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Journal of Thermal Biology 35 (2010) 422–427
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Condition, growth and food conversion in barbel, Barbus barbus (L.) juveniles under different temperature/diet combinations Rafa" Kamin´ski, Ewa Kamler n, Jacek Wolnicki, Justyna Sikorska, Jakub Wa"owski The Stanis!aw Sakowicz Inland Fisheries Institute, Pond Fishery Department, Z˙abieniec, 05-500 Piaseczno, Poland
a r t i c l e in f o
a b s t r a c t
Article history: Received 18 June 2010 Accepted 10 September 2010
Populations of a rheophilic cyprinid Barbus barbus have declined in last decades, which created a need of conservation aquaculture. Production of stocking material in controlled conditions calls for optimization of the two major factors, temperature and diet. Condition, growth and food conversion ratio in fish fed a formulated diet Aller Futura were compared with those on natural food—frozen Chironomidae larvae at 17, 21 and 25 1C. Groups of 60 early juveniles (0.6–3.7 g) were reared in each of 18 aquaria in which six experimental groups were run in triplicate. Daily food ratios were adjusted according to fish biomass, differences in hydration between the two diets and rearing temperature. No mortality occurred during the experiment. Condition coefficient K was significantly higher in fish fed Aller Futura compared to those fed Chironomidae irrespective of temperature tested; body deformities were not recorded. Relative growth rate at the same temperature was always higher in fish on the formulated diet than in those fed Chironomidae, and food conversion ratio was always suppressed, both suggesting an efficient utilization of Aller Futura for growth in B. barbus early juveniles. On both diets the coefficient K was depressed at 21 1C. Relative growth rate (RGR) was accelerated with temperature according the Krogh’s ‘‘normal curve’’ within the range 21–25 1C, while at lower temperatures (17–21 1C) the observed values of temperature coefficient Q10 were much higher than the theoretical Q10 values based on Krogh’s ‘‘normal curve’’. Food conversion ratios (FCR) were reduced on both diets at 21 and 25 1C. Theoretical optimum temperatures for food conversion were 22.0 and 23.6 1C. Summing up, responses of three independent indices: condition, growth and food utilization locate the optimum temperature for B. barbus between 21 and 25 1C. No evidence was found that the effect of temperature on these indices was substantially modified by the diet. & 2010 Elsevier Ltd. All rights reserved.
Keywords: Barbus barbus Diet Food conversion ratio Growth rate Temperature
1. Introduction The barbel, Barbus barbus (Linnaeus, 1758), is a rheophilic cyprinid widely distributed in European submontane and lowland reaches of fast running rivers. During recent decades its stocks have declined dramatically due to deterioration and/or loss of microhabitats. Moreover, B. barbus is an attractive game fish which remains under a strong angling pressure; thus overfishing contributed to the reduction of fish numbers (Philippart et al., 1989; Lusk, 1996; Penczak and Kruk, 2000; Poncin and Philippart, 2002; Kottelat and Freyhof, 2007). In some Central European countries its conservational status is relatively high; the fish has been defined as near threatened or vulnerable (Schiemer and Waidbacher, 1992; Witkowski et al., 1999; Lusk et al., 2004). Restocking programs are carried out to restore populations in
n
Corresponding author. Tel.: + 48 22 7562044; fax: +48 22 7562088. E-mail addresses: erka@infish.com.pl (R. Kamin´ski), kamler@infish.com.pl (E. Kamler), jawol@infish.com.pl (J. Wolnicki), justyks@infish.com.pl (J. Sikorska), jwalowski@infish.com.pl (J. Wa"owski). 0306-4565/$ - see front matter & 2010 Elsevier Ltd. All rights reserved. doi:10.1016/j.jtherbio.2010.09.003
European running waters. For example, 772,000 newly-hatched larvae and summer fry, as well as 282 kg of autumn fry and 1+ B. barbus were stocked in 2009 to rivers controlled by Polish Angling Association (Przemys"aw Mielcarski, personal information). All these make B. barbus stocking material highly desirable. Since the 1980s relatively numerous attempts have been made to optimize methods for B. barbus captive breeding (Philippart, 1982; Poncin et al., 1987; Poncin, 1989, 1992) and larval rearing under controlled conditions (Wolnicki and Go´rny, 1995; Wolnicki, 1997; Kujawa et al., 1998; Kujawa, 2004; Fiala and Spurny´, 2001; Policar et al., 2007). Surprisingly little is known about technologies for the controlled rearing of this species in early juvenile period. Wolnicki (1997) did not found any differences in growth and survival rates at three fish densities of B. barbus juveniles fed a formulated diet of FK at 25 1C. Policar et al. (2007) reported a better growth of B. barbus juveniles fed a formulated diet of ASTA in aquaria than in troughs at 21 1C. Temperature effects and combined temperature/diet effects have not been studied in B. barbus juveniles yet. The results of such studies cannot be extrapolated from larval to juvenile rearing. A shift of optimum temperatures between early larvae and juveniles was found by
´ ski et al. / Journal of Thermal Biology 35 (2010) 422–427 R. Kamin
Keckeis et al. (2001) in another rheophilic cyprinid, Chondrostoma nasus. Temperature and food are among the major extrinsic factors both in the field and in fish culture. At the same time these two are the most expensive components of the aquaculture process. In the study reported here the response of B. barbus early juveniles to a formulated diet was compared with that of natural food at 17, 21 and 25 1C. A question was posed: at what temperature condition, growth rate and food conversion ratio is optimized in B. barbus under controlled conditions? Further, a hypothesis was tested that the optimum temperature is affected by diet quality.
2. Materials and methods 2.1. Fish The experimental fish were offspring of B. barbus spawners aged 5 +, whose parents originated from the San River, but the spawners were born and kept for their whole life indoor under controlled conditions. Ovulation was induced by means of a mGnRH analogue Ovopel (Horva´th et al., 1997). Eggs (32.9 g) were hand-stripped from one female (578 g weight) and fertilized by the dry method in one batch with pooled milt from five males (160–280 g). Incubation was performed in small (0.3 dm3) Weiss glasses at 18 1C. For the first 30 days of external feeding larvae were fed live, freshly hatched Artemia nauplii at 25 1C, whereupon the fish were fed frozen Chironomidae larvae at 17–20 1C. Prior to the experiment B. barbus early juveniles (six-month old, 42.372.7 mm TL, 0.6070.11 g BW, 7SD) were pre-selected to minimize variability of body size. Groups of 60 fish were stocked in each aquarium.
