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Encina Et Al Puye DOI 20 5 2011
Encina Et Al Puye DOI 20 5 2011
Correspondence: F Encina-Montoya, Escuela de Ciencias Ambientales, Universidad Cato¤lica de Temuco, PO Box 15-D Temuco, Chile.
E-mail: fencina@uctemuco.cl
New Zealand, the ¢sheries are only based on the food intake (Blanco Cachafeiro 1984). Therefore, the
diadromous juveniles of G. maculatus (Campos 1970; aquaculture systems require the adjustment of max-
Valdebenito & Vega 2003). In Chile, the concentrated imum ¢sh densities and the £ux of water in response
research in G. maculatus, has been in the areas of to the maximum oxygen consumption rates shortly
taxonomy (Campos 1970; Campos et al. 1998; Murillo after feeding (R|¤ os 1979; Blanco Cachafeiro 1984). In
& Ruiz 2002), ecology, physiology (R|¤ os 1979; Murillo most studies, oxygen consumption is measured un-
& Ruiz 2002) and the development of intensive der conditions of: (a) routine metabolism, i.e. when
culture (Vega 1999; Barile, Borquez, Dantagnan, ¢sh are food restricted and perform spontaneously
Mardones, Quevedo, Salgado, Valdebenito & Vega in normal activity (Waller1992), and (b) under condi-
2003; Mardones et al. 2008). tions of feed-related metabolism (Speci¢c Dynamic
Intensive ¢sh cultures are limited by the oxygen Action ^ SDA), the metabolic rates increase due to
availability and the accumulation of metabolic food consumption and absorption (McCue 2006).
products, each of which are related to the food and Oxygen consumption rates are related to body weight
oxygen consumption (OC) rates of the species being through the allometric equation OC 5 aWb (Y 5aXb)
cultured (Piper1971,1983; Soderberg1995). In salmo- (Kleiber 1961; Beamish 1964a, b; Paloheimo & Dickie
nid cultures, for example, oxygen concentrations of 1966; Schmidt-Nielsen 1975). Research on the meta-
5 mg L 1 produce signs of asphyxia, and concentra- bolic characteristics of G. maculatus has only been
tions below 3 mg L 1 generate mortalities (Leitriz & carried out by R|¤ os (1979), who studied the energetic
Lewis 1980; Blanco Cachafeiro 1984; Valenzuela, budget and metabolic di¡erences in populations in
Alveal & Tarifeno 2002). The technical recommenda- the rivers and lakes of southern Chile.
tion for the levels of oxygen availability in ¢sh Currently, there is limited research describing and
farming must be higher than 8 mg O2 L 1; this con- quantifying the metabolic characteristics of G. macu-
centration is su⁄cient to avoid a¡ecting the indivi- latus, consequently placing constraints on the de-
dual respiratory rates (Leitriz & Lewis 1980). sign, implementation and the development of
The lethal e¡ects of low dissolved oxygen levels optimal culture for commercial hatcheries. In com-
on freshwater ¢sh can be indicated by two end mercial salmon cultures, the densities need to range
points. Firstly, LT 50 is the average time that causes from 10 to 100 kg m 3. However, in Chile, the experi-
50% mortality of the population due to the oxygen mental culture of G. maculatus has traditionally used
concentration supplied. Secondly, LC 50 is the oxy- low culture densities (2 kg m 3), which need to be
gen level that causes 50% mortality within the ¢xed increased due to the economic considerations in
time (USEPA 1986, as in the case of rainbow trout, commercial cultures.
Landman, Van der Heubel & Ling (2005) estimated The objective of this study is to determine the oxy-
a LC 50 of 1.62 mg O 2 L 1. The minimum oxygen con- gen consumption rates, water oxygen concentration
centration is an important factor when the oxygena- levels that produce signs of hypoxia and furthermore,
tion system fails in intensive cultivation or in the the average time elapsed between food intake and the
transportation of ¢sh. Therefore, when ¢sh are peak oxygen consumption rates in G. maculatus. The
exposed to conditions of hypoxia, they exhibit exter- accurate characterization of these physiological
nal signs, such as changes in the gill ventilation rates, parameters is a critical factor in the optimization of
which indicate physiological stress (McNeil & Closs carrying capacities for species under intensive cul-
2007). Consequently, when the hypoxic conditions ture conditions.
continue, the ¢sh lose equilibrium, turn upside down
and die.
