Aquaculture Research, 2011, 42, 835^843
Characterization of whitebait (Galaxias maculatus)
respiratory rates to optimize intensive culture carrying
capacities
Francisco Encina-Montoya1, Rolando Vega-Aguayo2, Alfonso Mardones-Lazcano2,Teresa Rueda1 &
Alfredo Tello1
1
Escuela de Ciencias Ambientales, Universidad Cato¤lica de Temuco,Temuco, Chile
2
Escuela de Acuicultura, Universidad Cato¤lica de Temuco,Temuco, Chile
Correspondence: F Encina-Montoya, Escuela de Ciencias Ambientales, Universidad Cato¤lica de Temuco, PO Box 15-D Temuco, Chile.
E-mail: fencina@uctemuco.cl
Abstract
Galaxias maculatus is an osmeriform native ¢sh of the
Southern Hemisphere, in which the crystalline lar-
vae is considered as a luxury delicacy, for this reason,
it has been commercially exploited in Chile, Argenti-
na and New Zealand. However, the ¢sheries have
been rapidly decreasing due to the overexploitation
and the predation of introduced species. Because of
these events, there is a need to determine a carrying
capacity for an intensive ¢sh culture. In order to opti-
mize stocking densities for ¢sh culture, this paper
proposes objectives to determine oxygen consump-
tion (OC) rates, dissolved oxygen concentrations that
produce signs of hypoxia and the average time
elapsed between food intake and peak OC in G. macu-
latus. In the oxygen experiments under routine meta-
bolism conditions, we found that G. maculatus adults
and whitebait showed signs of asphyxia at dissolved
oxygen concentrations between 1.3 and 2.2 mg L  1
and that adults tolerated dissolved oxygen levels as
low as1.3 mg L  1. The results showed that G. macula-
tus individuals with an average weight of 0.04 g con-
sumed 0.048 mg O2 h  1, whereas individuals with
an average weight of 1.4 g consumed 0.345 mg
O2 h  1. Galaxias maculatus increased the OC rate by
31%, from 0.39 to 0.51mg O2 h  1 g  1, occurring
14 min after food intake. The carrying capacities for
industrial cultures of G. maculatus, were estimated
using an allometric equation (OC 50.2363  W0.612),
a water £ow rate of 1m3 h  1 and an input oxygen
concentration of 10 mg L  1 at 12 1C. The density
culture of whitebait (4 g) can be allowed to reach
8^11kg m  3; therefore, these stocking densities
r 2010 Blackwell Publishing Ltd
doi:10.1111/j.1365-2109.2010.02692.
reduce the risk of hypoxia and mortality, ensuring
the appropriate growth and feed conversion rates.
Keywords: Galaxias maculatus, whitebait, oxygen
tolerance, respiration, carrying capacities
Introduction
Galaxias maculatus (Jenyns, 1842) is an osmeriform
¢sh that belongs to the Galaxiidae family and has a
circumpolar distribution. In Chile, G. maculatus can
be found in lakes, rivers and estuarine systems of
the Patagonia Region (Peredo & Sobarzo 1993, 1994)
between 32 and 531S (Campos 1970; Campos,
Dazarola, Dyer, Fuentes, GavilaŁn, Huaqu|¤ n, Mart|¤ nez,
Mele¤ndez, Pequeno, Ponce, Ruiz, Sielfeld, Soto,Vega &
Vila 1998; Murillo & Ruiz 2002), where G. maculatus
current conservation status is ranked as vulnerable
(Armesto, Ru|¤ z & Leo¤n-Lobos 1997; Glade 1988;
Campos et al. 1998) due to the overexploitation of
¢sheries and predation by the introduced salmonid
species. Research and commercial exploitation of this
species has been carried out in Chile, Argentina and
New Zealand, mainly due to the great value of
the crystalline larvae as a food product (US$28^
US$83 kg  1) (Mardones,Vega & Encina 2008).
The morphological stages of G. maculatus include
larvae and the juvenile stage (i.e. glass stage or white-
bait), where individuals usually measure between 4
and 6 mm and weigh o0.6 g. In the adult pigmented
stage, the individuals have an average length above
5 cm and weigh at least 0.6 g. However, in Chile and
835
Characterization of whitebait respiratory rates F Encina-Montoya et al.
New Zealand, the ¢sheries are only based on the
diadromous juveniles of G. maculatus (Campos 1970;
Valdebenito & Vega 2003). In Chile, the concentrated
research in G. maculatus, has been in the areas of
taxonomy (Campos 1970; Campos et al. 1998; Murillo
& Ruiz 2002), ecology, physiology (R|¤ os 1979; Murillo
& Ruiz 2002) and the development of intensive
culture (Vega 1999; Barile, Borquez, Dantagnan,
Mardones, Quevedo, Salgado, Valdebenito & Vega
2003; Mardones et al. 2008).
Intensive ¢sh cultures are limited by the oxygen
availability and the accumulation of metabolic
products, each of which are related to the food and
oxygen consumption (OC) rates of the species being