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The diet Aller Futura (Aller Aqua, Denmark, 62.1% crude protein, 11.9% crude fat, 8.5% carbohydrates, 10.5% ash and 7.0% moisture in raw matter; caloric value 22 kJ g 1 dry matter) or commercially available frozen Chironomidae larvae (Katrinex, Poland, 9.2%, 0.8%, 4.8%, 2.5% and 82.8%, respectively, in raw matter; 19 kJ g 1 dry matter) were fed to the groups F and C. Fish were fed manually at 8:00, 11:00, 14:00, 17:00 and 20:00, daily food ratios were equally distributed among the five feedings. During the days when fish measurements were performed the daily food ratios were distributed among three feedings at 14:00. 17:00 and 20:00. The demand for food in a poikilotherm is a result of an interplay of an intrinsic factor – body size with two extrinsic factors – temperature and diet. The daily food ratios were adjusted as usual according to the fish biomass (Fig. 1), and with differences in diet hydration between the Aller Futura dry diet and Chironomidae (Fig. 1, Table 1). In addition, adjustments of the daily ratios with the rearing temperature were made to mitigate a bias resulting from comparison of fish of contrasting size and ‘‘physiological age’’. These adjustments were made basing on Winberg’s (1956) temperature correction factors q (1.31, 0.920 and 0.659) for converting a rate at a temperature t (17, 21 and 25 1C, respectively) to 20 1C according to Krogh’s ‘‘normal curve’’ (Ege and Krogh, 1914). The temperature of 25 1C (recommended as the standard one for larvae and early juveniles of rheophilic cyprinids, review in Kamler and Wolnicki (2006)) was taken as a reference. Recalculated factors of 0.503 and 0.716 were used to
2.2. Experimental design Six feeding groups of B. barbus, each in triplicate, were fed a dry formulated diet Aller Futura (groups F) or a natural food—frozen Chironomidae larvae (groups C) at 17, 21 or 25 1C each (Table 1). Eighteen 20 dm3 glass aquaria were supplied with water from a recirculation system equipped with a polyethylene bed biofilter. Water flow through aquaria was about 0.3 dm3 min 1. Feces were siphoned out from the aquaria every day in the morning. The aquaria were equipped with a system preventing diet particles from being washed out. No uneaten food was found in the aquaria during inspections performed before each feeding. Photoperiod was 13L:11D, light intensity was approximately 700 lx at the water surface. Dissolved oxygen, measured twice daily, was kept at about 80–90% (Table 1). Total ammonia and nitrites were kept at o0.3 and o0.03 mg dm 3, respectively; conductivity between 382 and 417 mS cm 1; and pH between 7.7 and 8.1.
Fig. 1. Time course of daily food ratios of Aller Futura dry diet (F) and Chironomidae larvae (C) in six feeding groups of B. barbus juveniles at 17, 21 and 25 1C.
Table 1 Duration of the experiment, rearing temperature, dissolved oxygen, amount of food used during the experiment and mean relative feeding rate in six feeding groups of B. barbus juveniles fed Aller Futura diet (F) or Chironomidae larvae (C) Feeding group
F17 F21 F25 C17 C21 C25 a
Duration (days)
119 84 60 119 84 60
Temperature (1C)
Oxygen (%)
Food used (g aquarium 1)
Relative feeding rate (% fish biomass d 1)
Target
Actual 7SD
Mean 7SD
Mean
Target
17 21 25 17 21 25
17.0 7 0.1 21.0 7 0.2 24.9 7 0.2 16.9 7 0.1 21.0 7 0.2 25.0 7 0.2
94.07 6.0 87.2 7 9.3 81.5 7 11.1 93.1 7 5.8 91.3 7 8.3 86.8 7 11.0
99.9 136.0 142.0 654.9 700.5 760.4
1.48 2.11 2.95 11.30 16.09 22.46
Theoretical values computed using temperature correction factors (Winberg, 1956), further explanations in Section 2.2.
a
Actual 7 SD 1.477 0.05 2.067 0.00 2.967 0.02 11.21 7 0.05 15.77 7 0.18 22.46 7 0.12
´ ski et al. / Journal of Thermal Biology 35 (2010) 422–427 R. Kamin
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convert feeding rate from 25 to 17 1C and to 21 1C, respectively. Duration of rearing (119, 84 and 60 days at 17, 21 and 25 1C, Fig. 1, Table 1) was adjusted with temperature in a similar way. The actual (experimental) values were very close to the target (theoretical) ones (Table 1).
The minimum FCRwet was computed as FCRwetmin ¼ 2 A þ B1 topt þB2 topt .
The significance of differences was tested with one-way ANOVA followed by Duncan’s multiple range test, Po0.05.
2.3. Measurements and observations 3. Results On Day 1 of the experiment total length (TL, mm, up to the nearest 0.1 mm) and wet body weight (BW, g, up to 0.01 g) in all fish were individually determined after a short-term mild anaesthesia induced by 0.5 g dm 3 water solution of 2-phenoxyethanol (Myszkowski et al., 2003). All fish were measured and weighed individually (TL and BW) on six occasions during the experiment (Fig. 1) and were inspected for the presence of visible body deformities. Dry weight was taken after drying at 60 1C to constant weight in a desiccator over NaOH. 2.4. Data analysis
Condition coefficient (K) was calculated from wet body weight
(BW, g) and total length (TL, mm): K¼ 105 BW TL 3. A weightto length relationship computed for 180 initial B. barbus prior to the present experiment was BW ¼0.000017 TL2.79, which suggested an isometric growth. The relationship between BW and age (t, days) was approximated by an exponential equation: BWt ¼BW0eGt, where BW0 is theoretical wet body weight at time 0. Relative growth rate for wet body weight (RGRBW, % d 1, Ricker, 1975; Myszkowski, 1997) was calculated as RGRBW ¼ 100 (eG–1), where G ¼ln (BW2/BW1)(t2–t1) 1, and t is time (days). The common estimate for acceleration of growth rate with increase in temperature (Dt ¼t2–t1, 1C), the temperature 10=Dt coefficient, was taken from: Q10 ¼ RGRBW2 =RGRBW1 , where RGRBW2 and RGRBW2 are relative growth rates (% d 1) at temperatures t2 and t1, respectively. Theoretical Q10 values were computed from Backiel’s (1977) equation describing Krogh’s ‘‘normal curve’’: RGRBWt ¼ RGRBW20 (0.3e0.071t–0.24), where RGRBWt and RGRBW20 are relative growth rates at temperatures t and 20 1C, respectively. Food conversion ratio (FCR, g diet g fish weight gain 1) was expressed in two ways. As usual, it was expressed in terms of wet weight of both, diet and fish body (FCRwet, g g 1). Moreover, FCR was computed in terms of dry weight of diet and fish body (FCRdry, g g 1) in which an essential difference in hydration between Aller Futura and Chironomidae was accounted for. A second order polynomial FCRwet ¼A +B1t +B2t2 was used for the relation of FCRwet to temperature (t, 1C). The optimum temperature that minimizes FCR (topt, 1C) was taken from: topt ¼B1( 2B2) 1.