On the other hand, the OC rate of ¢sh is an impor-
Methods
tant parameter to design for intensive cultivation
systems. Klontz (1991) indicated that rainbow trout Diadromous individuals of G. maculatus were ob-
individuals’ (0.3 kg) oxygen consumption rates were tained from the Tolten Estuary in Chile’s IX Region
between 105 and 149 mg O2 kg 1 h 1 under routine (39114 07200 S and 73113 01100W) and transported to a la-
metabolic conditions and at a temperature range of boratory in the Universidad Cato¤lica de Temuco,
7^11 1C. In trout, the oxygen consumption increases where they were treated with Chloramines T, Oxyte-
after food ingestion, and the ¢sh require up to 70% tracycline and salt. The specimens were adapted to
more oxygen than that under routine metabolism culture conditions in 100 L ¢breglass tanks, with a
conditions; this occurs approximately 15 min after water £ow rate of 1.71L min 1 for 30 days. During
this period, the ¢sh were fed salmon starter pellets in dissolved oxygen concentration (DO) were mea-
without an oil bath. 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).
Experiment 1. Oxygen consumption and Each chamber was transferred to a 6 L aquarium and
critical oxygen level under routine conditions maintained in a thermo regulated bath at 15 1C, un-
Individuals of G. maculatus were not fed for a period der an arti¢cial 12/12 h light/dark cycle. Beginning
of 24 h before measuring their rate of OC and the cri- at 09:00 hours, each experiment lasted approxi-
tical dissolved oxygen level in the water. Primarily, mately 3 h.
¢sh were randomly chosen from the holding tank, Oxygen consumption per gram of ¢sh (mg O2
weighed and measured, and grouped into various h 1 g 1) was estimated according to Handy and De-
size classes, in which each group included 10 indivi- pledge (1999) using the following equation:
duals. In the ¢rst stage of this experiment, the oxygen ðð½DOt2 ½DOt1 Þ VÞ
OC ¼ ð1Þ
consumption was estimated; ¢ve ¢sh classes were se- ðB TÞ
lected with respect to length and body weight in a where ð½DOt2 ½DOt1 Þ is the di¡erence in the dis-
range of 2^7 cm and 0.02^1.5 g respectively (Table solved oxygen concentrations (mg O2 L 1) at times
1). According to Mardones et al. (2008), individuals t1 and t2,V is the tank volume in litres (L), B is the bio-
less than 5 cm in length and less than 0.5 g in weight mass in grams (g) and T is time in hours (h). The rela-
correspond to larvae and juveniles. In the second tion between oxygen consumption (mg O2 h 1) and
stage, the critical dissolved oxygen level was evalu- ¢sh weight (g) was graphed according to the follow-
ated; ¢ve ¢sh classes were selected with respect to ing equation (R|¤ os 1979; Schmidt-Nielsen 1986):
length and body weight in a range of 4^6.9 cm and
0.4^1.5 g respectively (Table 2). OC ¼ aW b ð2Þ
In Tables 1 and 2, the measurement and weight are where OC is the oxygen consumption in mg O2 h 1,
shown; according to the ranges speci¢ed, the larvae (a) and (b) are speci¢c constants and W is the ¢sh
were placed in chambers with a capacity of 300 mL weight (g). The critical dissolved oxygen level
and juveniles in chambers of 700 mL. The changes (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
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
solved oxygen concentrations until the ¢sh lost equi- tion and weight (mg O2 h 1), D is the ¢sh density
librium and turned upside down. (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).
Experiment 2. Time elapsed between food
intake and peak oxygen consumption (SDA)
Individuals of G. maculatus were taken from the hold- Data analysis
ing tank, weighed, measured and 16 specimens were
The average oxygen consumption rates were com-
selected with a mean weight of 1.2 g. The ¢sh were
pared between di¡erent sizes of G. maculatus using
placed in 700 mL hermetically closed oxygen meter
analysis of variance (ANOVA). Data normality assump-
chambers and fed with salmon starter feed with a ra-
tions were tested using the Anderson^Darling test
tion equivalent to 5% of the ¢sh biomass present in
and variance homogeneity was tested using Bartlett’s
each tank. After food ingestion, dissolved oxygen
test. Multiple comparisons were performed using Tu-
concentrations (mg O2 L 1) were measured for a per-
key’s test. The relationship between weight and the
iod of 3 h, similar to experiment 1. The time elapsed
oxygen consumption rate of di¡erent sizes of G. ma-
between food intake and peak oxygen consumption
culatus was assessed using Pearson’s correlation coef-
was estimated by the time at which the slope chan-
¢cient (r) (Zar 1984).
ged in the regression line between the oxygen con-
centration (mg O2 L 1) and time (h) for each of the
16 replicates.
