1 2 1

1 Department of Biology, American University of Beirut, Beirut, Lebanon; 2 Department of Fisheries and Allied Aquaculture,

Auburn University, Auburn, AL, USA

Redclaw crayfish (Cherax quadricarinatus, von Martens

1868) is a freshwater decapod crustacean with a number of

biological and commercial attributes that make it an excel-

lent aquaculture species. The redclaw aquaculture industry

has been growing rapidly since the mid-1980s in tropical

and subtropical regions of the world. Redclaw aquaculture

is mostly in extensive pond systems, but interest in devel-

oping more intensive systems is increasing. To support

continued intensification, development of high-quality

practical diet formulations and information about redclaw

nutrition requirements are necessary. A number of studies

have determined optimum dietary protein and lipid

requirements for juvenile redclaw. However, there is lim-

ited information on essential amino acid and fatty acid

requirements. Several studies report the presence of various

digestive enzymes that have been linked to the ability of

the species to digest a wide range of dietary components.

Furthermore, as in many other aquaculture species, there

is a need to replace fishmeal with other protein sources. A

number of studies explored the possibility of replacing fish

meal with various terrestrial plant sources of protein and

lipids and showed that redclaw can be offered diets con-

taining low-cost, plant-based ingredients without compro-

mising survival, growth and, to a certain extent,

reproduction. Formulated diets containing less expensive,

plant-based ingredients will contribute to a more profitable

and environmentally sustainable redclaw aquaculture

industry. Finally, there is also a paucity of information on

vitamin and mineral requirements of redclaw and little

information on nutrient requirements of broodstock. For

the redclaw aquaculture industry to thrive, we need to

have a better understanding of nutrient requirements at all

life stages.

KEY WORDS: Cherax quadricarinatus, crayfish, diet, feed,

nutrition, redclaw

Received 13 May 2011; accepted 6 December 2011

Correspondence: I. Patrick Saoud, Department of Biology, American

University of Beirut, Beirut, Lebanon. E-mail: is08@aub.edu.lb

Aquaculture of the Australian redclaw crayfish Cherax quad-

ricarinatus (von Martens 1868) is developing rapidly in tropi-

cal and some temperate regions of the world. Webster et al.

(2002) stated that aquaculture of the species was mainly

restricted to North-Eastern Australia, but redclaw aquacul-

ture has expanded into South-East Asia and Central/South

America and production is no longer restricted to Oceania.

The species grows well when offered diets developed for

other crustaceans, but nutritional requirement data specific

for redclaw have not been determined. As culture methodol-

ogy shifts from extensive and semi-intensive ponds into more

intensive systems and as hatchery production becomes more

common, we will need to develop species-specific feed formu-

lations (Huner et al. 1994; Medley et al. 1994; Webster et al.

1994, 2002; Curtis & Jones 1995). These diets should be less

expensive than traditional shrimp feeds, offer a complete

nutrient profile to the animal, be based on sustainable

sources of raw ingredients and be available wherever the

industry decides to grow. The present manuscript reviews

known nutritional requirements of redclaw crayfish based on

existing literature and the experience of the authors.

In natural ecosystems, crayfish have polytrophic feeding

habits and have been described as predators, omnivores

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ª 2012 Blackwell Publishing Ltd

2012 doi: 10.1111/j.1365-2095.2011.00925.x. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Aquaculture Nutrition

and/or detritivores (Momot et al. 1978; D’Abramo &

Robinson 1989; Jones 1990; Brown 1995a; Momot 1995;

Nystrom 2002; Garza de Yta et al. 2011), consuming a

variety of macrophytes, benthic invertebrates, algae and

detritus (Brown 1995a; Nystrom 2002). Jones (1990) sug-

gested that in general Cherax species are primarily detriti-

vores, a statement supported by the findings of Loya-

Javellana et al. (1993) who reported that C. quadricarinatus

demonstrates an ontogenetic shift from non-selective feed-

ing on decayed plant material or zooplankton to a selective

feeding on decayed plant material. Additionally, Jones

(1995) observed that juvenile C. quadricarinatus grow better

when feeding on fresh zooplankton than when offered for-

mulated flake diets (400 g kg�1 protein) but in both cases

grew better when diets were supplemented with vegetal

material. The feeding behaviour (omnivorous/detritivorous)

of redclaw appears to allow for the incorporation of a

broad range of animal- and plant-based ingredients into

formulations of practical diets for aquaculture (Jones 1990;

Campana-Torres et al. 2005, 2006, 2008; Pavasovic et al.

2007a).

Loya-Javellana et al. (1994) described the ontogeny of red-

claw foregut from embryonic stage to adult, while the

embryonic development of the digestive system of was

described by Meng et al. (2001). The digestive system of

decapod crustaceans, including redclaw, can be divided into

foregut, midgut and hindgut (Ceccaldi 1997; Meng et al.

2001). The foregut comprises the mouth (with associated

mandibles), oesophagus and a large part of the cardiac

stomach where the masticating parts are located. The

oesophagus is a short, straight vertical structure that con-

nects the mouth and the stomach. The cardiac stomach, an

oval like sac, is dorsal in the cephalothorax and leads into

the pyloric stomach (elliptically shaped), situated in a ven-

tro-posterior position in relation to the cardiac stomach.

The hepatopancreas (or midgut gland), a large, bilateral,

multilobate diverticulum of the midgut with a basic unit

called a blind tubule, occupies most of the cephalothoracic

cavity. The hepatopancreas has diverse functions including

synthesis and secretion of digestive enzymes, nutrient

absorption, storage of minerals, lipids and glycogen, and

distribution of stored reserves during the intermoult period

(Brown 1995a; Ceccaldi 1997; Verri et al. 2001). In most

crustaceans, the digestive epithelium of the hepatopancreas

is comprised of at least four different cell types: E, R, F

and B, and in some crustaceans, an M-cell is found (Jacobs

1928; Gibson & Barker 1979; Ceccaldi 1997; Verri et al.

2001). E-cells (embryonic) arise by mitotic division at the

distal tips of the each hepatopancreatic tubule and differen-

tiate giving rise to R-cells and F-cells (Dall & Moriarty

1983; Ceccaldi 1997; Verri et al. 2001). R-cells have

microvilli and also contain lipid droplets and glycogen, and

their primary role is storage (Dall & Moriarty 1983; Cec-

caldi 1997). F-cells (fibrillar cells), similar to R-cells, have

microvilli that might contribute to absorption. These cells

secrete and synthesize digestive enzymes and differentiate

into B-cells (Dall & Moriarty 1983; Ceccaldi 1997). B-cells

(blister cells) are associated with protein synthesis and

enzyme secretion (Verri et al. 2001). Another type of cells

found in some crustaceans is the M-cells (midget cells) that

might be involved in nutrient absorption and storage (Cec-

caldi 1997; Guillaume & Choubert 2001). The midgut, not

lined by chitin, begins at the posterior end of the stomach

and extends throughout the abdomen terminating at the

anus. The hindgut is almost straight and impregnated with

chitin, enlarging posteriorly into the rectum and terminates

at the anus (see Ceccaldi 1997).

Loya-Javellana et al. (1995) measured the effect of ani-

mal size and feeding frequency on the foregut evacuation

rates of redclaw. Results indicated that evacuation rates

did not differ significantly between size groups (medium,

large) nor between feeding frequency groups (fed daily, fed

every second day). However, the model specifications dif-

fered between feeding frequencies, i.e. ingesta was evacu-

ated linearly with time in the crayfish fed daily and

according to a curvilinear pattern in those fed every second

day, implying that crayfish are potentially capable of regu-

lating their digestive processes according to food availabil-

ity. Moreover, the return of appetite in redclaw is rapid;

the average return of appetite increased to >50% of the

satiation meal at 5–10 h postfeeding, when the residuum of

the previous meal was ca. 60% or less. The authors

reported that based on these results, redclaw can resume

feeding before a considerable proportion of an earlier meal

is processed in the foregut, suggesting that the species is

capable of optimizing the frequency of feeding during

active foraging periods.

A variety of digestive enzymes including proteases, lipas-

es and carbohydrases are found in the midgut gland (hepa-

topancreas) and gastric fluid of crayfish (Zwilling &

Neurath 1981; Brown 1995a; Hammer et al. 2000) includ-

ing redclaw (Figueiredo et al. 2001). Digestive enzymes are

synthesized and secreted into the digestive tract by F- and

B-cells in the midgut gland (Ceccaldi 1997; Verri et al.

2001). The presence of a variety of enzymes in juvenile red-

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Aquaculture Nutrition ª 2012 Blackwell Publishing Ltd

claw has been linked to the ability of the species to digest a

wide range of dietary components (Xue et al. 1999; Figuei-

redo et al. 2001; Lopez-Lopez et al. 2003, 2005; Pavasovic

et al. 2007a). This complex digestive enzyme activity is

affected by ontogeny (Figueiredo & Anderson 2003),

moulting (Fernandez et al. 1997; Vega-Villasante et al.