In all experimental groups a 100% survival rate was recorded. No deformed individuals were found. Condition coefficients (K) were consistently higher in fish fed on the formulated diet Aller Futura than in those fed on Chironomidae larvae at all the temperatures tested. On both the diets temperature 21 1C minimized the K values (Table 2). Relative growth rate (RGRBW) at the same temperature was always higher in fish on Aller Futura than in those fed on Chironomidae (Table 2). On both diets RGRBW increased with increase in temperature. Growth of B. barbus juveniles within the interval 0.5–3.5 g wet weight was well predicted by exponential models (Fig. 2, Table 3). Temperature-induced changes in the relative growth rate are summarized in Table 4. At the lower temperature range (17–21 1C) the observed Q10 values were much higher than the theoretical Q10 values based on Krogh’s ‘‘normal curve’’. In contrast, at the higher temperatures (from 21 to 25 1C) the observed Q10 values did not deviate much from the theoretical ones, and remained within the range 2–3 (Table 4). Food conversion ratio expressed in terms of wet weight (FCRwet) was higher in fish fed on Chironomidae by several fold compared with those fish on Aller Futura (Table 2); the theoretical minimum FCRwet on Chironomidae was higher by a factor of about
Fig. 2. Growth curves in B. barbus juveniles fed Aller Futura diet (F) or Chironomidae larvae (C) at 17, 21 and 25 1C. Parameters of the equations—see Table 3.
Table 2 Final total length (TL), body weight (BW), condition coefficient (K), relative growth rate for wet body weight (RGRBW) and food conversion ratio in terms of wet weight (FCRwet) or dry weight (FCRdry) in B. barbus juveniles fed Aller Futura diet (F) or Chironomidae larvae (C) at 17, 21 and 25 1C. Mean values7 SD of triplicate measurements. Group
TL (mm)
BW (g)
K
RGRBW (% d 1)
FCRwet (g g 1)
FCRdry (g g 1)
F17 F21 F25 C17 C21 C25
64.83 7 0.38c 78.65 7 0.34a 74.68 7 0.77b 59.15 7 0.37e 62.66 7 0.48d 62.97 7 0.03d
2.13 7 0.04c 3.67 7 0.04a 3.33 7 0.11b 1.55 7 0.02f 1.77 7 0.02e 1.88 7 0.02d
0.77 70.00b 0.75 70.01c 0.79 70.00a 0.75 70.00c 0.72 70.01d 0.75 70.01c
1.087 0.02e 2.197 0.02b 2.907 0.07a 0.817 0.02f 1.307 0.01d 1.937 0.01c
1.33 7 0.07c 0.967 0.01d 1.097 0.04c 13.90 7 0.22a 12.47 7 0.14b 12.30 7 0.24b
4.22 7 0.21c 3.047 0.04e 3.45 7 0.12d 8.16 7 0.13a 7.32 7 0.08b 7.22 7 0.14b
Within columns, results indicated by the same superscript lowercase letters are not significantly different (P4 0.05).
´ ski et al. / Journal of Thermal Biology 35 (2010) 422–427 R. Kamin
Table 3 Parameters of the exponential model BWt ¼BW0eGt describing growth curves in B. barbus juveniles fed Aller Futura diet (F) or Chironomidae larvae (C) at 17, 21 and 25 1C, n¼ 21 in each experimental group. Group
BW0
G
R2
F17 F21 F25 C17 C21 C25
0.604 0.627 0.653 0.606 0.629 0.635
0.0104 0.0215 0.0284 0.0080 0.0129 0.0189
0.9975 0.9970 0.9904 0.9979 0.9905 0.9901
Table 4 Comparison of temperature coefficients Q10 for growth in B. barbus juveniles. Temperature range (1C)
17–21 21–25
Observed Q10 computed from the Theoretical Q10 based on RGRBW values Krogh’s ‘‘normal curve’’ Aller Futura
Chironomidae
5.86 2.02
3.26 2.69
2.45 2.32
Table 5 Parameters of polynomials FCRwet ¼A + B1t + B2t2 describing the relation between food conversion ratio (FCRwet, g g 1 wet weight) and temperature (t, 1C) in B. barbus juveniles fed Aller Futura diet (F) or Chironomidae larvae (C). The optimum temperature that minimizes FCRwet (topt) and minimum food conversion ratio (FCRwetmin) are also shown. Diet
A
B1
B2
topt (1C)
FCRwetmin (g g 1)
F C
8.490 33.971
0.687 1.848
0.0156 0.0392
22.03 23.57
0.93 12.19
12 (see FCRwetmin in Table 5). This obvious difference only partly resulted from the difference in hydration between diets. The conversion from FCRwet to FCRdry did reduce the difference between these feeding groups but did not eliminate it. At comparable temperatures the FCRdry was higher in fish fed on Chironomidae by a factor of about two. The FCR expressed in terms of dry weight (FCRdry) revealed a difference between feeding groups F17 and F25, which could not be detected when FCRwet was used. Elevated values of both FCRwet and FCRdry were found at 17 1C on both diets (Table 2). Theoretical optimum temperatures that minimize FCRwet were found to be about 22.0 or 23.6 1C in fish fed on Aller Futura or Chironomidae, respectively (Table 5).
4. Discussion Condition coefficient K is a commonly used index that assumes isometric growth. The B-value 2.79 (i.e. close to 3) suggested that in B. barbus juveniles tested in the present experiment weight did increase with length in a cubic fashion, thus the condition coefficient K is expected to reflect well the condition variability among feeding groups. In many studies this coefficient has been used as a non-lethal, cheap and time-effective method to estimate condition in a large number of individuals. It also has been successfully used to predict proximate composition of whole body of fish juveniles. It was found to correlate positively and strongly with weight-specific total energy density, expressed as lipid % (Salam and Davies (1994) for Esox lucius; Herbinger and Friars (1991) as well as Sutton et al. (2000)
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for Salmo salar; Kamin´ski et al. (2005) for a cyprinid Eupallasella percnurus; Pangle and Sutton (2005) for Coregonus artedi) or as caloric value of body mass unit (Pangle and Sutton (2005) for C. artedi; Myszkowski et al. (2006) for Vimba vimba; and Kamler et al. (2006) for Tinca tinca). Moreover, strong, negative correlations were found in several species between condition coefficient K and whole body percentage of minerals (Salam and Davies, 1994; Kamin´ski et al., 2005; Kamler et al., 2006; Myszkowski et al., 2006). In the present study the K coefficient was higher in B. barbus fed on the formulated diet Aller Futura compared to those fed Chironomidae irrespective of temperature tested (Table 2). Thus, a higher lipid accumulation and depressed mineral content in the B. barbus fed Aller Futura than in fish on Chironomidae can be expected. High condition coefficients, excessive accumulation of lipids and deficiency of minerals are known to be associated with body deformities (Myszkowski et al.