Factors driving spatial variation in egg survival of an ecologically and culturally important forage fish

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Disentangling density‐dependent effects on egg production and survival from egg to recruitment in fish

Understanding of density‐dependent effects is key to achieving sustainable management of self‐regulating biological resources such as fish stocks. Traditionally, density‐dependent effects on population abundance in fish have been considered to occur from hatching to recruitment, based on the paradigm of proportionality between spawning stock biomass and total egg production. Here, we demonstrate how the existence of intraspecific and interspecific density dependence in egg production changes the current understanding of density‐dependent processes in the life history of fish, by disentangling density‐dependent effects on egg production and survival from egg to recruitment, using sardine (Sardinops melanostictus, Clupeidae) and anchovy (Engraulis japonicus, Engraulidae) as model species. For sardine, strong intraspecific density‐dependent effects occurred in egg production, but no density‐dependent effects occurred or if any they were weak enough to be masked by environmental factors from hatching to recruitment. In contrast, for anchovy, interspecific density‐dependent effects occurred in egg production. In the survival after hatching, anchovy experienced stronger intraspecific density‐dependent effects than currently recognized. This analysis could overturn the current understanding of density‐dependent effects in the life history, highlighting contrasts between the effects on individual quality and population abundance and between the model species. We propose to reconsider the basis of fisheries management and recruitment studies based on the revised understanding of density‐dependent effects in the life history of the respective species.     

Modelling Atlantic mackerel spawning habitat suitability and its future distribution in the north‐west Atlantic

We investigated the effect of environmental conditions on Atlantic mackerel spawning habitat in the southern Gulf of St. Lawrence (sGSL). Based on generalized additive models, we (i) modelled the optimal spawning habitat of mackerel in the sGSL using daily egg production (DEP) in June, (ii) predicted known and new potential present spawning habitats in the GSL and the north‐west Atlantic, and (iii) assessed how they respond to future climate change. Our findings showed that both mackerel presence–absence and given‐presence DEP were associated with sea surface temperature (10–16.5°C), salinity above 31 and depth < 120 m. Adding zooplankton showed a marked effect on the DEP given‐presence compared to the presence–absence. Predictions of spawning habitats under present (1999–2012) and future scenario (2066–2085) conditions were estimated from the presence–absence model without zooplankton, using physical conditions of the BNAM. Under present conditions, our model predicted well the main spawning habitat in the sGSL and other known secondary spawning habitats in the northern GSL (nGSL), the western and southern Newfoundland, and the north‐west Atlantic coast. Under future conditions, our study suggests that spawning habitats in the sGSL and the nGSL would expand. Our results, therefore, suggest that mackerel could benefit from a warmer GSL, minimizing the potential for a northward migration of the stock due to decreasing suitability of the sGSL as its main spawning ground, and a new but spatially limited potential habitat in Newfoundland coasts. These results can be used to inform stock management and develop adaptive management plans in the context of climate change.     

Seasonal effects of macrophyte growth on rainbow trout habitat availability and selection in a low‐gradient,groundwater‐dominated river

Rainbow trout habitat use is often described in high‐gradient, runoff‐driven, heterotrophic streams where geomorphic features and overhanging riparian vegetation provide channel complexity and cover. However, many rainbow trout populations thrive in rivers with contrasting aquatic habitat. We describe rainbow trout habitat use in a low‐gradient, groundwater‐dominated tailwater river where river flow management and macrophyte growth and senescence largely govern available trout habitat. In the summers of 2013 and 2014, available aquatic habitat (depth, velocity, macrophyte cover, substrate size) was quantified, while individual trout location was determined by radio telemetry and linked to environmental variables. Detailed habitat surveys indicate that macrophyte cover increases throughout the summer and is a strong determinant of in‐stream habitat characteristics. Paired logistic regression shows that adult rainbow trout prefer greater depths. Water depth increases with macrophyte abundance at both reach and local scales as plants restrict flow, and available trout habitat is linked to this seasonal pattern. When macrophyte abundance is high, adult trout show secondary preference for localised areas of lower macrophyte cover but otherwise show no selectivity for macrophyte cover, velocity or substrate size. Results suggest that submerged aquatic plants increased the quantity and quality of rainbow trout habitat as a source of channel complexity and cover. Macrophytes may play a similar role in other low‐gradient streams and should not be overlooked by fisheries managers considering habitat suitability.     