1999; Perera et al. 2008), diet composition (Lopez-Lopez

et al. 2005; Pavasovic et al. 2007a), circadian rhythms,

photoperiod and quality of light, temperature, stage of lar-

val development, changes during vitellogenesis (Ceccaldi

1997), feeding habits and even habitat (Figueiredo &

Anderson 2009).

Proteases, enzymes responsible for hydrolysis of peptide

bonds in protein, are present in the gut of crustaceans in

general. They include trypsin or a trypsin-like serine prote-

ase, astacin, chymotrypsin and exopeptidases [e.g. carboxy-

peptidases (A and B)] and aminopeptidases (New 1976;

Vogt et al. 1989; Brown 1995a; Ceccaldi 1997; Guillaume

1997; Navarrete del Toro et al. 2006; Figueiredo & Ander-

son 2009). However, it is generally accepted that most crus-

taceans lack pepsin and stomach acid (see Brown 1995a;

Guillaume 1997; Navarrete del Toro et al. 2006).

Total protease [two optimal pH peaks: 5.0 and 7.5 (gas-

tric fluid) and 4.0 and 7.0 (midgut gland)], trypsin-like

enzyme (EC 3.4.21.4), chymotrypsin-like enzyme (EC

3.4.21.1), carboxypeptidase A-like enzyme (EC 3.4.12.2),

carboxypeptidase B-like enzyme (EC 3.4.12.3) and low lev-

els of leucine aminopeptidase-like enzyme (EC 3.4.11.1)

(Figueiredo et al. 2001) are all found in the gut of crayfish

but might change in activity and concentration depending

on age and diet. Ontogenetic changes in C. quadricarinatus

cause total proteases, trypsin, leucine aminopeptidase and

carboxypeptidases A and B to exhibit high activity in juve-

niles and to decrease as the species grows (Figueiredo &

Anderson 2003).

Lipases are hydrolases that operate at the interface of

emulsified lipid substances (Vogt 2002). They break down

carboxyl ester bonds of triacylglycerols liberating carbox-

ylic acids and glycerol. Figueiredo et al. (2001) reported

lipase (EC 3.1.1.3) activity only in gastric fluid of adult

C. quadricarinatus, whereas Lopez-Lopez et al. (2003)

observed esterase–lipase activity in the hepatopancreas of

juvenile redclaw.

Although aquatic animals in general are not efficient at

utilizing carbohydrates as energy sources, some of the

omnivorous crustaceans exhibit some carbohydrate diges-

tion capabilities. Thus, some of the major carbohydrases

(amylases, laminarinases, chitinases) are found in the diges-

tive system of many crustaceans (Dall & Moriarty 1983;

Ceccaldi 1997). The activity of some of these carbohydrases

is age dependant (Figueiredo & Anderson 2003) and

change with developmental stages of redclaw. For example,

amylase and laminarinase activities are significantly greater

in large C. quadricarinatus than at other stages, whereas

protease activities decreased as the species grew. The carbo-

hydrases detected in the midgut gland and gastric fluid of

adult C. quadricarinatus also include a-amylase (EC

3.2.1.1), laminarinase (EC 3.2.1.6/EC 3.2.1.19), maltase

(EC 3.2.1.20) and several para-nitrophenyl glycosidases (Fi-

gueiredo et al. 2001). Xylanase activity was also reported

in the digestive system of redclaw crayfish (Xue 1998;

Crawford et al. 2005). The presence of these carbohydrases

would suggest that redclaw should be able to obtain a sub-

stantial amount of their metabolic energy needs from car-

bohydrates, yet research suggests that only a relatively

small portion of their energetic needs are obtained from

carbohydrates (see Pavasovic et al. 2006; Garza de Yta

et al. 2012). Additional work on carbohydrate digestibility

and assimilation by redclaw is warranted before definitive

statements can be made.

Some crustaceans have been reported to possess cellulas-

es (EC 3.2.1.4) (Yokoe & Yasumasu 1964; Kristensen 1972;

Brown 1995a; Xue et al. 1999; Figueiredo & Anderson

2003, 2009). Cellulase activity is also present in all stages

of growth in redclaw (Figueiredo & Anderson 2003), yet

we have no definitive proof that redclaw can use cellulose

nutritively. Enzymatic hydrolysis of cellulose to glucose

generally requires the synergistic action of three distinct

classes of cellulase enzymes: endoglucanases (endo-1,4-b-

glucanases (EC 3.2.1.4) that cleave randomly internal

b-1,4-glucosidic bonds; exoglucanases (exo-1,4-b-glucanases

(EC 3.2.1.91) that cleave the disaccharide cellobiose from

the non-reducing ends of the cellulose chains; and cellobias-

es (b-glucosidases, EC 3.2.1.21) that hydrolyse the cellobi-

ose to glucose (Wood 1985; Walker & Wilson 1991;

Woodward 1991; Beguin & Aubert 1994). Generally, higher

animals do not produce endogenous cellulases, but the

presence of symbiotic microorganisms in their alimentary

tracts produces the necessary enzymes for cellulose diges-

tion (Watanabe & Tokuda 2001).

The occurrence of cellulase in the midgut gland and gas-

tric fluid of redclaw (Byrne et al. 1999; Xue et al. 1999;

Figueiredo et al. 2001; Figueiredo & Anderson 2003;

Crawford et al. 2004; Pavasovic et al. 2006) is very interest-

ing. Cellulose, the principal constituent of most plant cell

walls, is known as the most abundant organic compound

and renewable energy source on earth (Aspinall 1980;

BeMiller 2008). Although the idea of using an abundant

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Aquaculture Nutrition ª 2012 Blackwell Publishing Ltd

and low-cost ingredient in aquafeeds is exciting (Byrne

et al. 1999; Crawford et al. 2004; Pavasovic et al. 2007a),

we believe that technological advances required to make

cellulose a dietary energy source for aquatic organisms are

yet to be described and might never be. Regardless, we will

review current literature on the subject.

The catalytic activity of cellulase in redclaw digestive

tracts is not markedly inhibited by antibiotic treatment,

despite a significant decrease in the gut bacterial popula-

tions (up to 94%), suggesting that the activity is innate in

the crayfish and not in microbial symbionts. Redclaw cellu-

lase enzymes demonstrated broad substrate specificity,

hydrolysing polysaccharides containing b-1,4 and mixed

b-1,4 and b-1,3 glycosidic bonds but with a higher prefer-

ence for soluble substrates (Xue et al. 1999). The occur-

rence and activity of cellulase in C. quadricarinatus is

consistent with the feeding behaviour of redclaw, which

consume significant amounts of plant materials and decom-

posing bacteria, fungi and animals (Byrne et al. 1999; Xue

et al. 1999). Byrne et al. (1999) isolated an endo-1,4-beta-

glucanase cDNA sequence (termed CqEG) from the hepa-

topancreas of redclaw, thus providing one of the first

endogenous cellulase sequences in crustaceans. Crawford

et al. (2004) complemented the study conducted by Byrne

et al. (1999) by presenting the genomic organization of

CqEG. According to the authors, the presence of an endog-

enous multigene glycosyl hydrolase family 9 in redclaw

indicates that partial breakdown of plant cell polysaccha-

rides is a significant evolutionary strategy for the species.

Results of their study suggested the presence of two func-

tional endoglucanase enzymes in redclaw that may be used

to obtain energy (glucose) from soluble cellulose (see also

Xue et al. 1999), a tool to allow access to other nutrients

within plant cells (Beguin & Aubert 1994) or to reduce

digestive viscosity of soluble polysaccharides leached from

plant cell walls (Crawford et al. 2004). Crawford et al.

(2005) reported that C. quadricarinatus has the capacity to

liberate glucose from carboxymethyl cellulose, indicating

that cellulose substrates can be a source of energy for cray-

fish. However, a study conducted by Pavasovic et al.

(2006) indicated that the presence of cellulase (higher activ-

ity in gastric fluid than midgut gland) in the gut of redclaw

is unlikely to hydrolyse a-cellulose into glucose and thus

would not allow for the supply of energy to the species.

Furthermore, the addition of a-cellulose to midgut gland

extracts did not change solution viscosity, suggesting that

insoluble non-starch polysaccharides do not increase visco-

sity of intestinal contents upon digestion, which in

turn would slow the passage of materials through the gut

(Pavasovic et al. 2006). The authors concluded that

although cellulase activity is present in redclaw, there are

no detectable nutritive benefits of including insoluble cellu-

lose (a-cellulose) in diet formulations of the species.

In addition to proteases, lipases and carbohydrases,

endonucleases probably also exist in redclaw. Endonuclease

activity has been reported in the digestive tract of various

other invertebrates including annelids, molluscs, echino-

derms and arthropods (chelicerates, insects and crusta-

ceans) (Yokoe & Yasumasu 1964; see also Watanabe &

Tokuda 2001 and references therein; Linton et al. 2006)

that are also probably members of the arsenal of digestive

enzymes in redclaw guts, but have yet to be isolated.