( 2002) for Carassius carassius; Helland et al. (2005) and Baeverfjord et al. (2008) for S. salar; Kamler et al. (2006) for T. tinca; review in Lall (2002)). It is well established that skeletal malformations are a major bottleneck in culture of early developmental stages of fish (review in Cahu et al., 2003), juvenile cyprinids in particular (review in Kamler and Wolnicki, 2006). The response to intensive feeding with the Aller Futura diet was found to be species-specific among 13 cyprinid species (Sikorska, 2009). Species such as Abramis brama, Leuciscus idus, L. leuciscus, Scardinius erythrophthalmus, C. carassius, Rutilus rutilus and L. cephalus were easy deforming fish in which daily ratio of about 2.5% of fish biomass resulted in 50–87% of malformed individuals after 60 days. Tinca tinca and V. vimba showed a medium susceptibility (11–24%), while B. barbus, C. nasus, Cyprinus carpio and Aspius aspius were considered as resistant species (0–3% of deformed individuals). Barbus barbus was the most resistant among the 13 cyprinids studied (Sikorska, 2009). Similarly, in the present experiment no body deformities of B. barbus on the formulated diet were observed. In this experiment B. barbus were not overfed, daily food ratios were strictly controlled (Table 1), and no uneaten food was left. The experimental conditions made reliable measurements of food conversion ratio (FCR) (Table 2). It is interesting that after the correction for hydration of both, diets and fish body, in fish fed Aller Futura the FCRdry remained lower than that in fish fed Chironomidae. That testifies to a good efficiency of utilization of the formulated diet for growth in this fish. In the natural conditions an important food component of B. barbus is aquatic macroinvertebrates, but also detritus and allochthonous debris. The latter formed 36% of B. barbus food in Vistula River near Warsaw (review in Brylin´ska, 2000). Irrespective of temperature the relative growth rates (RGRBW) of B. barbus fed Aller Futura were higher than those on Chironomidae (Table 2), which demonstrates once again that the formulated diet was well accepted by the B. barbus juveniles. In the present study the temperature effect on B. barbus performance was estimated using three independent parameters, condition coefficient (K), growth rate (RGRBW) and food conversion ratio (FCR). The condition coefficient K was significantly depressed at 21 1C on both diets (Table 2). In other words, at 21 1C fish were slender and were less endangered with potential overfeeding and its negative consequences as excessive fat storage, deficiency of minerals and body deformities. This suggests that 21 1C is a circum-optimum temperature. It is generally assumed that in aquatic species inhabiting temperate climate the Q10 values for circum-optimum temperatures remain at the level of 2–3 (thus do not deviate much from those based on Krogh’s ‘‘normal curve’’), while at sub-optimum
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´ ski et al. / Journal of Thermal Biology 35 (2010) 422–427 R. Kamin
temperatures the Q10 values are strongly elevated, exceeding those derived from Krogh’s ‘‘normal curve’’ (Kamler et al. (1998) for Q10 growth, Q10 developmental and Q10 metabolic in C. nasus; Kamin´ski et al. (2006) for Q10 developmental in E. percnurus; reviews in Rombough, 1988; Finn et al., 1995; Kamler, 2008). In the present work growth rate (RGRBW) was accelerated by temperature according to the Krogh’s ‘‘normal curve’’ within temperatures 21–25 1C, while at lower temperatures (17–21 1C) the Q10 values were much higher (Table 4). This indicates that the optimum temperature for growth of B. barbus juveniles remains within the range 21–25 1C. Food conversion ratios, both FCRwet and FCRdry, were reduced on both diets at 21 and 25 1C (Table 2), which suggests that efficiency of food utilization for growth was maximized within this range. Theoretical optimum temperatures were within a narrower range of about 22.0 and 23.6 1C, respectively, for fish fed the formulated diet and natural food (Table 5). The exact position of the optimum range remains to be verified experimentally using smaller temperature intervals. Taken together, conservation aquaculture seems to be a promising response to recent decline of B. barbus populations. There is a need to optimize captive breeding for restocking purposes. Temperature and food, the two important factors both in the field and in aquaculture, were considered. A question was posed: at what temperature performance of B. barbus is optimized under controlled conditions? Three independent indices: condition, growth and food utilization located the optimum temperature for B. barbus between 21 and 25 1C. Probably the optimum temperature is within a narrower range of about 22.0–23.6 1C. Further, a hypothesis was tested that the optimum temperature is affected by diet quality. No evidence was found that the response to temperature was substantially modified by the diet. A satisfactory performance of B. barbus early juveniles within 0.6–3.7 g was observed on the formulated diet Aller Futura, although elevated condition coefficient suggests that the formulated diet may result in an increase of fat deposits and decrease of mineral content.
Acknowledgements We are grateful to Przemys"aw Mielcarski from the Presidium of the Polish Angling Association for the information about B .barbus stocking. This study was supported by the Ministry of Science and Higher Education in Poland within the Project no. N311 297535 for 2008–2011. References Backiel, T., 1977. An equation for temperature dependent metabolism or for the ‘normal curve’ of Krogh. Pol. Arch. Hydrobiol. 24, 305–309. Baeverfjord, G., Hjelde, K., Helland, S., Refstie, S., 2008. Restricted dietary levels of phosphorus and zinc induce specific skeletal deformities in juvenile Atlantic salmon (Salmo salar L.). Eur. Aquacult. Soc. Spec. Publ. 37, 52–53. Brylin´ska, M., 2000. Freshwater Fishes of Poland New edn Wydawnictwo Naukowe PWN, Warszawa (in Polish). Cahu, C., Zambonino Infante, J., Takeuchi, T., 2003. Nutritional components affecting skeletal development in fish larvae. Aquaculture 227, 245–258. Ege, E., Krogh, A., 1914. On the relation between the temperature and the respiratory exchange in fishes. Int. Rev. Ges. Hydrobiol. Hydrog. 7, 48–55. Fiala, J., Spurny´, P., 2001. Intensive rearing of the common barbel (Barbus barbus L.) larvae using dry starter feeds and natural diet under controlled conditions. Czech. J. Anim. Sci. 46, 320–326. Finn, R.N., Rønnestad, I., Fyhn, H.J., 1995. Respiration, nitrogen and energy metabolism of developing yolk-sac larvae of Atlantic halibut (Hippoglossus hippoglossus). Comp. Biochem. Physiol. 111A, 647–671. Helland, S., Refstie, S., Espmark, A., Hjelde, K., Baeverfjord, G., 2005. Mineral balance and bone formation in fast-growing Atlantic salmon parr (Salmo salar) in response to dissolved metabolic carbon dioxide and restricted dietary phosphorus supply. Aquaculture 250, 364–376.
