soares et al., 2001). The pro-
duction of endo- and exo-polygalacturonase enzymes
by Aspergillus awamori grown on wheat (wheat grains
and milled wheat) was studied, and some of the fac-
tors influencing SSF, including particle size, chemical
composition, and moisture content, were evaluated
(Blandino et al., 2002). Production of polygalacturo-
nases and pectin lyase using Penicillium viridicatum
(Silva et al., 2005) and Fusarium moniliforme (Niture &
Pant, 2004) by SSF of agrowaste mixtures, such as orange
bagasse and wheat bran has been reported. Solid-state
For personal use only.
production of pectinases by A. niger strains using differ-
ent agro-industrial residues as substrates, such as wheat
bran, coffee pulp and husk, cranberry and strawberry
pomace, citrus peel, orange bagasse, sugarcane bagasse
and wheat bran have been reported (Antier et al., 1993;
Boccas et al., 1994; Castilho et al., 2000; Dhillon et al.,
2004; Fontana et al., 2005; Singh et al., 1999; Zheng &
Shetty, 2000).
Thermostable pectin lyase and polygalacturonase are
produced by the SSF of agro-industrial residues using
newly isolated thermophilic Thermoascus aurantiacus
(Martins et al., 2002). This fungus is able to produce high
levels of extracellular enzymes during SSF of mixtures
of sugarcane and orange bagasse. Recently, Patil and
Dayanand (2006) evaluated pectin-rich agro-wastes,
namely lemon peel, sorghum stem, and sunflower
head, as substrates for the production of pectinase by
Aspergillus niger DMF 27 and A. niger DMF 45 in SmF and
SSF systems, respectively. It was reported that Pencillium
sp. isolated from pectin industry waste produced polyga-
lacturonase by SSF of different agro-residues comprising
waste and byproducts, such as banana peel, wheat bran,
sugarcane bagasse, and orange bagasse. Production
and optimization of polygalacturonase from mango
(Mangifera indica L.) peel using Fusarium moniliforme
by SSF was reported (Kumar et al., 2010).
Lipases
Lipases (triacylglycerol acylhydrolases, EC 3.1.1.3) are
well known as efficient biocatalysts for the hydrolysis of
© 2012 Informa Healthcare USA, Inc.  
Bio-processing of agro-byproducts to animal feed  11
water-insoluble fatty-acid esters, being triacylglycerols
of long-chain fatty acids as natural substrates. Lipases
are nowadays widely used at industrial scale with appli-
cations as additives in detergents, the elaboration of
dietetic foods for use in the food industry, obtaining
bioactive molecules in the pharmaceutical industry
and pure optical compounds in chemical synthesis pro-
cesses, as well as modification of fats and lipids by hydro-
lysis and esterification reactions (Hernaiz & Sinisterra,
1999; Kazlauskas, 1994). Lipases may also be applied as
an additive to animal feed. The resulting breakdown of
fats in feed is assumed to facilitate the development of
lean and qualitatively high-value meat.
There are several reports on the extracellular lipase
production by fungi, such as Rhizopus sp., Aspergillus sp.,
and Penicillium sp. on different solid substrates, such as
gingelly-oil cake, babassu-oil cake, olive cake, soy cake,
coconut-oil cake, wheat bran, rice bran, almond meal,
and sugarcane bagasse (Adinarayana et al., 2004; Di
Luccio et al., 2004; Mahadik et al., 2002; Ul-Haq et al.,
2002). The use of mixed solid substrates, such as coconut-
oil cake:wheat bran (1:1) (Benjamin & Pandey, 1998) or
wheat bran:olive oil (9:1) (Nagy et al., 2006), will be more
advantageous for lipase production because of the pres-
ence of lipase inducers such as olive oil. Supplementation
of SSF medium with lipase inducers (i.e. olive oil) strongly
affected lipolytic enzyme production (Di Luccio et al.,
2004; Mahadik et al., 2002; Palma et al., 2000). Recently,
it has been reported that the lipase produced by A. niger
J-1 grown on wheat bran has a good stability in relation
to temperature and pH (Falony et al., 2006). The cultiva-
tion conditions and operation of tray and packed-bed
bioreactors for lipase production by Penicillium simplicis-
simum under SSF using babassu cake as a basal medium
was studied (Gutarra et al., 2005). It was reported that
the C/N (carbon/nitrogen) ratio (6–6·5 and 9–9·5) of the
medium is an important parameter for lipase production
by the yeast Candida rugosa in solid-state culture (Rao
et al., 1993). Domínguez et al. (2003) reported the great
potential of food and agro-industrial wastes (groundnut
and barley bran) as support-substrates for lipase produc-
tion in solid-state cultures of the yeast Yarrowia lipolytica.