Preliminary investigations on the effects of dietary lipid on the spawning performance and egg quality of black sea bass Centropristis striata L

Adult black sea bass Centropristis striata broodstock ( N =162) were fed three different dietary treatments: two commercially prepared diets with 45% protein and two different lipid levels (12% and 20%) (diets 1 and 2), and a diet of frozen Atlantic silversides Menidia menidia (SS, diet 3). Broodstock were held under controlled photothermal conditions and induced to spawn with an LHRHa pellet (72 μg kg⁻¹ bw). Dietary lipid had pronounced effects on spawning performance and egg quality. Diet 3 (SS) produced a significantly ( P <0.05) higher fertilization success (22.4%) than diets 1 (0.6%) and 2 (4.8%). The hatching success of fertilized eggs was similar in all diets (range=40–58.6%), but only two spawns from diet 1 (12% lipid) yielded viable yolk-sac larvae (YSL). Diet 3 (SS) also produced significantly more YSL per female (21.8 × 10³) than the diet 1 (0.3 × 10³). Eggs from diet 3 (SS) contained a significantly greater proportion of n-3 series fatty acids, with docosahexaenoic acid (DHA) as the largest fraction. Eggs from commercially prepared dietary treatments contained significantly more n-6 fatty acids. The poor spawning performance of fish fed diet 1 (12% lipid) may be related to higher levels of linoleic acid and lower levels of DHA in the diet.     

The effects of sperm–egg ratios on polyspermy in the blood clam,Tegillarca granosa

Polyspermy in Tegillarca granosa, which leads to failure or abnormal development of early embryos, appears to be directly related to varying sperm–egg ratios. We investigated the correlation between sperm–egg ratios and conditions of embryonic development by setting six strict sperm–egg ratios and subsequently scoring fertilization rate, abnormal rate, rates of trochophore and D‐shaped larvae. When sperm–egg ratios rose from 2 × 10²:1 to 1 × 10³:1, D‐shaped larvae rate dropped significantly from 78.8 ± 13.3% to 47.8 ± 14.9% (analysis of variance, P< 0.05), and to 3.6 ± 2.8% in the 2 × 10⁴:1 group. Using fluorescence microscope observation, polyspermy rates rose from 2.6 ± 0.8% to 72.9 ± 4.8% with increases in sperm–egg ratios. Sperm–egg ratios were therefore correlated with polyspermy rate, which seriously affected larval survival. Under the premise of higher fertilization rate, the sperm–egg ratios of ≤2 × 10²:1 appeared to be more beneficial for embryonic development of T. granosa.     

Investigation of the temporal effects of spawning season and maternal and paternal differences on egg quality in Atlantic cod Gadus morhua L. broodstock

A better understanding of the parameters affecting egg quality and larval survival is of importance for continued development of cod broodstock and efficient husbandry practices. Decision tree analysis (DTA) was applied to analyse 3 years of egg quality data in an effort to extract the most important variables (i.e. predictors) in explaining differences in egg quality. The effect of three predictors (spawning time, maternal and paternal differences) has been studied on early cleavage pattern parameters, egg diameters, fertilization and hatching rates and has shown that females are the dominant variable and that time has a limited and inconsistent impact on the data. When using maternal, paternal differences and batch number (instead of spawning time) as predictors, the results confirm that no particular relationship is found between batch order (i.e. order in time) and egg quality. Moreover, batches with a higher egg quality show a consistency in the parameters assessed (i.e. batches with higher rates of normality in any parameter tend to be normal for other parameters). This is confirmed by the significant correlations found between cleavage parameters. Our results highlight that spawning time is of less importance than female parent contribution in ensuring high rates of fertilization and larval hatch, and maximizing general egg quality.     

Effects of predation and intraspecific interactions on habitat use and foraging by brown trout in artificial streams

Abstract— We studied habitat use, foraging rates and behavior of 10 cm and 12 cm long brown trout, Salmo trutta , at two densities, 1.5 and 3.0 fish. m⁻², in artificial streams that contained either the amphipod, Gammarus pulex , alone or G. pulex together with the piscivore, northern pike, Esox lucius. Gammarus were stocked in and largely restricted to the pools at a density of 128 Gammurus. m⁻² . pool⁻¹ Large trout (12 cm) used pools more and riffles less when small trout (10 cm) were present than when small trout were absent. Small trout consumed fewer Gammarus when together with large trout than when alone, but showed no difference in habitat use in the presence and abscnce of large trout. Habitat use and number of Gammarus consumed per trout were not affected by trout density for either size-class when alone. For both size-classes of trout, use of pools and foraging rates were higher in the absence than in the presence of pike, and pike primarily resided in the pools. The number of aggressive interactions by both size-classes of trout decreased when pike was present. Our results indicate that for habitats that differ in food resources and predation risk, size structure may affect habitat use and foraging by brown trout.     