Results
The relationship between ¢sh weight and oxygen
consumption (mg O2 h 1) in G. maculatus (Fig. 1) is
Carrying capacity estimation
represented by the equation OC 5 0.144 W1.13, fol-
To estimate adequate densities for the culture of G. lowing the model described by Schmidt-Nielsen
maculatus, we considered that the output water (1975). Regression analysis showed a positive and sig-
needs an oxygen saturation of at least 60% ni¢cant correlation between weight and oxygen con-
(6.36 mg O2 L 1 at 12 1C). The conditions established sumption (r 5 0.94; Po0.05). Table 1 describes how
for the simulation were: ¢sh between 0.4 and 0.75 g, oxygen consumption increases with ¢sh weight. It
an input oxygen concentration of 10 mg L 1, a 1m3 can be seen that ¢sh with an average weight of
volume tank, a water £ow of 1m3 h 1 and a culture 0.04 g consume 0.048 mg O2 h 1, whereas ¢sh with
density in the range of 1^20 kg m 3. The output oxy- an average weight of1.4 g consume 0.345 mg O2 h 1.
gen concentrations were estimated using the follow- Therefore, oxygen consumption per unit weight de-
ing equations: creases as the weight increases. The smallest juvenile
Oxygen output ¼
Mass flow of oxygen in the water input Mass flow of oxygen consumed by fish ð3aÞ
VðLÞ
0.70 OC = 0.36 W
G. maculatus (lake)
Rios (1979)
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).
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,
R|¤ os (1979) reported that G. maculatus, like salmon, In Chile, the experimental cultures of whitebait
are e⁄cient carnivores that increase their metabolic G. maculatus have been carried out at low densities
rate when they are feeding at dawn and sunset. The (2 kg m 3). However, in commercial cultures, density
high G. maculatus food e⁄ciency values reported by is expected to greatly advance higher than10 kg m 3;
R|¤ os (1979) agreed with the results obtained for other therefore, the oxygen consumption rate after feeding
carnivore species (Winber 1956, in Welch 1968; Davis is relevant, because this determines the maximum
& Warren 1971). The maximum oxygen consumption oxygen requirement. It is generally recommended for
for Salvelinus alpinus, for example, occurs between 13 commercial ¢sh cultures that oxygen concentrations
and 17 min after feeding, being very similar to the must never decline below 8 mg O2 L 1 (Leitriz & Le-
times reported for other salmonids, e.g. trout (Blanco wis 1980; Valenzuela et al. 2002). Furthermore,WDOE
Cachafeiro 1984). Other factors that a¡ect oxygen (2002) concluded, when using dissolved oxygen con-
consumption after feeding are the amount and size centrations of 5^6 mg L 1, that the salmonid growth
of food ingested and the physiological stage of the ¢sh rates declined o22%. Thus, ¢eld studies demon-
(Rueda, Zamora and Mart|¤ nez (2001). In culture sys- strated that embryo survival is low when the
tems, the oxygen availability is a restrictive para- dissolved oxygen content is below 5 mg L 1, and
meter for feed conversion and hypoxic conditions; survival is greater at 8 mg L 1 (Carter 2005). The
the concentration of dissolved oxygen is regulated growth, food conversion e⁄ciency and swimming
by the rate of renewal water, air pumps or the addi- performance are related to dissolved oxygen levels
tion of pure oxygen. and the process becomes less e⁄cient when the
The relationship between oxygen consumption oxygen concentrations are below 5 mg L 1
and weight is represented by the model OC 5 aWb (Herrmann, Warren & Doudoro¡ 1962; Bjornn & Rei-
(Y 5aXb) (Kleiber 1961; Beamish 1964a, b; Paloheimo ser 1991; ODEQ 1995).
& Dickie 1966; Schmidt-Nielsen 1975; R|¤ os 1979). We Using the physiological information, the carrying
found oxygen consumption rates similar to S. trutta capacities for industrial culture of G. maculatus were
(Beamish 1964a, b) (Fig. 1). The results of R|¤ os (1979) estimated. The results showed that dissolved oxygen
showed no signi¢cant di¡erences between popula- concentrations in the culture units of G. maculatus of
tions of G. maculatus from Lake Rinihue and Cau estuarine origin need to be maintained at a level
Cau River. However, these results with an estuarine above 60% of oxygen saturation (6.3 mg O2 L 1).
population showed lower rates than those reported Therefore, the density culture can be allowed to
by R|¤ os (1979). The di¡erences in oxygen consump- reach 8^11kg m 3; these stocking densities reduce
tion may be due to origins, metabolic stage and age. the risk of hypoxia and mortality, ensuring the ap-
In this research, the ¢sh were captured from the Tol- propriate growth and feed conversion rates.
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 Acknowledgments
to the failure of the oxygen supply equipment, the The authors wish to thank the following for their
water distribution systems or pollution-related is- support: (a) Conicyt Fondef Project D96i1071,
sues. Under accidental conditions of anoxia (i.e. oxy- D99i1003, D02T1025, Fondecyt 1930134. (b) The Re-
gen values declining to 3 mg O2), changes in gill search Department of Universidad Cato¤lica de Temu-
ventilation rates can be expected. The results showed co, Chile, Projects DIUCT 2004-3-01, 2004-4-02. (c)
that the loss of equilibrium can be expected to occur Robert McDowall, Santiago Peredo and Robyn Miles
at oxygen concentrations between 1.3 and for revision of the manuscript.
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-
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