Tannases
Tannases (tannin acyl hydrolases, EC 3.1.1.20) cata-
lyze the breakdown of hydrolyzable tannins or gallic
acid esters. These enzymes are used as clarifying agents
in the industrial processing of fruit juices and coffee-
flavored soft drinks, in the manufacture of instant tea,
and the production of gallic acid in leather industry, for
manufacture of antibacterial agents and antioxidants
(Aguilar et al., 2007). Tannins form insoluble complexes
with proteins and interaction plays an important role in
nonruminants (rabbits, poultry among others) industry.
The tannin in many feed materials complex with dietary
and endogenous proteins as well as digestive enzymes,
thereby interfering with normal digestion, leading to
a drain of high-quality protein from the body (Belur &
 12  C. M. Ajila et al.
Mugeraya, 2011). Tannins are also known to interfere with
the absorption of iron and also damage the mucosal lin-
ing of the gastrointestinal tract. The use of tannase as an
ingredient of animal feed would improve the digestibility
of the feed (Lekha & Lonsane, 1997). The enzyme has
potential uses in the treatment of tannery effluents and
pretreatment of tannin-containing animal feed (Aguilar
et al., 2007; Murugan & Al-Sohaibani, 2010). Tannase is
synthesized in the presence of tannic acid by various fila-
mentous fungi, principally A. niger and Penicillium chrys-
ogenum, and also by some bacteria and yeasts (Mondal
et al.,  2001) and mainly by SmF and SSF (Lekha &
Lonsane, 1997). Agro-residues and forest products
serve well for being used as the substrate for production
of microbial enzymes by SSF. Sabu et al. (2006), have
Critical Reviews in Biotechnology Downloaded from informahealthcare.com by 206.162.180.2 on 03/01/12
reported the use of palm-kernel cake and tamarind-seed
powder as the substrate for the production of tannase
from A. niger ATCC 16620 by SSF. Battestin and Macedo
(2007) have reported the use of coffee husk and rice bran
as media ingredient for SSF of Paecilomyces variotii.
Production of tannase by modified SSF of tannin-rich
agro-residues comprising powdered fruits of Terminalia
chebula and powdered pod cover of Caesalpinia digyna
using improved strains of Aspergillus foetidus and R. ory-
zae was reported (Purohit et al., 2006). Tannase produc-
tion by Penicillium atramentosum KM under SSF using
cheap and locally available agro-residues, such as jamun
For personal use only.
and keekar leaves were reported recently (Selwal et al.,
2011). A comparative study was conducted on the suit-
ability of different agro-byproducts’ residue substrates
such as paddy husk, rice bran, millet husk, and groundnut
shell for the production of tannin acyl hydrolase produc-
tion using A. oryzae by SSF (Paranthaman et al., 2009),
and rice bran was found to be the best substrate for the
production of enzyme with higher activity. Bioconversion
of sugarcane bagasse and rice-straw powder for tannase
production and optimization of fermentation-processing
condition required for maximum tannase production was
studied (Paranthaman et al., 2010).
Xylanases
Xylanases are enzymes which degrade the linear poly-
saccharide β-1,4-xylan into xylose, thus breaking down
hemicellulose, one of the major components of plant-
cell walls. One rapidly developing area of application for
xylanases is their use as animal-feed additive for poultry
and pigs and other monogastric animals that do not have
xylanolytic microflora in their digestive tract. Xylanases
break down the nonstarch polysaccharides into high
fiber rye- and barley-based feeds, thus reducing viscos-
ity and increasing absorption (Bedford & Classen, 1992).