cultured (Piper1971,1983; Soderberg1995). In salmo-
nid cultures, for example, oxygen concentrations of
5 mg L  1 produce signs of asphyxia, and concentra-
tions below 3 mg L  1 generate mortalities (Leitriz &
Lewis 1980; Blanco Cachafeiro 1984; Valenzuela,
Alveal & Tarifeno 2002). The technical recommenda-
tion for the levels of oxygen availability in ¢sh
farming must be higher than 8 mg O2 L  1; this con-
centration is su⁄cient to avoid a¡ecting the indivi-
dual respiratory rates (Leitriz & Lewis 1980).
The lethal e¡ects of low dissolved oxygen levels
on freshwater ¢sh can be indicated by two end
points. Firstly, LT 50 is the average time that causes
50% mortality of the population due to the oxygen
concentration supplied. Secondly, LC 50 is the oxy-
gen level that causes 50% mortality within the ¢xed
time (USEPA 1986, as in the case of rainbow trout,
Landman, Van der Heubel & Ling (2005) estimated
a LC 50 of 1.62 mg O 2 L  1. The minimum oxygen con-
centration is an important factor when the oxygena-
tion system fails in intensive cultivation or in the
transportation of ¢sh. Therefore, when ¢sh are
exposed to conditions of hypoxia, they exhibit exter-
nal signs, such as changes in the gill ventilation rates,
which indicate physiological stress (McNeil & Closs
2007). Consequently, when the hypoxic conditions
continue, the ¢sh lose equilibrium, turn upside down
and die.
On the other hand, the OC rate of ¢sh is an impor-
tant parameter to design for intensive cultivation
systems. Klontz (1991) indicated that rainbow trout
individuals’ (0.3 kg) oxygen consumption rates were
between 105 and 149 mg O2 kg  1 h  1 under routine
metabolic conditions and at a temperature range of
7^11 1C. In trout, the oxygen consumption increases
after food ingestion, and the ¢sh require up to 70%
more oxygen than that under routine metabolism
conditions; this occurs approximately 15 min after
836
Aquaculture Research, 2011, 42, 835–843
food intake (Blanco Cachafeiro 1984). Therefore, the
aquaculture systems require the adjustment of max-
imum ¢sh densities and the £ux of water in response
to the maximum oxygen consumption rates shortly
after feeding (R|¤ os 1979; Blanco Cachafeiro 1984). In
most studies, oxygen consumption is measured un-
der conditions of: (a) routine metabolism, i.e. when
¢sh are food restricted and perform spontaneously
in normal activity (Waller1992), and (b) under condi-
tions of feed-related metabolism (Speci¢c Dynamic
Action ^ SDA), the metabolic rates increase due to
food consumption and absorption (McCue 2006).
Oxygen consumption rates are related to body weight
through the allometric equation OC 5 aWb (Y 5aXb)
(Kleiber 1961; Beamish 1964a, b; Paloheimo & Dickie
1966; Schmidt-Nielsen 1975). Research on the meta-
bolic characteristics of G. maculatus has only been
carried out by R|¤ os (1979), who studied the energetic
budget and metabolic di¡erences in populations in
the rivers and lakes of southern Chile.
Currently, there is limited research describing and
quantifying the metabolic characteristics of G. macu-
latus, consequently placing constraints on the de-
sign, implementation and the development of
optimal culture for commercial hatcheries. In com-
mercial salmon cultures, the densities need to range
from 10 to 100 kg m  3. However, in Chile, the experi-
mental culture of G. maculatus has traditionally used
low culture densities (2 kg m  3), which need to be
increased due to the economic considerations in
commercial cultures.
The objective of this study is to determine the oxy-
gen consumption rates, water oxygen concentration
levels that produce signs of hypoxia and furthermore,
the average time elapsed between food intake and the
peak oxygen consumption rates in G. maculatus. The
accurate characterization of these physiological
parameters is a critical factor in the optimization of
carrying capacities for species under intensive cul-
ture conditions.
Methods
Diadromous individuals of G. maculatus were ob-
tained from the Tolten Estuary in Chile’s IX Region
(39114 07200 S and 73113 01100W) and transported to a la-
boratory in the Universidad Cato¤lica de Temuco,
where they were treated with Chloramines T, Oxyte-
tracycline and salt. The specimens were adapted to
culture conditions in 100 L ¢breglass tanks, with a
water £ow rate of 1.71L min  1 for 30 days. During
r 2010 Blackwell Publishing Ltd, Aquaculture Research, 42, 835^843
Aquaculture Research, 2011, 42, 835^843 Characterization of whitebait respiratory rates F Encina-Montoya et al.