Information derived from studies on biochemical compo-

sition and digestive enzyme activities on utilization of yolk

during embryonic development may provide some clues of

the nutrient requirements for the embryos and therefore

can be used in understanding nutritional requirements of

brood stock (Yao et al. 2006; Luo et al. 2008a). Luo et al.

(2008a) studied five digestive enzymes (trypsin, pepsin,

lipase, amylase and cellulase) in embryonic redclaw, and

all showed changes in enzymatic activity closely correlated

with morphogenesis, hydrolysing the yolk and providing

construction substances and energy resources for formation

of tissues, organs and various systems. The activities of the

digestive enzymes were controlled by their genes and

expressed sequentially during development. Specific activi-

ties of pepsin and trypsin increased during early stages of

embryonic development, but pepsin activity decreased in

later stages (stage VI), while trypsin remained at high level

of activity (Luo et al. 2008b). Furthermore, chymotrypsin

activity peaked in stage IV and then decreased significantly

during the last stage of embryonic development. Low levels

of lipase activity were also reported during embryonic

development of redclaw (Luo et al. 2008a). Specific activity

of amylase changed in a ‘V’ curve, increasing during later

stages (stage VI). Cellulase activity during embryonic

development in redclaw was relatively low (Luo et al.

2008a).

Research on the nutritional requirements and practical diet

formulations for redclaw increased rapidly as the culture of

the species became established with further advances occur-

ring in the 21st century. Dietary requirements of some

nutrients have been determined for rapidly growing juve-

niles only, with limited information for larger redclaw

approaching market weight or for broodstock. This is

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Aquaculture Nutrition ª 2012 Blackwell Publishing Ltd

probably because most broodstock are collected from

extensively stocked farm ponds where the animals have

access to primary productivity to supplement possible defi-

ciencies in manufactured diets. Currently, diets for the

commercial production of redclaw are based on formula-

tions of other aquatic species, primarily penaeid shrimp

feed but sometimes prawn and fish feed (Cortes-Jacinto

et al. 2003, 2004, 2005; Garcıa-Ulloa et al. 2003; Thomp-

son et al. 2003a,b). Redclaw have the capacity to adapt

their digestive physiology in response to changes in their

nutrient requirement or dietary profile (Pavasovic et al.

2007b) and consequently have been reared on a wide range

of feed formulations. Redclaw diets could potentially be

quite inexpensive to manufacture, considering that formu-

lated diets with 200–300 g kg�1 crude protein and 50–

100 g kg�1 lipids, based primarily on vegetable rather than

animal ingredients, allow for good survival and growth of

the species (Cortes-Jacinto et al. 2004).

Proteins and amino acids are essential nutrients required for

maintenance, growth and reproduction in crustaceans as in

other animals (Guillaume 1997). Protein requirements of

crustaceans are affected by various factors including physio-

logical stage and size, dietary characteristics of protein

quantity and quality (e.g. digestibility), amount of non-

protein energy in the feed, environmental factors (e.g.

temperature) and methodology used for dietary protein

determination (D’Abramo & Robinson 1989; D’Abramo &

Sheen 1994; Guillaume 1997; Thompson et al. 2005, 2006;

Rodrıguez-Gonzalez et al. 2006a). In general, a mixture of

proteins of both animal and plant origin provide better

growth than either alone because the mixture often contains

a complementary blend of amino acids, which are more

likely to meet or exceed the requirements (D’Abramo &

Robinson 1989; Lovell 1998).

Most crayfish exhibit an ontogenetic diet shift where adult

crayfish incorporate greater levels of detritus and plants in

their diet as compared to juvenile crayfish that feed mostly

on invertebrates (Mason 1975; Loya-Javellana et al. 1993;

Lodge & Hill 1994; Momot 1995; Nystrom 2002). Such dif-

ferences in feeding habits between adult and juvenile cray-

fish have been attributed to slower growth of adult crayfish

and therefore lower protein requirements than in faster

growing juveniles (Lodge & Hill 1994).

Several studies have attempted to determine protein

requirements of juvenile and preadult C. quadricarinatus

reared indoors or outdoors (see Table 1). Anson & Rouse

(1996) evaluated growth response and survival of newly

detached (0.01 g) redclaw offered various commercial feeds

(shrimp feed, catfish feed with or without Artemia nauplii

supplement) ranging in protein content from a 320 g kg�1

protein catfish diet to a 400 g kg�1 shrimp diet. The

400 g kg�1 shrimp diet resulted in best growth for the ani-

mals. D’Agaro et al. (2001) evaluated the dietary protein

content (240 g kg�1 and 290 g kg�1; gross energy: 20.0–

20.4 MJ kg�1) on growth performance of juvenile C. quad-

ricarinatus reared in a recirculating system. No significant

differences in growth were reported among treatments,

probably because of protein-sparing effects from other

energy sources. Meade & Watts (1995) offered 0.01 g red-

claw a number of commercially available formulated diets

and found that a 300 g kg�1 crude protein, 100 g kg�1 fat

feed provided best weight gain and survival as compared to

all other treatments. However, the authors note that such

feeds do not provide complete nutritional needs of crayfish.

Jones & Ruscoe (1996a) evaluated growth performance

of juvenile redclaw in glass aquaria offered five formulated

diets (four commercial formulations and one experimental

reference formulation) and one natural diet containing

crude protein ranging from 100–447 g kg�1. Growth was

significantly greater in trials offered diets containing

365 g kg�1 protein (with fish meal as protein source) and

205 g kg�1 crude protein (entirely of non-animal material).

The authors concluded that redclaw does not seem to have

a specific requirement for high levels of proteins and that

they can be successfully cultured on a diet primarily com-

posed of material of plant origin. Similarly, Thompson

et al. (2005) examined the growth performance of juvenile

redclaw offered formulated practical diets containing

increasing percentages of dietary protein (300, 350 and

400 g kg�1). They found that juvenile redclaw can be

offered a 350 g kg�1 protein formulated practical diet with

a combination of plant-protein ingredients if fishmeal is

excluded.

Natural food and forage can also supplement formulated

diets and spare proteins in the prepared feed. Metts et al.

(2007) reported that juvenile redclaw stocked semi-inten-

sively and offered forage at a rate of 500 kg ha�1 month�1

may be able to utilize 130 g kg�1 protein diets. Thompson

et al. (2006) reported that juvenile redclaw offered diets

containing 280 g kg�1 crude protein with or without fish

meal had significantly greater weight gain compared to red-

claw offered 180 g kg�1 crude protein with or without fish

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Aquaculture Nutrition ª 2012 Blackwell Publishing Ltd

Table

1Optimaldietary

protein

level

forredclaw

(Cheraxquadricarinatus)

Reference

Initialsize

(g)

Protein

source

Protein

leve

ls

tested

Optimalprotein

leve

l(g

kg�1ofdiet)

Culture

type

Websteretal.(1994)

0.022

Mixture:Menhadenfish

meal,soyb

eanmeal(SBM),

shrimpheadmealandgroundco

rn

230–5

50

330(reco

mmended)

Aquaria,recirculatingsystem

Keefe

&Rouse

(1999)

0.02

Mixture:Corn,fish

andSB

Ms

230–4

30

430

280(calculated)

Individualco

ntainers,

recirculatingsystem

D’Agaro

etal.(2001)

9.7

Mixture:Fish

andSB

Msandwheatmiddlings

240–2

94

240(suggested)

Tanks,

recirculatingsystem

Hernandezetal.(2001)

0.2

and8.52

Anch

ovy

fish

meal

250–5

00

300–3

50

Tanks,

recirculatingsystem

Manomaitis

(2001)

0.07

Mixture:Menhadenfish

meal(67%),SB

M(46%),shrimp

meal(44%)andwheatflour(11%)

250–4

00

400

Semi-recirculatingsystem

3.22

250–4

60

300(suggested)

Cortes-Jacinto

etal.(2003)

1.08

Mixture:Sa

rdine,sorghum,soyb

ean,redcrab,squid

meals,wheatmealandgrenetine.

200–5

50

310

342(calculated)

Staticexp

erimentalsystem

Cortes-Jacinto

etal.(2004)

21.8:females

Mixture:Sa

rdine,sorghum,soyb

ean,redcrab,squid

meals,wheatmealandgrenetine.

220–4

50

220

Staticexp

erimentaltanks

23.1:males

256(calculated)

Cortes-Jacinto

etal.(2005)

0.71

Mixture:Sa

rdine,sorghum,soyb

ean,squid

meals,wheat

mealandgrenetine

260–3

60

310

Staticexp

erimentaltanks

Cortes-Jacinto

etal.(2009)

1.04

Mixture:Sa

rdine,sorghum,soyb

ean,redcrab,squid

meals,wheatmealandgrenetine

280–4

00

350

Staticexp

erimentaltanks

Dıazetal.(2006)

1–2

RangenandPurina

320–3

50

350

Recirculatingsystem

Rodrıguez-Gonza

lez

etal.(2006a)

23.0

Mixture:Sa

rdine,sorghum,soyb

ean,andsquid

meals,

wheatmealandgrenetine.