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Publ. 26, 155–156. Kujawa, R.J., 2004. Biological background for rearing rheophilic cyprinid larvae under controlled conditions. Dissertations and Monographs 88. Wyd. Uniw. Warmin´sko-Mazurskiego, Olsztyn (in Polish). Lall, S.P., 2002. The minerals. In: Halver, J.E., Hardy, R.W. (Eds.), Fish Nutrition 3rd Edition Elsevier, San Diego, CA, pp. 259–308. Lusk, S., 1996. Development and status of populations of Barbus barbus in the waters of the Czech Republic. Folia Zool. 45 (Suppl. 1), 39–46. Lusk, S., Hanel, L., Luskova´, V., 2004. Red list of the ichthyofauna of the Czech Republic: development and present status. Folia Zool. 53, 215–226. Myszkowski, L., 1997. Pitfalls of using growth rate coefficients. Pol. Arch. Hydrobiol. 44, 389–396. Myszkowski, L., Kamin´ski, R., Quiros, M., Stanny, A.L., Wolnicki, J., 2002. Dry dietinfluenced growth, size variability, condition and body deformities in juvenile crucian carp Carassius carassius reared under controlled conditions. Arch. Pol. Fish. 10, 51–61. Myszkowski, L., Kamin´ski, R., Wolnicki, J., 2003. Response of juvenile tench Tinca tinca (L.) to the anaesthetic 2-phenoxyethanol. J. Appl. Ichthyol. 19, 142–145. Myszkowski, L., Kamin´ski, R., Kamler, E., 2006. Compensatory growth and matter and energy deposition in Vimba vimba juveniles fed natural food or a formulated diet. Folia Zool. 55, 211–222. Pangle, K.L., Sutton, T.M., 2005. Temporal changes in the relationship between condition indices and proximate composition of juvenile Coregonus artedi. J. Fish Biol. 66, 1060–1072. Penczak, T., Kruk, A., 2000. Threatened obligatory riverine fishes in humanmodified Polish rivers. Ecol. Freshw. Fish 9, 109–117. Philippart, J.C., 1982. Mise au point de l’alevinage controˆ le´ du barbeau Barbus barbus (L.) en Belgique. Perspectives pour le rempoissonnement des rivie res. Cah. Ethol. Appl. 2, 173–202. Philippart, J.C., Melard, Ch., Poncin, P., 1989. Intensive culture of the common barbel, Barbus barbus (L.) for restocking. In: De Pauw, N., Jaspers, E., Ackefors, H, Wilkins, N. (Eds.), Aquaculture—A Biotechnology in Progress. European Aquaculture Society, Bredene, pp. 484–491. Policar, T., Koza´k, P., Hama´cˇkova´, J., Lepicˇova´, A., Musil, J., Kouˇril, J., 2007. Effects of short-time Artemia spp. feeding in larvae and different rearing environments in juveniles of common barbel (Barbus barbus) on their growth and survival under intensive controlled conditions. Aquat. Living Res. 20, 175–183. Poncin, P., 1989. Effects of different photoperiods on the reproduction of the barbel, Barbus barbus (L.), reared at constant temperature. J. Fish Biol. 35, 395–400. Poncin, P., 1992. Influence of the daily distribution of light on reproduction in the barbel, Barbus barbus (L.). J. Fish Biol. 41, 993–997. Poncin, P., Melard, Ch., Philippart, J., 1987. Use of temperature and photoperiod in the control of the reproduction of 3 European cyprinids: Barbus barbus (L.). Leuciscus cephalus (L.) and Tinca tinca (L.) reared in captivity. Preliminary results. Bull. Fr. Pˆeche Piscic. 304, 1–12. Poncin, P., Philippart, J.C., 2002. The role of aquaculture in fish conservation: a case study of Barbus barbus in Belgium. In: Collares-Pereira, M.J., Coelho, M.M., Cowx, I.G. (Eds.), Conservation of Freshwater Fishes: Options for the Future. Blackwell Science, Oxford, pp. 402–413. Ricker, W.E., 1975. Computation and Interpretation of Biological Statistics of Fish Populations. Department of the Environmental Fisheries and Marine Service, Ottawa. Rombough, P.J., 1988. Respiratory gas exchange, aerobic metabolism, and effects of hypoxia during early life. In: Hoar, W.S., Randall, D.J. (Eds.), Fish Physiology. Vol. XI Part A.. Academic Press, San Diego, pp. 59–161. Salam, A., Davies, P.M.C., 1994. Body composition of northern pike (Esox lucius L.) in relation to body size and condition factor. Fish. Res. 19, 194–204.
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Journal of Thermal Biology journal homepage: www.elsevier.com/locate/jtherbio
Condition, growth and food conversion in barbel, Barbus barbus (L.) juveniles under different temperature/diet combinations Rafa" Kamin´ski, Ewa Kamler n, Jacek Wolnicki, Justyna Sikorska, Jakub Wa"owski The Stanis!aw Sakowicz Inland Fisheries Institute, Pond Fishery Department, Z˙abieniec, 05-500 Piaseczno, Poland
a r t i c l e in f o
a b s t r a c t
Article history: Received 18 June 2010 Accepted 10 September 2010
Populations of a rheophilic cyprinid Barbus barbus have declined in last decades, which created a need of conservation aquaculture. Production of stocking material in controlled conditions calls for optimization of the two major factors, temperature and diet. Condition, growth and food conversion ratio in fish fed a formulated diet Aller Futura were compared with those on natural food—frozen Chironomidae larvae at 17, 21 and 25 1C. Groups of 60 early juveniles (0.6–3.7 g) were reared in each of 18 aquaria in which six experimental groups were run in triplicate. Daily food ratios were adjusted according to fish biomass, differences in hydration between the two diets and rearing temperature. No mortality occurred during the experiment. Condition coefficient K was significantly higher in fish fed Aller Futura compared to those fed Chironomidae irrespective of temperature tested; body deformities were not recorded. Relative growth rate at the same temperature was always higher in fish on the formulated diet than in those fed Chironomidae, and food conversion ratio was always suppressed, both suggesting an efficient utilization of Aller Futura for growth in B. barbus early juveniles. On both diets the coefficient K was depressed at 21 1C. Relative growth rate (RGR) was accelerated with temperature according the Krogh’s ‘‘normal curve’’ within the range 21–25 1C, while at lower temperatures (17–21 1C) the observed values of temperature coefficient Q10 were much higher than the theoretical Q10 values based on Krogh’s ‘‘normal curve’’. Food conversion ratios (FCR) were reduced on both diets at 21 and 25 1C. Theoretical optimum temperatures for food conversion were 22.0 and 23.6 1C. Summing up, responses of three independent indices: condition, growth and food utilization locate the optimum temperature for B. barbus between 21 and 25 1C. No evidence was found that the effect of temperature on these indices was substantially modified by the diet. & 2010 Elsevier Ltd. All rights reserved.