this period, the ¢sh were fed salmon starter pellets
without an oil bath.
Experiment 1. Oxygen consumption and
critical oxygen level under routine conditions
Individuals of G. maculatus were not fed for a period
of 24 h before measuring their rate of OC and the cri-
tical dissolved oxygen level in the water. Primarily,
¢sh were randomly chosen from the holding tank,
weighed and measured, and grouped into various
size classes, in which each group included 10 indivi-
duals. In the ¢rst stage of this experiment, the oxygen
consumption was estimated; ¢ve ¢sh classes were se-
lected with respect to length and body weight in a
range of 2^7 cm and 0.02^1.5 g respectively (Table
1). According to Mardones et al. (2008), individuals
less than 5 cm in length and less than 0.5 g in weight
correspond to larvae and juveniles. In the second
stage, the critical dissolved oxygen level was evalu-
ated; ¢ve ¢sh classes were selected with respect to
length and body weight in a range of 4^6.9 cm and
0.4^1.5 g respectively (Table 2).
In Tables 1 and 2, the measurement and weight are
shown; according to the ranges speci¢ed, the larvae
were placed in chambers with a capacity of 300 mL
and juveniles in chambers of 700 mL. The changes
in dissolved oxygen concentration (DO) were mea-
sured every 60 s using a Schott electrode (0.01mg
O2 L  1 sensitivity) and automatically recorded on a
Jeulin-Generis ESAO4 model 451101 data recorder
(S.A.V. Jeulin. B.P.1900. 27019, EŁvreux Cedex, France).
Each chamber was transferred to a 6 L aquarium and
maintained in a thermo regulated bath at 15 1C, un-
der an arti¢cial 12/12 h light/dark cycle. Beginning
at 09:00 hours, each experiment lasted approxi-
mately 3 h.
Oxygen consumption per gram of ¢sh (mg O2
h  1 g  1) was estimated according to Handy and De-
pledge (1999) using the following equation:
ðð½DOt2   ½DOt1 Þ  VÞ
OC ¼ ð1Þ
ðB  TÞ
where ð½DOt2   ½DOt1 Þ is the di¡erence in the dis-
solved oxygen concentrations (mg O2 L  1) at times
t1 and t2,V is the tank volume in litres (L), B is the bio-
mass in grams (g) and T is time in hours (h). The rela-
tion between oxygen consumption (mg O2 h  1) and
¢sh weight (g) was graphed according to the follow-
ing equation (R|¤ os 1979; Schmidt-Nielsen 1986):
OC ¼ aW b ð2Þ
where OC is the oxygen consumption in mg O2 h  1,
(a) and (b) are speci¢c constants and W is the ¢sh
weight (g). The critical dissolved oxygen level
(mg O2 L  1) was determined by measuring the dis-