220–3

70

320(calculated:300)

Tanks,

staticsystem

Rodrıguez-Gonza

lez

etal.(2009a)

25.5

females

Mixture:Sa

rdine,sorghum,wheat,

squid,redcrabmeals,

soyb

eanpasteandgrenetine

220–4

50

330(reco

mmended)

284–3

55

Tanks,

staticsystem

Thompsonetal.(2004)

4.6

Mixture:Menhadenfish

meal(67%),SB

M(50%),

Brewer’sgrainswithye

ast

(35%),wheatgluten(41%)

andwheatflour(14%)

220–4

20

220(reco

mmended)

Ponds

Thompsonetal.(2005)

1.12

Mixture:Anch

ovy

fish

(65%)meal,SB

M(48%),wheat

flour(12.5%,milo(11.5%),BGY(35%)andwheat

gluten(80%)

300–4

00

300(150gkg�1fish

meal)

350(0

gkg�1fish

meal)

Tanks,

recirculatingsystem

Thompsonetal.(2006)

5.75

Mixture:Menhadenfish

meal(62%),SB

M(48%),

distillers’grainswithsolubles(28%),milo(10%)and

wheatgluten(72%).

180–2

80

280(0

gkg�1fish

meal)

Ponds

Mettsetal.(2007)

6.25

Mixture:Fu

ll-fatSB

M,solvent-extractedSB

M,wheat

middsandwhole

wheat+alfalfahay

130–2

80

130(w

ithorwithout

alfalfahay)

Ponds

Pava

sovicetal.(2007b)

13.9

Fish

meal,gelatin

130–3

20

250

Individualcageswithin

tanks,

recirculatingsystem

Zenteno-Savınetal.(2008)

0.71

SameasCortes-Jacinto

etal.(2003)diets

260–3

60

310

Staticexp

erimentaltanks

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Aquaculture Nutrition ª 2012 Blackwell Publishing Ltd

meal (778% and 799%, respectively). They concluded that

pond-cultured redclaw performed well when offered diets

with 280 g kg�1 protein inclusion even if devoid of fish-

meal.

Hernandez et al. (2001) studied the effect of eighteen iso-

caloric (417.4–422.8 kcal 100 g�1) diets containing six levels

of protein inclusion (250, 300, 350, 400, 450 and 500 g kg�1)

each at lipid levels of 40, 80 and 120 g kg�1, on growth and

survival of hatchling and juvenile redclaw reared under con-

trolled conditions. The authors concluded that diets contain-

ing 300–350 g kg�1 protein (40–80 g kg�1 lipid) result in

best growth performance for both size classes. Manomaitis

(2001) offered juvenile redclaw diets with various protein

inclusion levels (250–400%) for 7 weeks. Final weight, spe-

cific growth rate (SGR) and percentage weight gain of the

juveniles were positively correlated with increasing protein

levels in the diet. However, a second similar trial with larger

juveniles resulted in no effect of dietary protein level on all

test factors. The author concluded that a diet of at least

400 g kg�1 crude protein should be offered to newly released

redclaw, whereas for larger juveniles, a diet containing

300 g kg�1 protein is sufficient.

Cortes-Jacinto et al. (2003) evaluated the response of

juvenile redclaw offered experimental diets containing seven

levels (200, 250, 310, 370, 430, 490 and 550 g kg�1) of

dietary protein and with 18.73–21.45 kJ g�1 gross energy

(protein to energy ratio: 10.7–25.6 mg kJ�1). Results showed

that highest mean weight (9.6 g) and SGR (3.64% day�1)

were achieved by offering a diet containing 310 g kg�1 crude

protein. The optimum dietary protein requirement, calcu-

lated from using a second-order polynomial (y = 1.142 +

0.484 � 0.0071x2, r2 = 0.952), was 342 g kg�1. Similar

results were achieved by a later study conducted by Cortes-

Jacinto et al. (2005) determining the effect of various protein

(260, 310 and 360 g kg�1) and lipid (40, 80 and 120 g kg�1)

levels, with gross energy content of 17.5–19.4 kJ g�1, on

growth of juvenile C. quadricarinatus. Best growth was

observed when using dietary protein inclusion of 310 g kg�1

(80 g kg�1 crude lipid) with gross dietary energy content of

19.69 kJ g�1. Similar results were observed by Dıaz et al.

(2006).

Dietary protein also appears to have an effect on redclaw

health. Zenteno-Savın et al. (2008) reported that diets con-

taining 310 g kg�1 crude protein satisfy nutritional require-

ments for optimal growth, while preventing diet-induced

oxidative stress and protecting the integrity of the immune

response in juvenile redclaw. Similarly, Cortes-Jacinto et al.

(2009) reported that a 350 g kg�1 protein diet stimulates

antioxidant response of superoxide dismutase (SOD) (SOD

is a cytosolic enzyme specific for scavenging superoxide

radicals and is involved in protective mechanisms within

injured tissues following oxidative processes and phagocy-

tosis) of juvenile redclaw.

For earthen pond culture, it is not necessary to supply

high dietary protein because redclaw supposedly obtain a

substantial proportion of their nutrient requirements from

natural food materials in the pond (Jones 1990; Jones &

Ruscoe 1996b). Jones & Ruscoe (1996b) stocked juvenile

redclaw in cages in a pond and offered diets containing

crude protein ranging from 100 to 447 g kg�1. Although

crayfish offered a reference crayfish diet (205 g kg�1 crude

protein) grew better than crayfish offered all other diets,

the authors suggested that the crayfish did not have a

direct use of the feed offered but obtained the bulk of their

nutrition from natural productivity of the pond benthos. In

a similar experiment, Thompson et al. (2004) found that

220 g kg�1 dietary protein was sufficient for redclaw cul-

ture.

In other experiments, Pavasovic et al. (2007b) reported

maximum growth of subadult redclaw offered diets con-

taining 250 g kg�1 crude protein with a strong positive cor-

relation between dietary protein and protein content in the

tail. However, other researchers did not observe a signifi-

cant effect of dietary protein on percentage protein in red-

claw tail muscle or even total body protein (Muzinic et al.

2004; Thompson et al. 2004).

A summary of the literature thus suggests that diets with

250 g kg�1 or greater protein inclusion are suitable for red-

claw growout in ponds with natural productivity. Diets

with 350 g kg�1 protein inclusion or greater are recom-

mended for redclaw grown in closed recirculation systems.

All diets should have a gross energy content of 18 kJ g�1,

minimum. These suggestions are supported by Cortes-Jac-

into et al. (2004) who propose a minimum protein inclusion

in redclaw diets of 220 g kg�1 with 15.21 kJ g�1 of digest-

ible energy.

No discussion of aquatic animal nutrition is complete

without mentioning broodstock diets. Broodstock nutrition

is of high importance for successful reproduction and egg

quality; adequate nutrients and energy in broodstock diets

are necessary for the onset of gonadal maturation, because

maternal nutrient intake during ovarian development is

critical and influences the composition of ovaries and the

nutritional status of eggs. Crustacean embryos rely exclu-

sively on the nutrients and energy supplied by the egg

(yolk) (Harrison 1997). In decapod crustaceans, protein is

a structural, functional and energy constituent of tissues

and plays an important role in spawning, fertilization and

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Aquaculture Nutrition ª 2012 Blackwell Publishing Ltd

normal development of embryos (Harrison 1990; Wouters

et al. 2001; Garcıa-Guerrero et al. 2003; Rodrıguez-Gonza-

lez et al. 2006a). Asgari (2004) reported that based on

spawning rate, fecundity, hatchability and egg size, a diet

containing 400–450 g kg�1 crude protein and 16.72 kJ g�1

energy is optimal for redclaw crayfish broodstock, yet

Rodrıguez-Gonzalez et al. (2006a) tested diets with lesser

protein inclusion and found no differences in survival, final

weight and fecundity of female broodstock. However,

regression analysis indicated that maximum spawning was

from females offered a 300 g kg�1 crude protein diet and

dietary protein levels of 320 g kg�1 had a significant effect

on egg quality but not on biochemical composition of the

eggs. Such findings were recently corroborated by Li et al.

(2010) who found that using diets with higher protein con-

tent improves redclaw female spawning, especially when

gonadosomatic index is >1.6. Rodrıguez-Gonzalez et al.

(2009a) separated maturation from gonadal development as

they relate to female broodstock diets and found that diets

with 220–450 g kg�1 crude protein result in maturation of

female redclaw, but a dietary protein range from 284 to

355 g kg�1 improved gonadal development and resulted in

more protein production in the hepatopancreas. In previ-

ous work, Rodrıguez-Gonzalez et al. (2006b) had observed

that external sources of protein and energy were vital for

nutrient accumulation in the gonad. Additionally, protein

contents in the gonad were correlated with gonadosomatic

index; at mature stages, higher protein concentration was

observed. These gonadal proteins were a result of an active

mobilization of energy reserves from exogenous sources,

incorporated into the oocytes by endocytosis (Abdu et al.