Keywords: Barbus barbus Diet Food conversion ratio Growth rate Temperature
1. Introduction The barbel, Barbus barbus (Linnaeus, 1758), is a rheophilic cyprinid widely distributed in European submontane and lowland reaches of fast running rivers. During recent decades its stocks have declined dramatically due to deterioration and/or loss of microhabitats. Moreover, B. barbus is an attractive game fish which remains under a strong angling pressure; thus overfishing contributed to the reduction of fish numbers (Philippart et al., 1989; Lusk, 1996; Penczak and Kruk, 2000; Poncin and Philippart, 2002; Kottelat and Freyhof, 2007). In some Central European countries its conservational status is relatively high; the fish has been defined as near threatened or vulnerable (Schiemer and Waidbacher, 1992; Witkowski et al., 1999; Lusk et al., 2004). Restocking programs are carried out to restore populations in
n
Corresponding author. Tel.: + 48 22 7562044; fax: +48 22 7562088. E-mail addresses: erka@infish.com.pl (R. Kamin´ski), kamler@infish.com.pl (E. Kamler), jawol@infish.com.pl (J. Wolnicki), justyks@infish.com.pl (J. Sikorska), jwalowski@infish.com.pl (J. Wa"owski). 0306-4565/$ - see front matter & 2010 Elsevier Ltd. All rights reserved. doi:10.1016/j.jtherbio.2010.09.003
European running waters. For example, 772,000 newly-hatched larvae and summer fry, as well as 282 kg of autumn fry and 1+ B. barbus were stocked in 2009 to rivers controlled by Polish Angling Association (Przemys"aw Mielcarski, personal information). All these make B. barbus stocking material highly desirable. Since the 1980s relatively numerous attempts have been made to optimize methods for B. barbus captive breeding (Philippart, 1982; Poncin et al., 1987; Poncin, 1989, 1992) and larval rearing under controlled conditions (Wolnicki and Go´rny, 1995; Wolnicki, 1997; Kujawa et al., 1998; Kujawa, 2004; Fiala and Spurny´, 2001; Policar et al., 2007). Surprisingly little is known about technologies for the controlled rearing of this species in early juvenile period. Wolnicki (1997) did not found any differences in growth and survival rates at three fish densities of B. barbus juveniles fed a formulated diet of FK at 25 1C. Policar et al. (2007) reported a better growth of B. barbus juveniles fed a formulated diet of ASTA in aquaria than in troughs at 21 1C. Temperature effects and combined temperature/diet effects have not been studied in B. barbus juveniles yet. The results of such studies cannot be extrapolated from larval to juvenile rearing. A shift of optimum temperatures between early larvae and juveniles was found by
´ ski et al. / Journal of Thermal Biology 35 (2010) 422–427 R. Kamin
Keckeis et al. (2001) in another rheophilic cyprinid, Chondrostoma nasus. Temperature and food are among the major extrinsic factors both in the field and in fish culture. At the same time these two are the most expensive components of the aquaculture process. In the study reported here the response of B. barbus early juveniles to a formulated diet was compared with that of natural food at 17, 21 and 25 1C. A question was posed: at what temperature condition, growth rate and food conversion ratio is optimized in B. barbus under controlled conditions? Further, a hypothesis was tested that the optimum temperature is affected by diet quality.
2. Materials and methods 2.1. Fish The experimental fish were offspring of B. barbus spawners aged 5 +, whose parents originated from the San River, but the spawners were born and kept for their whole life indoor under controlled conditions. Ovulation was induced by means of a mGnRH analogue Ovopel (Horva´th et al., 1997). Eggs (32.9 g) were hand-stripped from one female (578 g weight) and fertilized by the dry method in one batch with pooled milt from five males (160–280 g). Incubation was performed in small (0.3 dm3) Weiss glasses at 18 1C. For the first 30 days of external feeding larvae were fed live, freshly hatched Artemia nauplii at 25 1C, whereupon the fish were fed frozen Chironomidae larvae at 17–20 1C. Prior to the experiment B. barbus early juveniles (six-month old, 42.372.7 mm TL, 0.6070.11 g BW, 7SD) were pre-selected to minimize variability of body size. Groups of 60 fish were stocked in each aquarium.
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The diet Aller Futura (Aller Aqua, Denmark, 62.1% crude protein, 11.9% crude fat, 8.5% carbohydrates, 10.5% ash and 7.0% moisture in raw matter; caloric value 22 kJ g 1 dry matter) or commercially available frozen Chironomidae larvae (Katrinex, Poland, 9.2%, 0.8%, 4.8%, 2.5% and 82.8%, respectively, in raw matter; 19 kJ g 1 dry matter) were fed to the groups F and C. Fish were fed manually at 8:00, 11:00, 14:00, 17:00 and 20:00, daily food ratios were equally distributed among the five feedings. During the days when fish measurements were performed the daily food ratios were distributed among three feedings at 14:00. 17:00 and 20:00. The demand for food in a poikilotherm is a result of an interplay of an intrinsic factor – body size with two extrinsic factors – temperature and diet. The daily food ratios were adjusted as usual according to the fish biomass (Fig. 1), and with differences in diet hydration between the Aller Futura dry diet and Chironomidae (Fig. 1, Table 1). In addition, adjustments of the daily ratios with the rearing temperature were made to mitigate a bias resulting from comparison of fish of contrasting size and ‘‘physiological age’’. These adjustments were made basing on Winberg’s (1956) temperature correction factors q (1.31, 0.920 and 0.659) for converting a rate at a temperature t (17, 21 and 25 1C, respectively) to 20 1C according to Krogh’s ‘‘normal curve’’ (Ege and Krogh, 1914). The temperature of 25 1C (recommended as the standard one for larvae and early juveniles of rheophilic cyprinids, review in Kamler and Wolnicki (2006)) was taken as a reference. Recalculated factors of 0.503 and 0.716 were used to
2.2. Experimental design Six feeding groups of B. barbus, each in triplicate, were fed a dry formulated diet Aller Futura (groups F) or a natural food—frozen Chironomidae larvae (groups C) at 17, 21 or 25 1C each (Table 1). Eighteen 20 dm3 glass aquaria were supplied with water from a recirculation system equipped with a polyethylene bed biofilter. Water flow through aquaria was about 0.3 dm3 min 1. Feces were siphoned out from the aquaria every day in the morning. The aquaria were equipped with a system preventing diet particles from being washed out. No uneaten food was found in the aquaria during inspections performed before each feeding. Photoperiod was 13L:11D, light intensity was approximately 700 lx at the water surface. Dissolved oxygen, measured twice daily, was kept at about 80–90% (Table 1). Total ammonia and nitrites were kept at o0.3 and o0.03 mg dm 3, respectively; conductivity between 382 and 417 mS cm 1; and pH between 7.7 and 8.1.
Fig. 1. Time course of daily food ratios of Aller Futura dry diet (F) and Chironomidae larvae (C) in six feeding groups of B. barbus juveniles at 17, 21 and 25 1C.
Table 1 Duration of the experiment, rearing temperature, dissolved oxygen, amount of food used during the experiment and mean relative feeding rate in six feeding groups of B. barbus juveniles fed Aller Futura diet (F) or Chironomidae larvae (C) Feeding group
F17 F21 F25 C17 C21 C25 a
Duration (days)
119 84 60 119 84 60
Temperature (1C)
Oxygen (%)
Food used (g aquarium 1)
Relative feeding rate (% fish biomass d 1)
Target
Actual 7SD
Mean 7SD
Mean
Target
17 21 25 17 21 25
17.0 7 0.1 21.0 7 0.2 24.9 7 0.2 16.9 7 0.1 21.0 7 0.2 25.0 7 0.2
94.07 6.0 87.2 7 9.3 81.5 7 11.1 93.1 7 5.8 91.3 7 8.3 86.8 7 11.0
99.9 136.0 142.0 654.9 700.5 760.4
1.48 2.11 2.95 11.30 16.09 22.46
Theoretical values computed using temperature correction factors (Winberg, 1956), further explanations in Section 2.2.
a
Actual 7 SD 1.477 0.05 2.067 0.00 2.967 0.02 11.21 7 0.05 15.77 7 0.18 22.46 7 0.12
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convert feeding rate from 25 to 17 1C and to 21 1C, respectively. Duration of rearing (119, 84 and 60 days at 17, 21 and 25 1C, Fig. 1, Table 1) was adjusted with temperature in a similar way. The actual (experimental) values were very close to the target (theoretical) ones (Table 1).