Table 1 Oxygen consumption for di¡erent Galaxias maculatus weights under routine conditions

O2 average O2 average
Length Weight Mean No. of consumption consumption
Life stage class (cm) class (g) weight (g) repli cates (mg O2 h  1) SD (mg O2 h  1 g  1) SD

Whitebait 2.0–3.0 0.02–0.06 0.04 6 0.048 0.005 1.324 0.488

Whitebait 4.0–4.9 0.40–0.60 0.54 5 0.157 0.002 0.298 0.051
Adult 5.0–5.9 0.70–0.90 0.86 12 0.217 0.002 0.252 0.039
Adult 6.0–6.4 1.00–1.20 1.12 8 0.354 0.005 0.295 0.062
Adult 6.5–7.0 1.30–1.50 1.40 3 0.345 0.007 0.247 0.072

Table 2 Critical oxygen level for di¡erent Galaxias maculatus sizes under routine metabolism conditions

Average O2
Length Weight Average No. of critical level
Life stage class (cm) class (g) weight (g) replicates (mg O2 L  1) SD

Whitebait 4.0–4.9 0.4–0.6 0.5 6 2.0 0.7

Adult 5.0–5.4 0.7–0.8 0.8 6 2.2 1.2
Adult 5.5–5.9 0.9–1.7 1.0 8 1.8 0.5
Adult 6.0–6.4 1.2–1.3 1.3 8 1.7 0.8
Adult 6.5–6.9 1.3–1.5 1.4 3 1.3 0.3