Xylanases are able to decrease the viscosity of chyme in
the gastrointestinal tract and stimulate the digestion and
absorption of nutrient and also able to release nutrient
and improve the availability of feedstuff, and ameliorate
the micro-ecosystem in the gastrointestinal tract. There
are many reports on the production of xylanse by SSF
of agro-residues using different microorganisms. Uma
 
Maheswari and Chandra (2000) isolated a mesophilic
Streptomyces cuspidosporus strain which was cultivated
under solid-state conditions on wheat bran. The crude
enzyme preparation, which contained mainly xylanase
but also amylase and phytase activities, was applied in
predigestion of poultry feed and was found to be effective
in releasing sugars and soluble phosphorous. A two-stage
process that combined SSF and ensiling was used for
bio-conversion of corn straw, in order to increase nutri-
tional value and palatability for animal feed. Milagres et
al. (2004) investigated and optimized xylanase produc-
tion by Thermoascus aurantiacus ATCC 204492 by SSF
on sugarcane bagasse using a glass-column reactor with
forced aeration. The new Bacillus circulans strain was
grown under SSF on industrial fibrous soy residue in a
horizontal rotated drum bioreactor, and the procedure
for xylanase extraction was investigated and optimized
(Heck et al., 2002). Alkalophilic Bacillus subtilis produced
high levels of xylanase using easily available inexpensive
agricultural waste residues, such as wheat bran, wheat
straw, rice husk, sawdust, gram bran, groundnut, and
maize bran in SSF. Among these, wheat bran was found
to be the best substrate (Sanghi et al., 2008).
Phytases
Phytases (myo-inositol hexakisphosphate phospho-
hydrolases, EC 3.1.3.8) catalyze the release of phos-
phate from phytate, which is one of the major forms of
phosphorus predominantly occurring in cereal grains,
legumes, and oilseeds. Monogastric animals, such as pigs
and poultry, need a certain amount of phosphorous for
their skeletal growth, but they lack the enzymes needed
to efficiently digest phytate in their feed. Consequently,
they excrete large amounts of phosphorous into the envi-
ronment, thus causing an important pollution problem.
Supplementation of phytase to their feed appears as an
interesting strategy for upgrading the nutritional quality
of phytate-rich feed, reducing inorganic P supplementa-
tion and consequently fecal P excretion (Pandey et al.,
2001). The addition of microbial phytase to the feed-
stuff of monogastric animals was described as early as
25 years ago. Addition of phytase to feed substantially
improved phosphorus utilization and reduced excretion
of phosphorus in the feces and also counteracted the
antinutritional properties of phytate. Supplementation
of phytase in animal feed increased phosphorous avail-
ability by 60% and also reduced or even eliminated the
need for supplemental dietary inorganic phosphorous.
It also decreased the excretion of phosphorous in the
feces by over 30% and also counteracted the antinutri-
tional properties associated with unhydrolyzed phytate.
Phytases have a big share in enzyme business due to their
widespread application as a feed supplement (Greiner &
Konietzny, 2006; McCoy, 1998; Vohra & Satyanarayana,
2003).
Although a broad range of microorganisms can pro-
duce phytase, the expression levels of phytase in them
are too low for economic considerations. Due to the

high cost of phytase production, application of phytase
has not been widespread. For economic considerations,
inorganic phosphorous supplement of monogastric ani-
mal feedstuff is still the method of choice. Commercially
viable procedures for the production of a large quantity
of phytase would be of tremendous value both from the
feed-conversion and environmental points of view.