The minimum FCRwet was computed as FCRwetmin ¼ 2 A þ B1 topt þB2 topt .
The significance of differences was tested with one-way ANOVA followed by Duncan’s multiple range test, Po0.05.
2.3. Measurements and observations 3. Results On Day 1 of the experiment total length (TL, mm, up to the nearest 0.1 mm) and wet body weight (BW, g, up to 0.01 g) in all fish were individually determined after a short-term mild anaesthesia induced by 0.5 g dm 3 water solution of 2-phenoxyethanol (Myszkowski et al., 2003). All fish were measured and weighed individually (TL and BW) on six occasions during the experiment (Fig. 1) and were inspected for the presence of visible body deformities. Dry weight was taken after drying at 60 1C to constant weight in a desiccator over NaOH. 2.4. Data analysis
Condition coefficient (K) was calculated from wet body weight
(BW, g) and total length (TL, mm): K¼ 105 BW TL 3. A weightto length relationship computed for 180 initial B. barbus prior to the present experiment was BW ¼0.000017 TL2.79, which suggested an isometric growth. The relationship between BW and age (t, days) was approximated by an exponential equation: BWt ¼BW0eGt, where BW0 is theoretical wet body weight at time 0. Relative growth rate for wet body weight (RGRBW, % d 1, Ricker, 1975; Myszkowski, 1997) was calculated as RGRBW ¼ 100 (eG–1), where G ¼ln (BW2/BW1)(t2–t1) 1, and t is time (days). The common estimate for acceleration of growth rate with increase in temperature (Dt ¼t2–t1, 1C), the temperature 10=Dt coefficient, was taken from: Q10 ¼ RGRBW2 =RGRBW1 , where RGRBW2 and RGRBW2 are relative growth rates (% d 1) at temperatures t2 and t1, respectively. Theoretical Q10 values were computed from Backiel’s (1977) equation describing Krogh’s ‘‘normal curve’’: RGRBWt ¼ RGRBW20 (0.3e0.071t–0.24), where RGRBWt and RGRBW20 are relative growth rates at temperatures t and 20 1C, respectively. Food conversion ratio (FCR, g diet g fish weight gain 1) was expressed in two ways. As usual, it was expressed in terms of wet weight of both, diet and fish body (FCRwet, g g 1). Moreover, FCR was computed in terms of dry weight of diet and fish body (FCRdry, g g 1) in which an essential difference in hydration between Aller Futura and Chironomidae was accounted for. A second order polynomial FCRwet ¼A +B1t +B2t2 was used for the relation of FCRwet to temperature (t, 1C). The optimum temperature that minimizes FCR (topt, 1C) was taken from: topt ¼B1( 2B2) 1.
In all experimental groups a 100% survival rate was recorded. No deformed individuals were found. Condition coefficients (K) were consistently higher in fish fed on the formulated diet Aller Futura than in those fed on Chironomidae larvae at all the temperatures tested. On both the diets temperature 21 1C minimized the K values (Table 2). Relative growth rate (RGRBW) at the same temperature was always higher in fish on Aller Futura than in those fed on Chironomidae (Table 2). On both diets RGRBW increased with increase in temperature. Growth of B. barbus juveniles within the interval 0.5–3.5 g wet weight was well predicted by exponential models (Fig. 2, Table 3). Temperature-induced changes in the relative growth rate are summarized in Table 4. At the lower temperature range (17–21 1C) the observed Q10 values were much higher than the theoretical Q10 values based on Krogh’s ‘‘normal curve’’. In contrast, at the higher temperatures (from 21 to 25 1C) the observed Q10 values did not deviate much from the theoretical ones, and remained within the range 2–3 (Table 4). Food conversion ratio expressed in terms of wet weight (FCRwet) was higher in fish fed on Chironomidae by several fold compared with those fish on Aller Futura (Table 2); the theoretical minimum FCRwet on Chironomidae was higher by a factor of about
Fig. 2. Growth curves in B. barbus juveniles fed Aller Futura diet (F) or Chironomidae larvae (C) at 17, 21 and 25 1C. Parameters of the equations—see Table 3.
Table 2 Final total length (TL), body weight (BW), condition coefficient (K), relative growth rate for wet body weight (RGRBW) and food conversion ratio in terms of wet weight (FCRwet) or dry weight (FCRdry) in B. barbus juveniles fed Aller Futura diet (F) or Chironomidae larvae (C) at 17, 21 and 25 1C. Mean values7 SD of triplicate measurements. Group
TL (mm)
BW (g)
K
RGRBW (% d 1)
FCRwet (g g 1)
FCRdry (g g 1)
F17 F21 F25 C17 C21 C25
64.83 7 0.38c 78.65 7 0.34a 74.68 7 0.77b 59.15 7 0.37e 62.66 7 0.48d 62.97 7 0.03d
2.13 7 0.04c 3.67 7 0.04a 3.33 7 0.11b 1.55 7 0.02f 1.77 7 0.02e 1.88 7 0.02d
0.77 70.00b 0.75 70.01c 0.79 70.00a 0.75 70.00c 0.72 70.01d 0.75 70.01c
1.087 0.02e 2.197 0.02b 2.907 0.07a 0.817 0.02f 1.307 0.01d 1.937 0.01c
1.33 7 0.07c 0.967 0.01d 1.097 0.04c 13.90 7 0.22a 12.47 7 0.14b 12.30 7 0.24b
4.22 7 0.21c 3.047 0.04e 3.45 7 0.12d 8.16 7 0.13a 7.32 7 0.08b 7.22 7 0.14b
Within columns, results indicated by the same superscript lowercase letters are not significantly different (P4 0.05).
´ ski et al. / Journal of Thermal Biology 35 (2010) 422–427 R. Kamin
Table 3 Parameters of the exponential model BWt ¼BW0eGt describing growth curves in B. barbus juveniles fed Aller Futura diet (F) or Chironomidae larvae (C) at 17, 21 and 25 1C, n¼ 21 in each experimental group. Group
BW0
G
R2
F17 F21 F25 C17 C21 C25
0.604 0.627 0.653 0.606 0.629 0.635
0.0104 0.0215 0.0284 0.0080 0.0129 0.0189
0.9975 0.9970 0.9904 0.9979 0.9905 0.9901
Table 4 Comparison of temperature coefficients Q10 for growth in B. barbus juveniles. Temperature range (1C)
17–21 21–25
Observed Q10 computed from the Theoretical Q10 based on RGRBW values Krogh’s ‘‘normal curve’’ Aller Futura
Chironomidae
5.86 2.02
3.26 2.69
2.45 2.32
Table 5 Parameters of polynomials FCRwet ¼A + B1t + B2t2 describing the relation between food conversion ratio (FCRwet, g g 1 wet weight) and temperature (t, 1C) in B. barbus juveniles fed Aller Futura diet (F) or Chironomidae larvae (C). The optimum temperature that minimizes FCRwet (topt) and minimum food conversion ratio (FCRwetmin) are also shown. Diet
A
B1
B2
topt (1C)
FCRwetmin (g g 1)
F C
8.490 33.971
0.687 1.848
0.0156 0.0392
22.03 23.57
0.93 12.19
12 (see FCRwetmin in Table 5). This obvious difference only partly resulted from the difference in hydration between diets. The conversion from FCRwet to FCRdry did reduce the difference between these feeding groups but did not eliminate it. At comparable temperatures the FCRdry was higher in fish fed on Chironomidae by a factor of about two. The FCR expressed in terms of dry weight (FCRdry) revealed a difference between feeding groups F17 and F25, which could not be detected when FCRwet was used. Elevated values of both FCRwet and FCRdry were found at 17 1C on both diets (Table 2). Theoretical optimum temperatures that minimize FCRwet were found to be about 22.0 or 23.6 1C in fish fed on Aller Futura or Chironomidae, respectively (Table 5).