Effects of simulated light intensity,habitat complexity and forage type on predator–prey interactions in walleye Sander vitreus

Predator‐prey interactions can be influenced by the behaviour of individual species as well as environmental factors. We conducted laboratory experiments to test for the influences of two abiotic factors (light intensity and habitat complexity) on predator–prey interactions between walleye Sander vitreus and two prey species, bluegill Lepomis macrochirus and golden shiner Notemigonus crysoleucas. Three light intensities were simulated (day, twilight and night) in the presence or absence of simulated vegetation. Observations of predator behaviour indicated that walleye increased activity and foraging success with decreasing light levels and had most success capturing dispersed, closer prey. While schooling could not be maintained as light levels diminished, prey decreased predation vulnerability by moving into vegetation or higher in the water column. Throughout all treatments, bluegill were more evasive to capture as the number of strikes was similar on both prey but capture rates were higher for golden shiner. Although light intensity and simulated habitat complexity affected predator and prey behaviour, these factors did not interact to influence foraging success of walleye. To fully understand predator and prey behaviours in fishes, an understanding of species‐specific responses to abiotic and biotic factors is necessary.     

Perceptual constraints on stream fish habitat selection: effects of food availability and water velocity

Abstract– Habitat selection theory assumes that individuals either know the quality of habitats a priori or can learn it through experience. This work tests the effects of total food supply and of water velocity on the ability of drift-feeding stream fish ( Rhinicthys atratulus ) to discriminate between two patches in a laboratory flow-tank that differ only in the amount of food in the patch. If the fish can discriminate between the patches, their distributions should be predicted by ideal free distribution models; otherwise, their distributions should be random. The results show that at high total food supply levels, fish distributions differ from random in general agreement with ideal free distribution models; total food supply decreases, fish distributions become random, indicating that total food supply levels affect the ability of fish to discriminate between habitats that differ only in food supply. Increased water velocity was expected to affect fish distributions similarly, but its effect was not significant. The results suggest that habitat selection theory should consider both differences between habitats and the ability of fish to perceive those differences.     

The effects of dietary HUFA level on flame angelfish (Centropyge loriculus) spawning,egg quality and early larval characteristics

This study was conducted to determine the effects of dietary highly unsaturated fatty acids (HUFA) on flame angelfish (Centropyge loriculus) reproduction, and egg and larval quality. In the experiment, formulated diets containing 1.8, 2.9 or 3.6% n‐3 HUFA were fed to flame angelfish broodstock (n = 4) for 5 months. Mean fecundity (daily egg production), egg fertilization rates and embryo viability were used as indicators of egg quality. In addition, mean egg diameter, oil globule diameter, per cent hatch, larval size at hatch, per cent survival to yolk exhaustion and larval size at yolk exhaustion were recorded for each treatment. Flame angelfish fed the diet containing 3.6% n‐3 HUFA exhibited significantly increased fecundity, fertilization rates and embryo viability than fish that were fed the other two formulated diets. Egg diameter, egg oil globule diameter, larval size at hatch, larval survival to yolk exhaustion and larval size at yolk exhaustion from the 3.6% n‐3 HUFA broodstock treatment group were not significantly different from those derived from the Control broodstock treatment. These data revealed that flame angelfish egg quality could respond rapidly (within weeks) to maternal dietary changes. Results from this study further support that dietary HUFA composition can significantly affect broodstock reproductive performance as well as subsequent performance of eggs and larvae.     

The effects of salinity on reproductive development and egg and larvae survival in the spotted scat Scatophagus argus under controlled conditions

In the present study, we report the first successful instance of controlled reproduction in Scatophagus argus, which has recently emerged as a new aquaculture resource. The controlled reproduction process for S. argus was optimized with regard to salinity acclimation. Gonadal maturation was affected by salinity in both sexes. Levels of plasma 17β‐estradiol (E₂) and 11‐ketotestosterone (11‐kT) were salinity dependent and increased significantly with the duration of acclimation. Plasma levels of gonadal steroids were higher in fish held at 25‰ salinity. The highest gonadosomatic indices (GSI), 15.1 ± 1.6 in the female and 6.4 ± 1.2 in the male, were also observed at 25‰ salinity. Nevertheless, the optimal salinity for S. argus embryonic development and larval culture was 15‰. Thus, the salinity requirement for gonadal maturation and early development are quite different. The use of advanced reproductive technologies combining salinity acclimation and stimulation of luteinizing hormone‐releasing hormone analog (LHRH‐A₂) resulted in a fertilization rate of 83.2%–91.3% and embryonic survival rates of over 90%. Embryos of S. argus at the 2‐cell, blastula, gastrula and pharyngula stages were observed. Most embryos hatched after 21.0 hr of incubation at 28.0 ± 1.0°C. The development of larvae into juveniles was completed at 40–45 days posthatch (dph). In this study, we provide information about the controlled reproduction of S. argus and identify the optimal environmental parameters for S. argus embryonic and larval culture, with the aim of developing reliable reproductive techniques for its mass production.     