Phytase can be produced in SSF by filamentous fungi
on selected feed ingredients and the crude product may
be mixed in feed rations as a value-added supplement
(Pandey et al., 2001; Bogar et al., 2003). Fungal phytase
has received more attention due to high-production
yields and acid tolerance for feed production (Kim et
al., 1998). Nampoothiri et al. (2004) studied the produc-
tion of thermostable phytase by a thermophilic strain,
Critical Reviews in Biotechnology Downloaded from informahealthcare.com by 206.162.180.2 on 03/01/12
Thermoascus aurantiacus in submerged fermentation
and optimized various process parameters for phytase
using synthetic medium supplemented with wheat
bran or wheat-bran extract. Various filamentous fungi,
such as Mucor, Aspergillus, and Rhizopus sp. have been
reported to produce phytase and accessory enzymes
by SSF. Although several strains of bacteria, yeasts, and
fungi have been used for production of phytase, two
strains of Aspergillus sp., A. niger,and A. ficuum, have
most commonly been employed for commercial produc-
tion (Pandey et al., 2001; Sabu et al., 2005). The influence
of microbial phytases on digestibility and other related
For personal use only.
properties of sorghum, wheat, maize, and barley based-
feeds for broiler chickens has been evaluated, individu-
ally or in combination with glycanases (Wu et al., 2003;
Wu et al., 2004). Papagianni et al. (2000) compared phy-
tase production by A. niger in SmF and SSF, based on the
influence of medium composition and fungal morphol-
ogy using wheat bran as substrate. Krishna and Nokes
(2001) cultivated A. niger under SSF on wheat bran and
full-fat soybean flour, and indicated significant influ-
ence on age of liquid inoculums and cultivation time
on enzyme levels. SSF using agricultural residues, such
as wheat bran, mustard cake, cowpea meal, groundnut
cake, coconut cake, cotton cake and black-bean flour
using thermotolerant strain A. niger NCIM 563 produced
phytase, and maximum extracellular phytase activity
was obtained with cowpea meal. Addition of KH2PO4
and Triton X-100 increased enzyme yield (Mandviwala
& Khire 2000). The influence of moisture content, glu-
cose, P, surfactants, and gamma irradiation on phytase
production by A. niger A-98 by SSF on rapeseed meal has
been studied by EI-Batal and Karem (2001). Bogar et al.
(2003) utilized a strain of the same microorganism, as
well as Mucor racemosus and Rhizopus oligosporus for
producing phytase in SSF on several natural feed ingre-
dients: canola meal, cracked corn, soybean meal, and
wheat bran. Mixture of oil cakes and wheat bran was
found to be good SSF support for phytase production
by Mucor racemosus (Roopesh et al., 2006). The extra-
cellular phytase production by A. niger NCIM 563 was
evaluated in the medium containing various agriculture
residues under submerged fermentation conditions and
© 2012 Informa Healthcare USA, Inc.  
Bio-processing of agro-byproducts to animal feed  13
phytase production was affected by inorganic phosphate
content of agriculture residues which ranged from 2.8
to 8 mg/g (Bhavsar et al., 2008). Citrus pulp is an agro-
industrial residue from the citrus-processing industry
with low inorganic phosphorus content applied in ani-
mal feed. A new bioprocess method was developed to
produce and purify a new phytase generated on citrus-
pulp fermentation by Aspergillus niger FS3 (Spier et al.,
2010). Extracellular production of phytase by solid-state
cultures of Malbranchea sulfurea and Aspergillus niveus
was reported using cost-effective medium, such as wheat
bran, corn bran, soybean meal, and cotton-seed meal.
There are reports that phytase obtained from A. niger
has been extensively tested and it has been found to be
nontoxic and safe for use in food and feed (Zyta, 1992).
US Food and Drug Administration have approved gener-
ally–regarded–as–safe (GRAS) petition for use of phytase
in food and phytase has been marketed as a feed additive
in the USA since 1996 (Wodzinski & Ullah, 1996).
Potential of enzymes as feed and food enzymes is
well known in the biotechnological industries due to
their myriad of application. The higher cost of enzymes
becomes one of the significant factors which determine
the expense in animal-feed industries. Global enzyme
demand is increasing day by day due to its strong influ-
ence in the specialty enzyme segment and positive
growth in different industrial applications. The cost and
availability of substrates also play an important role in
the enzyme production industries. Agro-residues are
abundant low-cost materials and are easily available.
Production of enzymes from agro-wastes by SSF will not
only be cost effective but it will also reduce the environ-
ment pollution generated by the agro-wastes which need
to be disposed.