4. Discussion Condition coefficient K is a commonly used index that assumes isometric growth. The B-value 2.79 (i.e. close to 3) suggested that in B. barbus juveniles tested in the present experiment weight did increase with length in a cubic fashion, thus the condition coefficient K is expected to reflect well the condition variability among feeding groups. In many studies this coefficient has been used as a non-lethal, cheap and time-effective method to estimate condition in a large number of individuals. It also has been successfully used to predict proximate composition of whole body of fish juveniles. It was found to correlate positively and strongly with weight-specific total energy density, expressed as lipid % (Salam and Davies (1994) for Esox lucius; Herbinger and Friars (1991) as well as Sutton et al. (2000)
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for Salmo salar; Kamin´ski et al. (2005) for a cyprinid Eupallasella percnurus; Pangle and Sutton (2005) for Coregonus artedi) or as caloric value of body mass unit (Pangle and Sutton (2005) for C. artedi; Myszkowski et al. (2006) for Vimba vimba; and Kamler et al. (2006) for Tinca tinca). Moreover, strong, negative correlations were found in several species between condition coefficient K and whole body percentage of minerals (Salam and Davies, 1994; Kamin´ski et al., 2005; Kamler et al., 2006; Myszkowski et al., 2006). In the present study the K coefficient was higher in B. barbus fed on the formulated diet Aller Futura compared to those fed Chironomidae irrespective of temperature tested (Table 2). Thus, a higher lipid accumulation and depressed mineral content in the B. barbus fed Aller Futura than in fish on Chironomidae can be expected. High condition coefficients, excessive accumulation of lipids and deficiency of minerals are known to be associated with body deformities (Myszkowski et al.( 2002) for Carassius carassius; Helland et al. (2005) and Baeverfjord et al. (2008) for S. salar; Kamler et al. (2006) for T. tinca; review in Lall (2002)). It is well established that skeletal malformations are a major bottleneck in culture of early developmental stages of fish (review in Cahu et al., 2003), juvenile cyprinids in particular (review in Kamler and Wolnicki, 2006). The response to intensive feeding with the Aller Futura diet was found to be species-specific among 13 cyprinid species (Sikorska, 2009). Species such as Abramis brama, Leuciscus idus, L. leuciscus, Scardinius erythrophthalmus, C. carassius, Rutilus rutilus and L. cephalus were easy deforming fish in which daily ratio of about 2.5% of fish biomass resulted in 50–87% of malformed individuals after 60 days. Tinca tinca and V. vimba showed a medium susceptibility (11–24%), while B. barbus, C. nasus, Cyprinus carpio and Aspius aspius were considered as resistant species (0–3% of deformed individuals). Barbus barbus was the most resistant among the 13 cyprinids studied (Sikorska, 2009). Similarly, in the present experiment no body deformities of B. barbus on the formulated diet were observed. In this experiment B. barbus were not overfed, daily food ratios were strictly controlled (Table 1), and no uneaten food was left. The experimental conditions made reliable measurements of food conversion ratio (FCR) (Table 2). It is interesting that after the correction for hydration of both, diets and fish body, in fish fed Aller Futura the FCRdry remained lower than that in fish fed Chironomidae. That testifies to a good efficiency of utilization of the formulated diet for growth in this fish. In the natural conditions an important food component of B. barbus is aquatic macroinvertebrates, but also detritus and allochthonous debris. The latter formed 36% of B. barbus food in Vistula River near Warsaw (review in Brylin´ska, 2000). Irrespective of temperature the relative growth rates (RGRBW) of B. barbus fed Aller Futura were higher than those on Chironomidae (Table 2), which demonstrates once again that the formulated diet was well accepted by the B. barbus juveniles. In the present study the temperature effect on B. barbus performance was estimated using three independent parameters, condition coefficient (K), growth rate (RGRBW) and food conversion ratio (FCR). The condition coefficient K was significantly depressed at 21 1C on both diets (Table 2). In other words, at 21 1C fish were slender and were less endangered with potential overfeeding and its negative consequences as excessive fat storage, deficiency of minerals and body deformities. This suggests that 21 1C is a circum-optimum temperature. It is generally assumed that in aquatic species inhabiting temperate climate the Q10 values for circum-optimum temperatures remain at the level of 2–3 (thus do not deviate much from those based on Krogh’s ‘‘normal curve’’), while at sub-optimum
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temperatures the Q10 values are strongly elevated, exceeding those derived from Krogh’s ‘‘normal curve’’ (Kamler et al. (1998) for Q10 growth, Q10 developmental and Q10 metabolic in C. nasus; Kamin´ski et al. (2006) for Q10 developmental in E. percnurus; reviews in Rombough, 1988; Finn et al., 1995; Kamler, 2008). In the present work growth rate (RGRBW) was accelerated by temperature according to the Krogh’s ‘‘normal curve’’ within temperatures 21–25 1C, while at lower temperatures (17–21 1C) the Q10 values were much higher (Table 4). This indicates that the optimum temperature for growth of B. barbus juveniles remains within the range 21–25 1C. Food conversion ratios, both FCRwet and FCRdry, were reduced on both diets at 21 and 25 1C (Table 2), which suggests that efficiency of food utilization for growth was maximized within this range. Theoretical optimum temperatures were within a narrower range of about 22.0 and 23.6 1C, respectively, for fish fed the formulated diet and natural food (Table 5). The exact position of the optimum range remains to be verified experimentally using smaller temperature intervals. Taken together, conservation aquaculture seems to be a promising response to recent decline of B. barbus populations. There is a need to optimize captive breeding for restocking purposes. Temperature and food, the two important factors both in the field and in aquaculture, were considered. A question was posed: at what temperature performance of B. barbus is optimized under controlled conditions? Three independent indices: condition, growth and food utilization located the optimum temperature for B. barbus between 21 and 25 1C. Probably the optimum temperature is within a narrower range of about 22.0–23.6 1C. Further, a hypothesis was tested that the optimum temperature is affected by diet quality. No evidence was found that the response to temperature was substantially modified by the diet. A satisfactory performance of B. barbus early juveniles within 0.6–3.7 g was observed on the formulated diet Aller Futura, although elevated condition coefficient suggests that the formulated diet may result in an increase of fat deposits and decrease of mineral content.
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