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Characterization of whitebait respiratory rates F Encina-Montoya et al.
solved oxygen concentrations until the ¢sh lost equi-
librium and turned upside down.
Experiment 2. Time elapsed between food
intake and peak oxygen consumption (SDA)
Individuals of G. maculatus were taken from the hold-
ing tank, weighed, measured and 16 specimens were
selected with a mean weight of 1.2 g. The ¢sh were
placed in 700 mL hermetically closed oxygen meter
chambers and fed with salmon starter feed with a ra-
tion equivalent to 5% of the ¢sh biomass present in
each tank. After food ingestion, dissolved oxygen
concentrations (mg O2 L  1) were measured for a per-
iod of 3 h, similar to experiment 1. The time elapsed
between food intake and peak oxygen consumption
was estimated by the time at which the slope chan-
ged in the regression line between the oxygen con-
centration (mg O2 L  1) and time (h) for each of the
16 replicates.
Carrying capacity estimation
To estimate adequate densities for the culture of G.
maculatus, we considered that the output water
needs an oxygen saturation of at least 60%
(6.36 mg O2 L  1 at 12 1C). The conditions established
for the simulation were: ¢sh between 0.4 and 0.75 g,
an input oxygen concentration of 10 mg L  1, a 1m3
volume tank, a water £ow of 1m3 h  1 and a culture
density in the range of 1^20 kg m  3. The output oxy-
gen concentrations were estimated using the follow-
ing equations:
Oxygen output ¼
Mass flow of oxygen in the water input  Mass flow of oxygen consumed by fish
VðLÞ
½DO  QÞ  ½ðOC  D  W  VÞ  CF
O2 output ðmg L1 Þ ¼
V
ð3bÞ
where O2 output is the tank water oxygen output
(mg L  1), DO is the tank water input dissolved oxy-
gen concentration (mg h  1), Q is the water £ow
(L h  1), OC is the oxygen consumption from Eq. (2):
allometric relationship between oxygen consump-
838
Aquaculture Research, 2011, 42, 835–843
tion and weight (mg O2 h  1), D is the ¢sh density
(g L  1),W is the weight (g), CF is the correction factor:
rate adjustment due to increased oxygen consump-
tion while passing from routine metabolism to activa-
tion metabolism,V is the tank volume (L).
Data analysis
The average oxygen consumption rates were com-
pared between di¡erent sizes of G. maculatus using
analysis of variance (ANOVA). Data normality assump-
tions were tested using the Anderson^Darling test
and variance homogeneity was tested using Bartlett’s
test. Multiple comparisons were performed using Tu-
key’s test. The relationship between weight and the
oxygen consumption rate of di¡erent sizes of G. ma-
culatus was assessed using Pearson’s correlation coef-
¢cient (r) (Zar 1984).
Results
The relationship between ¢sh weight and oxygen
consumption (mg O2 h  1) in G. maculatus (Fig. 1) is
represented by the equation OC 5 0.144  W1.13, fol-
lowing the model described by Schmidt-Nielsen
(1975). Regression analysis showed a positive and sig-
ni¢cant correlation between weight and oxygen con-
sumption (r 5 0.94; Po0.05). Table 1 describes how
oxygen consumption increases with ¢sh weight. It
can be seen that ¢sh with an average weight of
0.04 g consume 0.048 mg O2 h  1, whereas ¢sh with
an average weight of1.4 g consume 0.345 mg O2 h  1.
Therefore, oxygen consumption per unit weight de-
creases as the weight increases. The smallest juvenile
ð3aÞ
¢sh, with an average of 0.04 g, consumed
1.324 mg O2 h  1 g  1, whereas, adults, between 0.86
and 1.40 g, consumed between a range of 0.252 and
0.295 mg O2 h  1 g  1. Signi¢cant di¡erences (P value
o0.5) were found in the average oxygen consump-
tions (mg O2 h  1 g  1) between the smallest juve-
niles (2.0^3.0 cm and 0.04 g) compared with the
other length classes.
Galaxias maculatus exhibited signs of asphyxia and
lost equilibrium under routine metabolic conditions
r 2010 Blackwell Publishing Ltd, Aquaculture Research, 42, 835^843
Aquaculture Research, 2011, 42, 835^843 Characterization of whitebait respiratory rates F Encina-Montoya et al.

0.70 OC = 0.36 W
G. maculatus (lake)
Rios (1979)

Oxygen consumption rate (mg O2 h–1)

0.60
OC = 0.0.38 W
G. maculatus (river)
0.50 Rios (1979)
OC = 0.14 W
Salvelinus fontinalis
0.40 Beamish (1964)

OC = 0.20 W
0.30 Salmo trutta
Beamish (1964)

0.20 OC = 0.30 W
Oncorhynchus rhodurus
Miura et al. (1976)
0.10 OC = 0.2363 W .
G. maculatus (estuary)
Our study results
0.00
0 0.5 1 1.5 2 2.5
W (g)

Figure 1 Comparison for di¡erent species of the relation between oxygen consumption (mg O2 h  1) and ¢sh weight (g)
based on the OC 5 aWb equation under a routine condition. The literature results were recalculated from mL O2 h  1 to mg
O2 h  1 to compare with our study (1.0 mg O2 h  1 51.428 mL O2 h  1, Bayne, Brown, Burns, Dixon, Ivanovici, Living-
stone, Lowe, Moore, Stebbing and Widdows (1985).