2000). Based on current knowledge, we suggest that brood-

stock females be offered diets with 350 g kg�1 protein and

a minimum of 18 kJ g�1 gross energy, a part of which

comes from fish oil to supply the necessary omega-3 HU-

FAs.

Determination of the exact amino acid requirements in

crustaceans is difficult (Shiau 1998), and this is probably

the reason for the paucity of reports on the specific amino

acid requirements of redclaw. In general, the essential

amino acid requirements for most crustaceans include argi-

nine, histidine, isoleucine, leucine, lysine, methionine, phen-

ylalanine, threonine, tryptophan and valine (D’Abramo &

Robinson 1989; Brown 1995b; Guillaume 1997) plus aspar-

agine for crayfish (Brown 1995b). Tyrosine and cysteine are

considered semi-essential in the diet as they potentially

spare the requirement of phenylalanine and methionine,

respectively (Guillaume 1997). There is a significant corre-

lation between the dietary amino acid requirements of a

species and the pattern of amino acids in whole body tissue

(Cowey & Tacon 1983; Wilson & Poe 1985). Consequently,

dietary amino acid requirements of growing animals are

often assumed to be similar to the amino acid composition

of the tissue proteins formed during growth. Mitchell

(1950) suggested that an animal’s amino acid requirements

might first be deduced from the amino acid composition of

its tissues. However, our experience suggests that when

using body composition as reference of requirement, one

would overestimate dietary requirement of essential amino

acids and underestimate requirement of other protein com-

ponents.

Muzinic et al. (2004) evaluated the amino acid composi-

tion of practical diets containing various levels of soybean

meal (SBM) and brewer’s grains with yeast as replacements

for fish meal, and results suggested that the amino acid levels

in a 400 g kg�1 crude protein diet were adequate for good

growth and survival of juvenile redclaw crayfish whichever

protein source was used. Similarly, Thompson et al. (2005)

noted that a complementary blend of SBM and other plant-

protein sources used to replace FM in a 350 g kg�1 protein

diet appeared to provide sufficient levels of essential amino

acids to meet requirements of redclaw. In pond-cultured red-

claw, diets containing 280 g kg�1 crude protein with or

without fish meal may sufficiently satisfy the requirements of

essential amino acids of male and female redclaw (Thomp-

son et al. 2006) probably because natural productivity sup-

plements the formulated feeds being offered. Consequently,

and based on their response to diets without fishmeal, one

may assume that methionine and lysine requirements of red-

claw are relatively low. Such assumptions are yet to be

empirically tested.

Knowledge of energetic utilization of farmed organisms is

necessary for the development of cost-effective diets.

Energy from non-protein sources (lipids, carbohydrates)

relative to protein levels must be supplied into diets in suf-

ficient amounts to insure that protein is used for tissue syn-

thesis as protein is considered the most expensive major

component of crustacean diets (D’Abramo & Robinson

1989; Cuzon & Guillaume et al. 1997; Cho et al. 2005). If

the non-protein energy to protein ratio is insufficient, die-

tary protein may be catabolized and used as an energy

source to satisfy maintenance before somatic growth. Con-

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Aquaculture Nutrition ª 2012 Blackwell Publishing Ltd

versely, if dietary energy to protein ratio is in excess, feed

consumption may be reduced, resulting in a decrease in

protein intake and other essential nutrients required for

maximum growth. Excessively high ratios of energy to

nutrients can also lead to deposition of large amounts of

body fat (Cuzon & Guillaume 1997).

A review of various published results concerning dietary

protein to energy requirements of redclaw suggests that

optimal growth is obtained when the animals are offered

feed with a protein to energy content between 16 and

20 mg kJ�1 and a crude protein content between 310 and

350 g kg�1 of the diet by weight (D’Agaro et al. 2001;

Cortes-Jacinto et al. 2003, 2005, 2009). D’Agaro et al.

(2001) found no significant differences in growth of redclaw

when offered diets containing protein to energy ratio of 50

and 60 mg kcal�1 (240 and 294 g kg�1 protein, respec-

tively) and attributed their results to the protein-sparing

capacity of the energy in the diets. This protein-sparing

effect was also observed by Hernandez et al. (2001). Values

for protein to energy ratio for optimal reproductive activity

and gonadal development in redclaw were 18 ± 2 mg kJ�1

(Rodrıguez-Gonzalez et al. 2011) and 17.16 mg kJ�1

(Rodrıguez-Gonzalez et al. 2006a), respectively, within the

range observed as necessary for juvenile growth.

Dietary lipids play an important role in crustacean nutri-

tion as they provide energy and essential fatty acids

(EFAs), sterols, phospholipids and fat-soluble vitamins nec-

essary for proper functioning of physiological processes

and maintenance of biological structure and function of

cell membranes (D’Abramo & Robinson 1989; Sargent

et al. 1989; D’Abramo 1997; Teshima 1997). Lipid used as

energy source can also spare dietary proteins and reduce

nitrogenous waste production (D’Abramo & Robinson

1989; Lim & Sessa 1995; Cho & Bureau 2001). However,

high dietary lipid levels can cause significant reductions in

growth rate, feed consumption and also might reduce the

utilization of other nutrients resulting in reduced growth

(D’Abramo 1997). Additionally, an increase in dietary lipid

levels was linked to increases in the lipid content of midgut

glands (hepatopancreas) (D’Abramo 1997).

In general, nutritional studies with crustaceans indicate

that lipid content of formulated diets should range between

50 and 80 g kg�1 of feed by weight to ensure optimal

growth and survival (D’Abramo 1997). The lipid level

required for optimal growth is influenced by several factors

including quality and quantity of protein, availability,

quantity and quality of other sources of energy and ade-

quate provision of EFAs (D’Abramo 1997) as well as the

ability of the organism to digest carbohydrates and use glu-

cose in its metabolism. Lipids are often supplemented in

excess of minimal requirements to spare protein for somatic

growth. Such a protein-sparing effect of lipids was reported

in hatchling and juvenile redclaw offered diets containing

40–80 g kg�1 lipid (300–350 g kg�1 protein) (Hernandez

et al. 2001), suggesting that this range of lipid inclusion to

redclaw diets is suitable.

A few studies investigated dietary lipid requirements of

redclaw under laboratory conditions (Hernandez et al. 2001;

Cortes-Jacinto et al. 2005; Zenteno-Savın et al. 2008), and

all seem to agree that a diet containing 80 g kg�1 dietary

lipid with approximately 300 g kg�1 protein and gross

energy 17.5–19.1 kJ g�1 is suitable for good growth perfor-

mance of juvenile C. quadricarinatus while preventing diet-

induced oxidative stress and protecting the integrity of the

immune function.

Although lipids are necessary in redclaw diets, it appears

that natural productivity can replace dietary lipids to a cer-

tain extent. Hernandez-Vergara et al. (2003) evaluated the

effect of different dietary lipid levels (42, 82 and

123 g kg�1) on growth, survival and proximate composi-

tion of juvenile redclaw reared semi-intensively in outdoors

tanks and observed no effect of treatment on the various

parameters. Accordingly, it seems that in the presence of

some natural productivity, a diet containing 42 g kg�1 die-

tary lipid (17.58 kJ g�1, 300 g kg�1 crude protein) is suffi-

cient for growth and survival of juvenile redclaw.

Differences in lipid metabolic routes between sexes where

females have higher carcass lipid content than males are

often reported (e.g. Hernandez-Vergara et al. 2003). This is

generally attributed to storage of lipids for ova develop-

ment or vitellogenesis. Yet, studies on developing adequate

diets for maturation of broodstock redclaw are rare (Rodrı-

guez-Gonzalez et al. 2006a,b, 2009a,b). Considering that

lipids are the main energy sources during ontogeny of crus-

taceans and also structural components of cell membranes

(Holland 1978; Harrison 1997), lack of research on the sub-

ject seems surprising. However, when one considers the

ease of collecting egg-bearing females from ponds, one

understands the lack of interest in broodstock maintenance.

Nonetheless, as the industry grows and biosecurity issues

become more important and infectious diseases appear,

indoor closed system hatcheries will become necessary and

with them special broodstock diets.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Aquaculture Nutrition ª 2012 Blackwell Publishing Ltd

According to Rodrıguez-Gonzalez et al. (2006b), the

lipid requirements for the developing gonad of female red-

claw mainly originates from the diet. Rodrıguez-Gonzalez

et al. (2009b) studied the effect of dietary lipid levels (40,

80 and 120 g kg�1) on female redclaw crayfish and their

eggs. Results indicated no significant differences in survival,

final weight or fecundity. However, dietary lipid content

influenced size and weight of eggs, with greatest egg weight

obtained from females offered the 87 g kg�1 lipid diet. In a

similar study on the effects of dietary lipids on female red-

claw reproduction, Li et al. (2010) observed a significant

correlation between lipid transportation in the hepatopan-

creas and the ovaries, but it appeared that the lipid reserves

in the hepatopancreas could not meet the requirements of

ovaries. The authors concluded that the lipid requirements

of gonads come only partly from the diet.