Production of flavor and color as feed additives
Palatability refers to the group of sensations that the food
stimulates in the animal. It is a difficult task to combine
the maximum nutritional value of a portion of food with
maximum desirability for the animal. Animals mainly
rely on the senses of smell and taste to evaluate food.
In combination with flavors, the attractiveness of the
feed can be significantly increased for young animals,
and thus ensure optimum feed intake during the criti-
cal rearing phase. Most of the flavor additives increase
the animal feed intake, while simultaneously promoting
the growth and improving the digestion and absorption
capacity, the food conversion ratio and the palatability
of the unconventional feed.
Flavor is usually the result of the presence, within
complex matrices, of many volatile and nonvolatile
components possessing diverse chemical and physico-
chemical properties. The nonvolatile compounds mainly
contribute to the taste and the volatile ones influence
both taste and aroma. Volatile organic chemicals, such
as flavors and aromas are the sensory principles of many
consumer products and govern their acceptance and
market success (Berger, 2009). Flavors produced using
 14  C. M. Ajila et al.
microorganisms currently compete with those from
traditional agricultural sources. According to Berger
(2009), more than 100 commercial aroma chemicals
are derived using biotechnology either through the
screening for overproducers, the elucidation of meta-
bolic pathways and precursors, or by the application of
conventional bioengineering. Market development has
been particularly dynamic for the flavor enhancer glu-
tamate (Leuchtenberger et al., 2005) which is produced
by the fermentation of sugar sources, such as molasses,
sucrose, or glucose using high-performance strains of
Corynebacterium glutamicum and Escherichia coli.
SSF could be potentially used for the production of fla-
vors (Soccol & Vandenberghe, 2003). Microorganisms, such
as Neurospora sp. (Pastore et al., 1994), Zygosaccharomyces
Critical Reviews in Biotechnology Downloaded from informahealthcare.com by 206.162.180.2 on 03/01/12
rouxii (Sugawara et al., 1994) and Aspergillus sp. (Ito et al.,
1990) have been used for the production of flavors by SSF.
Christen et al. (2000) reported the production of volatile
compounds such as acetaldehyde and 3-methylbutanol
by the edible fungus, Rhizopus oryzae during SSF on tropi-
cal agro-industrial substrates. Soares et al. (2000) reported
the production of strong pineapple aroma when SSF was
carried out using coffee husk as a substrate by Ceratocystis
fimbriata. Alberto de Araujo et al. (2002) studied the pro-
duction of an unsaturated lactone with a strong coconut-
like aroma (6-pentyl-α-pyrone), using sugarcane bagasse.
Several parameters, such as initial substrate pH, addition
For personal use only.
of glucose, cultivation temperature, and initial substrate
moisture and inoculum size were studied during the pro-
duction of fruity aroma compounds by Kluyveromyces
marxianus SSF using cassava bagasse or giant palm bran
as a substrate (Medeiros et al., 2001).
SSF was found to be very suitable for the production
of pyrazynes, which possess a nutty and roasty flavor,
and are used as a food additive for flavoring. Besson et al.
(1997) and Larroche et al. (1999) studied the biosynthesis
of 2,5-dimethylpyrazine and tetramethylpyrazine using
SSF cultures of Bacillus subtilis on soybeans. Conversely,
the production of dairy flavor compounds, such as
butyric acid, lactic acid, and diacetyl in mixed cultures of
Lactobacillus acidophilus and Pediococcus pentosaceus
growing on a semisolid maize-based culture has been
reported (Escamilla-Hurtado et al., 2005).
An alternative route for the production of the natural
food colorants is through the application of biotechno-
logical tools by employing the microorganisms. There is a
growing demand for natural colors due to their presumed
safety. The selection of substrate for SSF process depends
upon several factors, mainly related with cost and avail-
ability. In the SSF process, the solid substrate supplies the
nutrients to the microbial culture growing in it. There is
an increased interest in natural pigments to replace some
currently used synthetic dyes, as the latter has been asso-
ciated with toxic effects in foods. The red pigment of the
fungus, Monascus is widely used in Asia for centuries as a
food colorant (Kim et al., 2002).