Table 3 Time period necessary for a change in the oxygen (Fig. 2), using the allometric equation (OC 5 0.2363
consumption rate of Galaxias maculatus individuals of 1.2 g  W0.612). Considering commercial cultures of
after food intake G. maculatus whitebait harvest, ¢sh with an average
weight of 0.5 g (range 0.4^0.75 g) and a correction
Time (min)
factor of 1.3 are needed to ensure the oxygen supply,
when the mg O2 h  1 g  1
consumption before mg O2 h  1 g  1
because the oxygen consumption increased by 31%
rate changes 14.4 min after 14.4 min after food intake. We found that ¢sh oxygen con-
sumption reaches critical levels (5^6.3 mg O2 L  1) at
Average 14.4 0.39 0.51
lower culture densities. Considering the critical
Standard 3.8 0.22 0.26
deviation
range, lighter individuals (0.4 g) need to be cultured
at densities of 8^11kg m  3 and heavier individuals
(0.75 g) at densities of 10^14 kg m  3 (Table 4).

when oxygen concentrations decreased between 2.0

and 1.3 mg O2 L  1 (Table 2) for juveniles of 0.5 g
Discussion
(4.3 cm) and adults of1.4 g (6.8 cm). The values of oxy-
gen concentration at which asphyxia signs developed Several studies have described the taxonomy, ecology
in the di¡erent stages were estimated to be between and physiology of G. maculatus; however, there are in-
1.3 and 2.2 mg L  1. There were no statistical di¡er- su⁄cient data to determine the development of inten-
ences between the di¡erent weight classes. Galaxias sive culture. In order to achieve high productivity in
maculatus of1.2 g increase their oxygen consumption intensive ¢sh culture systems, expressed in terms of
rate 14.4  3.8 min after food intake from 0.39 to growth rates or feed conversion rates, it is desirable
0.51mg O2 h  1 g  1 (Table 3), which represents an to combine high density with satiation feeding and
increase of 31% in the oxygen consumption rate. optimize parameters such as water temperature, oxy-
The di¡erences observed between the oxygen con- gen concentration and metabolic product concentra-
sumption rates before and after feeding were statisti- tions (Piper 1971, 1983; Soderberg 1995). There is an
cally signi¢cant (P value o0.05). extensive body of literature on the e¡ects of low
In this study, the output oxygen (mg L  1) was dissolved oxygen on freshwater ¢sh (Doudoro¡ &
estimated for di¡erent culture densities of whitebait Shumway 1970; Davis 1975; USEPA 1986; Dean &

r 2010 Blackwell Publishing Ltd, Aquaculture Research, 42, 835^843 839

Characterization of whitebait respiratory rates F Encina-Montoya et al. Aquaculture Research, 2011, 42, 835–843

0.40 g y = –0.44x + 10 at which G. maculatus individuals lost equilibrium

0.50 g y = –0.40x + 10 and turned upside down.
11
0.75 g y = –0.34x + 10 The average critical dissolved oxygen concentra-
10
tion level was in the range of 1.3 and 2.0 mg O2 L  1,
9 60% Oxygen Saturation
and no signi¢cant di¡erences were observed be-
Oxygen Output (mg L–1)