Polyunsaturated fatty acids (PUFA) of the C18 series (lino-

lenic (18:3n-3) and linoleic (18:2n-6) acids) and n-3 and n-6

highly unsaturated fatty acids [eicosapentaenoic acid

(EPA), docosahexaenoic acid (DHA) and arachidonic acid

(ArA)] are considered essential in crustacean diets (see

D’Abramo & Robinson 1989; D’Abramo 1997; Venero

et al. 2008). No studies on specific EFA requirements of

redclaw were found; however, a few studies evaluated diets

containing various levels of fatty acids on growth perfor-

mance of the species.

Thompson et al. (2003a) reported that a mixture of 5%

cod liver oil and 1% corn oil added to the diets probably

met the EFA requirements of juvenile redclaw. The blend

of these oils provides a mix of PUFA such as linoleic

(18:2n-6), linolenic (18:3n-3), oleic (18:1n-9) acids and

highly unsaturated fatty acids such as eicosapentaenoic

(20:5n-3) and docosahexaenoic (22:6n-3) acids, that is suffi-

cient for redclaw survival and growth. Similarly, Thompson

et al. (2003b) evaluated practical diets with and without

supplemental lecithin and/or cholesterol offered to juvenile

redclaw. The authors indicated that diets with 0 g kg�1

supplemental lecithin and/or cholesterol contained a combi-

nation of PUFA and HUFA in the diet, which satisfied the

EFA requirements of juvenile redclaw. Thompson et al.

(2010) examined the effect of different sources of lipids (lin-

seed oil, canola oil, corn oil, beef tallow or menhaden oil)

that differ in fatty acid profile on growth response of juve-

nile redclaws. Results showed that whole-body fatty acid

composition of redclaw differed among animals offered the

various lipid sources, generally reflecting the fatty acid

composition of the diets. Plant oils rich in a-linolenic acid

(18:3n-3), linoleic acid (18:2n-6) and oleic acid (18:1n-9)

perform as well as menhaden oil containing high levels of

n-3 HUFA when offered to juvenile redclaw reared indoors

and lacking natural productivity. The authors concluded

that redclaw can be fed diets containing plant-based oils

with high levels of 18-carbon unsaturated fatty acids. Muzi-

nic et al. (2004) reported that practical diets containing

various levels of SBM and brewer’s grains with yeast, as

replacements for fish meal, have both n-6 and n-3 highly

unsaturated fatty acids such as linoleic (18:2n-6), eicosapen-

taenoic (20:5n-3) and docosahexaenoic (22:6n-3) acids that

may satisfy the EFA requirements of juvenile redclaw.

The fatty acid profile during early embryonic develop-

ment of redclaw shows that the major fatty acids, oleic/

vaccenic (18:1), palmitic (16:0), linoleic (18:2n-6) and pal-

mitoleic (16:1n-7) remain major during later developmental

stages and are required in larger quantities than other fatty

acids (Alimon et al. 2003). Monounsaturated fatty acids

constituted the major moiety of the fatty acid profile, and

the PUFA were dominated by linoleic (n-6) series (low n-3

to n-6 ratio) (Alimon et al. 2003). Luo et al. (2008a)

reported that the predominant fatty acids of both neutral

and polar lipids of redclaw during embryonic development

were C16:0, C18:0, C18:1n-9 and C18:3n-3.

Saturated fatty acids (16:0 and 18:0) and monosaturated

fatty acids (16:1n-7 and 18:1n-9) are generally used for

energetic purposes, whereas PUFA (20:5n-3 and 22:6n-3)

are important as structural components of cell membranes

and in the development of the central nervous system (Luo

et al. 2008a). However, even during vitellogenesis, there are

high proportions of monounsaturated fatty acids in the

ovaries and hepatopancreas, suggesting their use as major

sources of energy (Li et al. 2010). Such information would

suggest that broodstock diets could be formulated to con-

tain more vegetable oils to be used for energy during vitel-

logenesis without compromising development of eggs,

which require some n-3 HUFAs found in expensive but

necessary fish oils.

Phospholipids are added to the diet of crustaceans for vari-

ous reasons such as a source of energy; a major component

of cell membranes; emulsification of lipid aggregates during

digestion and absorption; and because they play a major

role in lipid transportation in the haemolymph (Coutteau

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Aquaculture Nutrition ª 2012 Blackwell Publishing Ltd

et al. 1997; Teshima 1997). Thompson et al. (2003a) evalu-

ated the effect of different levels of supplemental soybean

lecithin on growth and survival of juvenile redclaw crayfish

and found no effect. The authors suggested that diets con-

taining enough lipids and 5 g kg�1 choline chloride by

weight may be sufficient for redclaw crayfish growth. Simi-

lar observations were made by Thompson et al. (2003b) in

a study examining growth performance of juvenile redclaw

reared over an 8-week period and offered diets with and

without supplemental lecithin and/or cholesterol.

It is generally accepted that crustaceans are unable to syn-

thesize sterols de novo and require an exogenous dietary

source for growth, development and/or survival (see Teshi-

ma 1997; Kanazawa 2001). Cholesterol is the major essen-

tial sterol in crustacean nutrition with an important role as

a cell constituent, metabolic precursor of steroid hormones

and moulting hormones (see Brown 1995b; Teshima 1997;

Shiau 1998). Dietary cholesterol requirement for optimal

growth performance in various crustacean species ranges

from 1.2 to 20 g kg�1 of the dry weight of a diet (Teshima

1997; Kanazawa 2001).

Studies on cholesterol requirements of C. quadricarinatus

are few. Hernandez et al. (2004) evaluated the effect of die-

tary cholesterol on growth and survival of redclaw and

observed no significant differences among treatments of

various cholesterol supplementations but noticed slight

growth increase in redclaw offered a diet with 5 g kg�1

cholesterol inclusion. A growth study by Thompson et al.

(2003b) whereby they offered C. quadricarinatus juveniles

(0.2 g) practical diets with or without supplemental lecithin

and/or cholesterol showed no significant differences in

weight gain among treatments. The authors interpreted

their results to suggest that redclaw could be farmed using

feeds less expensive than traditional marine shrimp feeds,

currently used in redclaw culture.

Carotenoids, a family of over 600 natural lipid-soluble pig-

ments, are the most universally widespread (e.g. bacteria,

algae, plants and animals) and structurally diverse pigment-

ing agents. They are synthesized through the isoprenoid

pathway, which also produces diverse compounds such as

EFAs, steroids, sterols and vitamins A, D, E and K (see

Meyers & Latscha 1997; Linan-Cabello et al. 2002). Crusta-

ceans are unable to synthesize carotenoids de novo and must

obtain them from the diet (Meyers & Latscha 1997). Yet,

most crustaceans contain and utilize carotenoid pigments;

mainly in the carapace, eyes, blood eggs, midgut gland and

ovaries (Sagi et al. 1995; Meyers & Latscha 1997). Func-

tions of carotenoids include pigmentation, sources of provi-

tamin A, antioxidants, positive effects on development,

growth, maturation, reproduction and enhancement of

immunity (see Meyers & Latscha 1997). The most common

pigments, derived from diets or from metabolic transforma-

tion of precursor carotenoids, are astaxanthin, b-carotene,

echinenone and canthaxanthin (Meyers & Latscha 1997).

Astaxanthin has been described as the most frequent end

product of carotenoid metabolism in crustacean (Meyers &

Latscha 1997).

Rouse & Rash (1999) reported that astaxanthin added to

diets offered to juvenile redclaw resulted in an increase in

survival and growth by 20%. However, in a study by Har-

paz et al. (1998) where the effect of three carotenoid

sources (dried alga cells prepared from Dunaliella salina

(source of b-carotene), synthetic astaxanthin and alfalfa

meal) on growth and pigmentation of juvenile C. quadrica-

rinatus was evaluated, no significant differences were found

on growth and survival. Redclaws receiving carotenoid-

enriched diets exhibited better pigmentation than those

receiving carotenoid-free diets. The authors suggested how-

ever that adding alfalfa meal and artificial astaxanthin to

redclaw diets produce desired body coloration.

Carotenoids play an essential role before and after gona-

dal maturation processes (reviewed by Linan-Cabello et al.

2002). Sagi et al. (1995) noted that target tissues for carot-

enoid accumulation in C. quadricarinatus are the ovary and

cuticle. In a study by Linan-Cabello et al. (2004), the effects

of carotenoid (b-carotene and astaxanthin) and vitamin A

injections were correlated with the ontogenic development

of oocytes in female redclaw. Their results showed that

retinol palmitate had the greatest inductive effect on the

primary vitellogenic phase and on the indicators of onto-

genic oocyte development. Accordingly, we deduce that

carotenoids and retinols are essential nutritive additives

during gonadic maturation of redclaw and help giving

adults a coloration that would help in marketing. Addi-

tional work should evaluate the effects of dietary carot-

enoid supplementation on immune responses and general

physiology of redclaw.