Monascus has been cultivated on rice (forming ang-
kak or red rice) for the pigment production. Rice is the
 
best substrate for pigment production, compared to other
typical cereals, tubers, and leguminous plants (Carvalho
et al., 2003). Corn, wheat, and cassava have been found
to be good substrates for the production of bio-pigment.
Even though cassava bagasse gave a low pigment yield,
its low price might compensate for its low yield since it
was an agro-industrial byproduct. Jackfruit-seed powder
has been identified as a potent substrate for edible red
pigment production by SSF using Monascus purpureus
(Babitha et al., 2007). Pigment production by Monascus
purpureus in submerged fermentation by using grape
waste as growth substrate was optimized by employing
factorial design and response surface techniques (Silveira
et al., 2008). There has been also interest in the use of tech-
nologies using microorganisms for the production of color
from food-processing byproducts, such as Rhodotorula,
Cryptococcus, Phatlia, Rhodozyma, Monosaus purpureus,
Bacillus spp.,and Xanthomonas campestris.
There is a growing interest for the development of low-
cost technologies for the production of natural pigments
and flavors due to the high-cost technology for the indus-
trial production of pigments and flavor. The production
of flavors and natural pigments by SSF by using agro-
industrial residues as substrates will be very cost effective
and also will add value to the industry.
The quantitative and qualitative potential of raw mate-
rials which are suitable for the production of animal feed
varies widely between economic classes and regions
throughout the world. There are several benefits derived
from the use of feed from agro-byproducts. The major
benefits are (1) environmental sanitation and sustainable
management of environmental pollution; (2) improve-
ment in livestock health and productive agriculture; (3)
development of secondary rural industries; (4) creation
of new employment; and (5) reduction of the import of
animal feed by utilizing the local agro-wastes.
Conclusion and future aspects
Agricultural- and food-industry residues comprise a
major portion of (approximately over 30%) worldwide
agricultural productivity. These wastes mainly consist
of fruit and vegetable wastes, sugar-industry wastes, etc.
They represent valuable biomass and potential solutions
to the problems of animal nutrition and worldwide sup-
ply of protein and calories if appropriate technologies can
be deployed for their valorization by protein enrichment.
SSF, ensiling and high-solid or slurry process are some of
the major technologies employed for protein enrichment
of agro-wastes. Technologies to be developed for the bet-
ter utilization should consider factors, such as peculiari-
ties of individual wastes and the environment in which
they are generated, reprocessed, and used. In particular,
utilization of such technologies needs to deliver products
that are safe not only for animal feed use, but also from
the perspective of human feeding.
Most of the agro-byproducts and wastes have substan-
tial potential value as a diet ingredient in livestock diet;

this kind of utilization will be economically worthwhile
especially in developing countries where traditional
and other expensive animal feed can be replaced. Agro-
industry should be developed on the basis of fundamen-
tal and applied research in the area of feed science and
processing technology. Research needs to be carried out
under local circumstances for information on processing
equipment, processing conditions, chemical and nutri-
tional diet, and its effects on animal growth performance.
The use of organisms that are generally recognized as
safe (GRAS) for the protein enrichment and reprocessing
of wastes will enhance user confidence.
The utilization of agro-byproducts, which are rich in
nutrients, for the production of animal feed will result in
reduced food costs and lower prices for animal products.
Critical Reviews in Biotechnology Downloaded from informahealthcare.com by 206.162.180.2 on 03/01/12
This will also contribute to self-sufficiency in nutrient
from locally available waste. Systems of feed production
from wastes make possible a vertical, mutually comple-
mented integration of animal production and in turn
solve the problem of waste and environmental pollution.
It will also help to ensure that these products will be of a
real value as an outlet for the main products from which
they have been derived. Utilization of agro-wastes for ani-
mal feed will serve as a sound basis for continued techno-
logical development and management of animal farming.
For personal use only.
Declaration of Interest
The authors are sincerely thankful to the Natural Sciences
and Engineering Research Council of Canada (Discovery
Grant 355254, Canada Research Chair), FQRNT (ENC
125216), MAPAQ (No. 809051) and Inde Initiative 2010
(Ministère de l’Éducation, du Loisir et du Sport) for
financial support. The views or opinions expressed in this
article are those of the authors.
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