8 5.0 mg O2 L–1
tween juveniles and adults; these results agree with
7
the LC50 concentrations of Dean and Richardson
6 (1999) and Landman et al. (2005).
5 McNeil and Closs (2007) reported that £athead ga-
4 laxias (Galaxias rostratus) increase their gill ventila-
3 tion rate at 3 mg O2 L  1 and do not survive at
2 concentrations below 1mg O2 L  1. On the other
1 hand, Dean and Richardson (1999) found when
0 G. maculatus juveniles were exposed to 1mg O2 L  1
0 5 10 15 20 25 for 48 h, they presented a mortality of 61%, whereas
Density (kg m–3) adults su¡ered a mortality of 38%. In addition, Land-
Figure 2 Output oxygen estimation (mg L  1) for di¡er- man et al. (2005) found an LC50 of 2.65 mg O2 L  1 for
ent culture densities (kg m  3) of whitebait weighing 0.4, whitebait (mean weight of 0.38 g and length of 5 cm)
0.5 and 0.75 g. being less resistant to Oncorhynchus mykiss, with an
LC50 of 1.61mg O2 L  1, reporting that there are no
di¡erences between whitebait stages in their sensitiv-
Table 4 Estimated culture densities of Galaxias maculatus ity to dissolved oxygen, whereas Burleson, Wilhelm
(0.4, 0.5^0.75 g) for 5 and 6.4 mg O2 L  1 and Smatresk (2001) suggested that G. maculatus
juvenile whitebait are more tolerant than adults to
Estimated density Estimated density hypoxia. The lack of red blood cells is an important
(kg m  3) for (kg m  3) for
factor that can explain the sensitivity of hypoxia in
5 mg O2 L  1 6.4 mg O2 L  1
Weight (g) output output juvenile whitebait (Landman et al. 2005).
In order to reduce the e¡ects of hypoxia, some ¢sh
0.40 11.4 8.3
species rise to the water surface layer, which is rich in
0.50 12.4 9.0
oxygen, they gasp for air or breathe and agitate in the
0.75 14.6 10.6
water (Kramer 1987). This behaviour is de¢ned as
Aquatic Surface Respiration, which involves breath-
ing the oxygen-rich surface layer, a common re-
Richardson 1999; Karna 2003; Landman et al. 2005; sponse of ¢sh to hypoxia even when no air-
McNeil & Closs 2007). These papers summarize stu- breathing organ is present. This has been described
dies on the lethal concentrations of dissolved oxygen by McNeil and Closs (2007) for G. rostratus, a species
as LT50 or LC50 for many species, and their objectives very closely related to G. maculatus, which do not
are mainly related to pollution studies and water have an air-breathing organ; however, it is possible
quality standards for di¡erent uses. This is the reason that they may be able to breathe through their skin,
why measured endpoints depend on the objectives of as they have no scales at any stage of their life.
study. According to Landman et al. (2005), the LT50 is During routine metabolism, ¢sh move sponta-
not a good parameter for comparing the response to neously in the water column (Beamish & Mookherjii
minimum oxygen levels, particularly in sensitive 1964; Waller 1992), and during active metabolism,
species, because mortality occurs during the ¢rst there is an increase in muscle activity associated
45 min in one or two concentrations. The LC50 also with the swimming speed during feeding and diges-
presents drawbacks, because the acute response only tion (Brett 1962,1972). In this study, we measured the
occurs when the homeostasis levels have been ex- time elapsed between food intake and the beginning
ceeded. For this reason, the chronic endpoints are of digestion, which can be assessed by measuring
considered to be relevant ecological and physiologi- oxygen consumption. The results showed that at
cal responses. Therefore, in this study, under routine 15 1C, 14.4  3.8 min after feeding, ¢sh weighing
metabolic conditions, the critical dissolved oxygen 1.2 g increase their oxygen consumption rate from
concentration level was de¢ned as the concentration 0.39 to 0.51mg O2 h  1 g  1. Considering this factor,