Carbohydrates are generally an inexpensive source of

energy for crustacean feeds (Shiau 1997). Although carbo-

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hydrates are not considered essential nutrients, they are

incorporated into feeds to reduce costs, for their binding

activity during feed manufacturing and possibly for

protein-sparing effects (D’Abramo & Robinson 1989; Guil-

laume & Choubert 2001; Wouters et al. 2001; Pillay &

Kutty 2005). Additionally, carbohydrates play several roles

in crustacean metabolism including glycogen storage, chitin

synthesis and the formation of steroids and fatty acids

(Parvathy 1971; Dall et al. 1990; Ali 1993; Sanchez-Paz

et al. 2006).

Crustaceans are able to utilize more complex carbohy-

drates (e.g. starch, chitin) than simple sugars (e.g. glucose)

(Shiau 1997), yet it is generally accepted that freshwater

crayfish have poor polysaccharide digestibility and are unli-

kely to obtain substantial nutrition from fibrous material

(Shiau 1997; Pavasovic et al. 2006). However, dietary

fibres, water-soluble and water-insoluble polysaccharides

delay stomach emptying by increasing viscosity of the diet

(see Shiau 1997 and references therein), thus assisting in

digestion.

Some polysaccharides such as carboxymethyl cellulose

cannot be digested even though some cellulase activity was

detected in the gut of redclaw. Pavasovic et al. (2006) dem-

onstrated cellulase activity in redclaw digestive systems, yet

observed no detectable nutritive benefits of including insolu-

ble cellulose (a-cellulose) in diet formulations for the species.

Dietary inclusions of a-cellulose above 120 g kg�1 signifi-

cantly reduced survival rate, feed conversion efficiencies and

general growth performance of redclaw. Jones & Ruscoe

(1996c) assessed growth performance of redclaw maintained

in earthen ponds and offered five diets containing various

carbohydrate sources (maize, wheat, rice, sorghum, lupin

and barley). No significant differences in survival and growth

were observed among treatments. This suggests that the

source of carbohydrate is not of particular importance in

redclaw nutrition. In a recent experiment in Mexico, we

assessed the effect of stargrass hay supplementation to for-

mulated feed diets on redclaw growth and found no nutritive

value beyond what the animal obtains from formulated feed.

Dietary carbohydrates also do not appear to affect or

improve gonadal maturation (Rodrıguez-Gonzalez et al.

2006b). Nevertheless, carbohydrates will always constitute a

good portion of formulated redclaw diets because they are

an inexpensive source of energy and filler.

Vitamins and minerals are essential micronutrients neces-

sary for normal life processes in crustaceans. Deficiencies in

vitamins and/or minerals lead to slower growth, negatively

affect reproduction and/or eventual mortality in crusta-

ceans (Conklin 1997; Davis & Lawrance 1997). Informa-

tion on specific vitamin and mineral requirements of

redclaw is scarce to non-existent. It is assumed that vitamin

and mineral requirements are similar to those of other crus-

taceans (D’Abramo & Robinson 1989). However, gonadal

maturation was shown to be affected by vitamin levels.

Linan-Cabello et al. (2004) reported that retinol has a sig-

nificant effect in oocyte maturation of C. quadricarinatus

and is an essential nutritive additive for gonadal matura-

tion. Luo et al. (2004) found that excessive vitamin E

affected reproduction of C. quadricarinatus and that opti-

mal content of vitamin E was 192 mg kg�1. Additionally,

the authors speculated that vitamin E could protect C20:

5n-3 and C22: 6n-3, necessary for the development of the

nervous system, from oxidizing and improved the accumu-

lation of important amino acids and fatty acids in fertilized

eggs.

The nutritive value of a feed ingredient is based on its

chemical composition and on an animal’s capacity to

digest, absorb and utilize it. Digestibility is the quantity of

the nutrient or energy in the ingested feedstuff that is not

excreted in the faeces (NRC 1993; Lee & Lawrence 1997;

Guillaume & Choubert 2001). In general, freshwater crusta-

ceans have higher apparent digestibility efficiency (ADE)

and apparent crude protein digestibility (ACPD) values for

high carbohydrate feeds than marine crustaceans, and both

have high ACPD values for animal meals and purified pro-

teins (Lee & Lawrence 1997).

Campana-Torres et al. (2005) evaluated dry matter and

protein digestibility of juvenile C. quadricarinatus offered

diets supplemented with 150 g kg�1 of three plant-derived

(soy paste, textured wheat and sorghum meal) and four

animal-derived (two sardine meals, squid meal and red crab

meal) ingredients. They found that plant-derived ingredi-

ents and corresponding diets had better digestibility than

animal ingredients (see Table 2). In a subsequent study,

Campana-Torres et al. (2006) reported that mean carbohy-

drate and lipid digestibilities of vegetal ingredients and cor-

responding diets were better than carbohydrate and lipid

digestibilities of animal ingredients although some of the

animal ingredients (e.g. red crab) had high lipid digestibility

(see Table 2). The authors concluded that redclaw are able

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Aquaculture Nutrition ª 2012 Blackwell Publishing Ltd

to efficiently consume diets containing a variety of plant-

and animal-derived ingredients, with better digestion effi-

ciency of plant-derived ingredients (see also Campana-Tor-

res et al. 2008). Similarly, Pavasovic et al. (2007a)

evaluated the potential use of several ingredients (fish meal,

meat and bone meal, poultry meal, SBM, canola meal,

lupin meal and brewer’s yeast) in dietary formulations for

adult redclaw. Results showed that apparent digestibility of

dry matter, crude protein and gross energy was better for

SBM diets than diets containing meat and bone meal

(Table 2). Accordingly, it seems that redclaw has the

capacity to utilize a broad range of dietary ingredients

including animal, single cell and plant matter in their diet.

However, protein digestibility seems to be affected by other

dietary ingredients. Supplementation of diets with either

300 g kg�1 a-cellulose or Fuller’s earth significantly reduces

apparent dry matter digestibility (ADMD) and apparent

protein digestibility (APD) of redclaw (Pavasovic et al.

2006).

Redclaw have the ability to modify their digestive

enzyme secretions in response to different ingredients in the

diet over time (Lopez-Lopez et al. 2005; Pavasovic et al.

2007a). Therefore, as they age and their food changes, they

adapt to the new dietary sources, particularly starches

(Lopez-Lopez et al. 2005). However, digestibility of nutri-

ents is slightly better in juveniles than in preadults (Cam-

pana-Torres 2001; Campana-Torres et al. 2006, 2008). This

deterioration of digestibility with age might be because of

faster metabolism in early stages of development (Guil-

laume 1997).

Diets constitute a major expenditure in aquaculture. Fish

meal and other marine meals (krill, shrimp, squid and scal-

lop meals) are used as protein sources in many finfish and

crustacean diets, as they are considered excellent sources of

high-quality proteins, highly unsaturated fatty acids, vita-

mins, minerals and attractants (Tacon & Akiyama 1997;

Webster et al. 2008). Fish oils have been used as a dietary

lipid source in commercial aquafeeds because of their ready

availability, fair price and abundance of EFAs (Turchini

et al. 2009). However, owing to aquaculture expansion,

competition from other agricultural sectors, uncertainty in

long-term availability (e.g. overfishing), yearly fluctuations

in supply, quality and potential price variation, there has

been considerable interest in partial or total replacement of

fish meal and other marine meals and fish oil with less

expensive plant-protein meals and oils (see Naylor et al.

2000, 2009; Venero et al. 2008; Webster et al. 2008).

Fishmeal replacement Few studies have evaluated the

replacement of fish meal in redclaw diets. SBMreplacement

of fishmeal at various levels resulted in marginal growth

differences among redclaw juveniles, but organisms offered

a fishmeal-based diet had better growth and more frequent

moulting than those offered graded levels of SBM (Garcıa-

Ulloa et al. 2003; Muzinic et al. 2004). Similarly, Gutierrez

& Rodrıguez (2010) examined the effect of protein source

(SBM) on growth of juvenile C. quadricarinatus reared in

Table 2 Apparent dry matter digestibility (ADMD) coefficients for protein (APD), carbohydrates (ACD), lipids (ALD) and gross energy

(AGED) of the various nutrient sources for redclaw (Cherax quadricarinatus)

Feedstuff % ADMD % APD % ACD % ALD % AGED References

Sardine meal (580 g kg�1 crude protein) 62.6 47.4 18.7/41.13 42.0/60.63 – 1, 2, 3

Sardine meal (670 g kg�1 crude protein) 83.2 72.4 28.0/12.33 83.5/84.53 – 1, 2, 3

Red crab meal 79.0 53.8 32.6/34.13 92.1/97.53 – 1, 2, 3

Squid meal 80.1 70.8 18.6/15.63 84.6/60.63 – 1, 2, 3

Textured wheat 88.9 90.5 87.5/83.93 95.0/96.43 – 1, 2, 3

Soy paste 90.8 91.8 88.5/81.63 93.6/95.13 – 1, 2, 3

Sorghum 90.8 89.6 94.4/93.63 85.6/77.33 – 1, 2, 3

Fish meal 80.2 89.8 – – 86.9 4

Meat and bone meal (MBM) 65.3 83.4 – – 74.5 4

Poultry meal (PM) 77.9 87.2 – – 84.9 4

Soybean meal (SBM) 89.3 94.6 – – 93.8 4

Canola meal (CM) 74.5 91.0 – – 80.9 4

Lupin meal (LM) 83.8 94.6 – – 89.6 4

Brewer’s yeast (BY) 85.6 92.6 – – 86.6 4

1 Campana-Torres et al. 2005 (using juveniles: 3.6 ± 1.3 g).2 Campana-Torres et al. 2006 (using juveniles: 3.62 ± 1.3 g).3 Campana-Torres et al. 2006 (using preadults: 10 ± 0.8 g).4 Pavasovic et al. 2007a (using adults: 94.5 ± 3.5 g).