Negative effect of zebra mussels on foraging and habitat use by lake sturgeon (Acipenser fulvescens)

• 1. Lake sturgeon (Acipenser fulvescens) are threatened or endangered throughout much of their range. Juvenile sturgeon utilize sandy and silty habitats extensively during their growth. Invasive zebra mussels change the nature of sandy and silty habitats because they settle on and coat the habitat with the shells of living and dead individuals. The potential impacts of this increased habitat complexity on lake sturgeon is unknown.
• 2. Juvenile lake sturgeon habitat choice was assessed in laboratory experiments, and zebra mussel impact on the foraging success of juvenile lake sturgeon on three different prey species was measured.
• 3. Sturgeon foraging on chironomids was virtually eliminated by 95% zebra mussel cover of the sand floor of the foraging arena, and 50% cover reduced foraging significantly. Foraging on more mobile prey items (amphipods and isopods) was essentially eliminated by either 95% or 50% zebra mussel cover of the arena floor. In habitat choice experiments, sturgeon avoided the zebra‐mussel‐covered habitat more than 90% of the time.
• 4. This combination of zebra mussel avoidance and reduced foraging in the presence of zebra mussels may be detrimental to sturgeon restocking programmes utilizing smaller sturgeon in zebra‐mussel‐infested waterways.

Copyright © 2006 John Wiley & Sons, Ltd.     

LHRHa‐induced ovulation of the endangered‐Caspian brown trout (Salmo trutta caspius) and its effect on egg quality and two sex steroids: testosterone and 17α‐hydroxyprogestrone

To induce synchronized ovulation, migrating wild Caspian brown trout (Salmo trutta caspius) females were treated with two interperitoneal injections of Des‐Gly¹⁰, d ‐Ala⁶ LHRH (LHRHa), given 3 days apart. Two injections of 100 μg kg^?1 body weight of this hormone effectively induced ovulation. Within 27 days from the second injection, all fish injected with 100 μg kg^?1 LHRHa had ovulated compared with 54.5% of the controls. The mean time to ovulation was reduced significantly (P<0.05) from 31.67±4.84 days in control fish and 28.83±7.31 days in sham‐treated fish to 16.36±1.61 days in fish injected with 100 μg kg^?1 LHRHa. The fertilization rate in 50 and 100 μg kg^?1 LHRHa‐injected fish was significantly lower than that in the control fish (P<0.05). In fish injected with 50 and 100 μg kg^?1 LHRHa, significant (P<0.05) changes in testosterone (T) and 17α‐hydroxyprogestrone (OHP) levels were observed. After the second LHRHa injection, the fish injected with 100 μg kg^?1 showed the highest serum levels of testosterone and OHP. These results demonstrate that the use of LHRHa can effectively reduce the mean time to ovulation and induce synchronized ovulation in Caspian brown trout.     

Synchronization of ovulation in cultured northern whitefish (Coregonus peled,Gmelin 1788) using [D‐Arg6Pro9Net]‐sGnRH analogue and its effect on egg quality

Female northern whitefish were treated with salmon [D‐Arg⁶Pro⁹Net]‐sGnRHa emulsified in Freund′s incomplete adjuvant as a sustained release treatment (sGnRHa‐FIA) or with [D‐Arg⁶Pro⁹Net]‐sGnRHa dissolved in physiological saline as acute release treatment (sGnRHa‐PS). Females in four experimental groups (A, B, C, D) and one control group (E) were injected intraperitoneally as follows: A and B – sGnRHa‐FIA at doses of 50 and 25 μg kg^?1, respectively; C – double injection (DI) of sGnRHa‐PS at 25 μg kg^?1 administered 3 days apart; D – single injection (SI) of sGnRHa‐PS at 25 μg kg^?1; and E – control group receiving a 1:1 mixture of FIA and physiological saline. All treatments induced and synchronized ovulation. Mean time to ovulation was significantly reduced (P < 0.05) in GnRHa‐treated groups compared with control. Hormone treatments did not affect the relative fecundity. Slight differences were found in ovarian fluid pH between group A and D (P < 0.05). Except for group C, egg size was significantly reduced (P < 0.01) in hormonally synchronized groups compared with the control. Survival to the eyed stage of eggs from females in groups A, B and C was significantly lower (P < 0.01) than in groups D and E. Per cent hatched alevins was lower (P < 0.01) in groups A, B and C than in groups D and E. We conclude that synchronization of ovulation in northern whitefish can be induced by a single acute injection of [D‐Arg⁶Pro⁹NEt]‐sGnRHa at 25 μg kg^?1 BW if given near the natural spawning time determined by water temperature below 2°C.