840 r 2010 Blackwell Publishing Ltd, Aquaculture Research, 42, 835^843

Aquaculture Research, 2011, 42, 835^843 Characterization of whitebait respiratory rates F Encina-Montoya et al.
R|¤ os (1979) reported that G. maculatus, like salmon,
are e⁄cient carnivores that increase their metabolic
rate when they are feeding at dawn and sunset. The
high G. maculatus food e⁄ciency values reported by
R|¤ os (1979) agreed with the results obtained for other
carnivore species (Winber 1956, in Welch 1968; Davis
& Warren 1971). The maximum oxygen consumption
for Salvelinus alpinus, for example, occurs between 13
and 17 min after feeding, being very similar to the
times reported for other salmonids, e.g. trout (Blanco
Cachafeiro 1984). Other factors that a¡ect oxygen
consumption after feeding are the amount and size
of food ingested and the physiological stage of the ¢sh
(Rueda, Zamora and Mart|¤ nez (2001). In culture sys-
tems, the oxygen availability is a restrictive para-
meter for feed conversion and hypoxic conditions;
the concentration of dissolved oxygen is regulated
by the rate of renewal water, air pumps or the addi-
tion of pure oxygen.
The relationship between oxygen consumption
and weight is represented by the model OC 5 aWb
(Y 5aXb) (Kleiber 1961; Beamish 1964a, b; Paloheimo
& Dickie 1966; Schmidt-Nielsen 1975; R|¤ os 1979). We
found oxygen consumption rates similar to S. trutta
(Beamish 1964a, b) (Fig. 1). The results of R|¤ os (1979)
showed no signi¢cant di¡erences between popula-
tions of G. maculatus from Lake Rinihue and Cau
Cau River. However, these results with an estuarine
population showed lower rates than those reported
by R|¤ os (1979). The di¡erences in oxygen consump-
tion may be due to origins, metabolic stage and age.
In this research, the ¢sh were captured from the Tol-
ten River Estuary, whereas R|¤ os (1979) had studied
the ¢sh from the Valdivia River.
Under culture conditions, dissolved oxygen con-
centrations below 5 mg O2 L  1 can only occur due
to the failure of the oxygen supply equipment, the
water distribution systems or pollution-related is-
sues. Under accidental conditions of anoxia (i.e. oxy-
gen values declining to 3 mg O2), changes in gill
ventilation rates can be expected. The results showed
that the loss of equilibrium can be expected to occur
at oxygen concentrations between 1.3 and
2.0 mg O2 L  1. Landman et al. (2005) reported that
an oxygen concentration of 2.65 mg O2 L  1 produced
a mortality rate of 50% (LC50) of G. maculatus indivi-
duals and 1.62 mg O2 L  1 for rainbow trout. How-
ever, in this research in Chile, we found that the G.
maculatus population was shown to be more tolerant
to hypoxia than the G. maculatus and rainbow trout
populations that had been studied by Landman et al.
(2005).
r 2010 Blackwell Publishing Ltd, Aquaculture Research, 42, 835^843
In Chile, the experimental cultures of whitebait
G. maculatus have been carried out at low densities
(2 kg m  3). However, in commercial cultures, density
is expected to greatly advance higher than10 kg m  3;
therefore, the oxygen consumption rate after feeding
is relevant, because this determines the maximum
oxygen requirement. It is generally recommended for
commercial ¢sh cultures that oxygen concentrations
must never decline below 8 mg O2 L  1 (Leitriz & Le-
wis 1980; Valenzuela et al. 2002). Furthermore,WDOE
(2002) concluded, when using dissolved oxygen con-
centrations of 5^6 mg L  1, that the salmonid growth
rates declined o22%. Thus, ¢eld studies demon-
strated that embryo survival is low when the
dissolved oxygen content is below 5 mg L  1, and
survival is greater at 8 mg L  1 (Carter 2005). The
growth, food conversion e⁄ciency and swimming
performance are related to dissolved oxygen levels
and the process becomes less e⁄cient when the
oxygen concentrations are below 5 mg L  1
(Herrmann, Warren & Doudoro¡ 1962; Bjornn & Rei-
ser 1991; ODEQ 1995).
Using the physiological information, the carrying
capacities for industrial culture of G. maculatus were
estimated. The results showed that dissolved oxygen
concentrations in the culture units of G. maculatus of
estuarine origin need to be maintained at a level
above 60% of oxygen saturation (6.3 mg O2 L  1).
Therefore, the density culture can be allowed to
reach 8^11kg m  3; these stocking densities reduce
the risk of hypoxia and mortality, ensuring the ap-
propriate growth and feed conversion rates.
Acknowledgments
The authors wish to thank the following for their
support: (a) Conicyt Fondef Project D96i1071,
D99i1003, D02T1025, Fondecyt 1930134. (b) The Re-
search Department of Universidad Cato¤lica de Temu-
co, Chile, Projects DIUCT 2004-3-01, 2004-4-02. (c)
Robert McDowall, Santiago Peredo and Robyn Miles
for revision of the manuscript.
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