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Aquaculture Nutrition ª 2012 Blackwell Publishing Ltd

individual containers. Based on their results, best growth

was obtained with a mixture of 50% fish meal and 50%

SBM.

Saoud et al. (2008) evaluated the response of juvenile

redclaw offered six practical diets (260 g kg�1 crude pro-

tein, 70 g kg�1 crude lipid) replacing fishmeal with poultry

by-product meal at various inclusion levels. Yuniarti et al.

(2011) performed a similar experiment but substituted

golden apple snail (Pomacea canaculata) meal for fishmeal.

No significant differences in survival and growth were

detected among treatments in both experiments. The

authors concluded that poultry by-product meal and

golden apple snail meal are potential candidates for fish

meal replacement in redclaw diets. Similar results were

reported by Garza de Yta et al. (2012) who evaluated

growth response of juvenile C. quadricarinatus, reared in

tanks, offered soybean-based diets (350 g kg�1 crude pro-

tein, 71 g kg�1 lipids) containing either fish meal, poultry

by-product meal, ground pea meal or distillers’ dried grains

with solubles (DDGS) meal as protein source. No signifi-

cant differences were found in survival, growth or feed con-

version ratio (FCR) of redclaw crayfish. This might be

because natural productivity contributed to supplementa-

tion of minor deficiencies in essential amino acids. Thomp-

son et al. (2005) reported that juvenile redclaw reared in a

recirculating system can be offered practical diets contain-

ing 350 g kg�1 crude protein with no fish meal if a combi-

nation of less expensive plant-protein ingredients such as

SBM, wheat, brewer’s grains with yeast is added to the

diet. In pond culture of juvenile redclaw, practical diets

containing 280 g kg�1 crude protein with no fish meal but

containing a combination of plant-protein ingredients

(SBM, distillers’ DDGS and milo) was adequate for good

growth (Thompson et al. 2006).

Forage (detrital) crops (e.g. rice, hay) are often used in

freshwater crayfish cultivation (Ackefors 2000; Salame &

Rouse 2000; Jones et al. 2002) for presumed benefits such

as supplementation of direct and indirect sources of food

and supplying protective cover for moulting crayfish as

they seek refuge from predators. Fletcher & Warburton

(1997) tested fresh and decomposed duckweed (Spirodela

sp.) as feed for juvenile crayfish redclaw and found that

decomposed duckweed supported crayfish growth as well

as commercial pellets did. The authors suggested that prep-

aration of diets using detrital aquatic plants may be a cost-

effective method of increasing redclaw production. Salame

& Rouse (2000) evaluated forage-based feeding strategies

for redclaw reared in earthen ponds. Juvenile redclaw

stocked at a density of 4 m�2 were offered two feeding

regimes: manufactured pellets and pellets + forage (star-

grass (Cynodon plectostachyum) and janeiro grass (Erio-

chloa polystachya)) at a rate of 100 kg ha�1 month�1.

Survival and yield were greater in ponds receiving pellets

and forage than survival and yield in ponds receiving pel-

lets only. However, Metts et al. (2007) performed similar

experiments but found no benefit from forage supplementa-

tion. Generally, results of the majority of studies that have

been performed do not support using forage in redclaw

aquaculture.

Fish oil replacement Fish oils rich in HUFA are tradition-

ally used in aquatic animal feeds. Aquaculturists would like

to replace them with terrestrial plant oils such as linseed oil,

canola oil and corn oil, rich in linolenic acid (18:3n-3), lino-

leic acid (18:2n-6) and oleic acid (18:1n-9). Thompson et al.

(2010) evaluated growth response and fatty acid composi-

tion of juvenile redclaw crayfish offered diets containing

various lipid sources such as linseed oil, canola oil, corn oil,

beef tallow or menhaden oil. Crayfish offered plant-derived

oils performed as well as those offered feeds based on men-

haden oil containing high levels of n-3 HUFA. The diet

containing beef tallow had a higher percentage of saturated

fatty acids and resulted in poor growth. The authors con-

cluded that menhaden oil can be replaced by plant-based

oils with high levels of 18-carbon unsaturated fatty acids in

diets of juvenile redclaw, thus reducing costs for producers.

We suggest further research before definitive conclusions are

made but believe that because redclaw are freshwater organ-

isms, they probably can perform well without dietary EPA

and DHA inclusion. It is possible that they can elongate

and desaturate a-linolenic acid.

Crustaceans exhibit relatively slow and intermittent feeding

activity that has an impact on food acquisition and pro-

cessing (Loya-Javellana et al. 1995; Houser & Akiyama

1997). These behavioural characteristics affect physical

properties including water stability and durability of the

pellets (Meyers & Zein-Eldin 1975; Lim & Cuzon 1994;

Houser & Akiyama 1997; Obaldo et al. 2002). Pellets need

to be firmly bound to avoid breaking up into small parti-

cles that results in leaching of nutrients into water, reduc-

tion in water quality, poor animal growth, inefficient feed

conversion and low survival (Lim & Cuzon 1994; Houser

& Akiyama 1997; Obaldo et al. 2002). Binders affect pellet

stability in three ways: they reduce void spaces resulting in

a more compact and durable pellet; act as adhesives stick-

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Aquaculture Nutrition ª 2012 Blackwell Publishing Ltd

ing particles together; and exert a chemical action on the

ingredients and alter the nature of the feed resulting in a

more durable pellet (Lim & Cuzon 1994; DeSilva & Ander-

son 1995). Presently, diets of redclaw consist of steam

pressed highly conglomerated pellets of about 5 mm in

diameter (see Ruscoe et al. 2005). Redclaw can only ingest

large grain fragments; fine granules are propelled forward

and away from the mouth by currents induced by the

scaphognathites as water is passed through the gill chamber

(Ruscoe 2002). Moreover, redclaw move out of shelter

when food is offered, grasp large food items and return to

the relative safety of the artificial shelter where consump-

tion and digestion can be completed without fear of preda-

tion (Loya-Javellana et al. 1993). This means that many

pellets will lay on the pond bottom for a while before redc-

laws come searching for a second helping. Such feeding

behaviour requires a very water durable pellet to ensure

optimal FCR and growth.

Ruscoe et al. (2005) assessed the water stability of practi-

cal crayfish research diets manufactured using various bind-

ers (agar, gelatin, carboxymethylcellulose and carrageenan).

They found that rate of dry matter loss decreased over time

and that carrageenan and carboxymethylcellulose are better

binders than agar and gelatin. The 50 g kg�1 binder con-

centration slowed the decay rate by as much as 62% as

compared with 30 g kg�1 binder concentration. Addition-

ally, 10% moisture alginate-bound pellets are more stable

than 50% binder-bound pellets. Growth and survival are

not influenced by diet moisture although slightly better

growth was noted with moist diets offered to redclaw cray-

fish (Ruscoe et al. 2000). According to Ruscoe et al.

(2002), moist diets allow the manipulation and ingestion of

appropriately sized pieces as determined by the animal,

without the losses associated with abrasion, rasping and

subsequent fragmentation of hard-pellet diets. This is in

agreement with the morphological evidence suggesting that

the mouthparts of juvenile redclaw are well suited to the

ingestion of soft, moist foods (Loya-Javellana & Fielder

1997), where pappo-serrate setae on the labrum allow for

gentle prey manipulation by pushing prey down in front of

the mouth opening (Garm 2004).

Currently, there are few if any commercial feeds specifically

formulated and manufactured for redclaw crayfish. Progress

has been made over the past decade, but there are still

knowledge gaps in relation to nutrient requirements for red-

claw. Some of the areas that require further research include

essential amino acid requirements, vitamin and mineral

requirements, pelleting technology to produce a dry but mal-

leable pellet, an estimation of optimal feed regimens, brood-

stock nutrient requirements and formulations of diets using

regionally available ingredients with least cost formulations.

Presently, we recommend that semi-intensive farms use sink-

ing diets containing 350 g kg�1 crude protein, 60 g kg�1 lip-

ids, 18–20 MJ kg�1 digestible energy with crustacean

vitamin and mineral premix supplement and a water stability

of at least 30 min. Broodstock diets should contain fish oil

and carotenoid pigments. We believe redclaw aquaculture

has reached critical mass and is growing. With appropriate

feeds, nursery technology and growout protocols, produc-

tion is set to increase in the near future.

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