A three-spined stickleback, Gasterosteus aculeatus L. from the North Atlantic

An unusual occurrence of a three-spined stickleback, Gasterosteus aculeatus , is described from Ocean Weather Station India in the North-east Atlantic. Previous records of G. aculeatus caught well offshore are summarised and reasons for these occurrences are suggested. Mention is made of the possibility of sticklebacks breeding in the open ocean.     

Phylogeography of isolated freshwater three-spined stickleback Gasterosteus aculeatus populations in the Adriatic Sea basin

Analyses of mitochondrial (mt) DNA and microsatellite variation were carried out to examine the relationships between 10 freshwater populations of three-spined sticklebacks Gasterosteus aculeatus along the eastern coast of the Adriatic Sea. Partial sequences of the mtDNA control region and cytochrome b gene, in addition to 15 microsatellite loci, were used to analyse populations from four isolated river catchments. Results uncovered an Adriatic lineage that was clearly divergent from the European lineage, and confirmed that the most divergent and ancient populations are located within the Adriatic lineage as compared with other European populations. Two northern Adriatic populations formed independent clades within the European mitochondrial lineage, suggesting different colonization histories of the different Adriatic populations. Nuclear marker analyses also indicated deep divergence between Adriatic and European populations, albeit with some discordance between the mtDNA phylogeny of the northern Adriatic populations, further highlighting the strong differentiation among the Adriatic populations. The southern populations within the Adriatic lineage were further organized into distinct clades corresponding to respective river catchments and sub-clades corresponding to river tributaries, reflecting a high degree of population structuring within a small geographic region, concurrent with suggestions of existence of several microrefugia within the Balkan Peninsula. The highly divergent clades and haplotypes unique to the southern Adriatic populations further suggest, in accordance with an earlier, more limited survey, that southern Adriatic populations represent an important reservoir for ancient genetic diversity of G. aculeatus.     

Schooling affects growth in the three-spined stickleback, Gasterosteus aculeatus

Three-spined stickleback fry (mean length 20 mm, mean weight 14 mg) were reared for 14 days alone and in groups of six in a constant per capita water volume. The fish originated from two habitats (rock-pools, sea) of different predation pressure. The fry were fed nauplii of Artemia and commercial aquarium fish food ad libitum. Specific growth rates of solitary and schooling fry differed and were also affected by their origin. The specific growth rates of solitary fry from the sea averaged 1.0% day⁻¹ (length) and 6.0% day⁻¹ (weight) and those of solitary rock-pool fry 1.1 and 6.7%, respectively. For group-reared fish the corresponding values were 1.2% (length) and 6.6% (weight), and 1.3% (length) and 7.6% (weight). The finding that schooling sticklebacks grow faster than solitary ones implies that grouping enhances fitness in stickleback fry under the conditions of our experiments.     

A temperature-dependent growth model for the three-spined stickleback Gasterosteus aculeatus

Specific growth rates of individually reared juvenile three-spined sticklebacks Gasterosteus aculeatus were investigated under laboratory conditions to parameterize a complete temperature-dependent growth model for this species. To test the applicability of experimentally derived optima in growth response rates to natural conditions, the effects of commercial pellets and natural prey on growth rates were investigated. In addition, to test for seasonal effects on growth, laboratory trials were performed in both spring and winter. Growth took place from 5 to 29° C with a temperature for optimum growth reaching a sharp peak at 21° C. Modelled optimal temperature for maximum growth was estimated to be 21.7° C and lower and upper temperatures for growth were estimated to be 3.6 and 30.7° C, respectively. There were no significant differences in growth rates between fish reared on invertebrates or commercial pellets. Seasonal effects on growth were pronounced, with reduced growth rates in the winter despite similar laboratory conditions. On average, 60% higher growth rates were achieved at the optimum temperature in summer compared to the winter. The strong seasonality in the growth patterns of G. aculeatus indicated here reduces the applicability of the model derived in this study to spring and summer conditions.     

Divergence in reproductive ecology of the three-spined stickleback,Gasterosteus aculeatus

An anadromous population (trachurus form) and three freshwater populations (leiurus form) of the three-spined stickleback,Gasterosteus aculeatus, in central Japan were compared with one another in the reproductive traits: body length at maturity, egg size, clutch size, the swell of abdomen and the number of eggs per nest. The anadromous fish which become larger in body size at maturity spawn eggs smaller in size and greater in number than the freshwater fish. The abdomen swell of gravid females expressed by the proportion of abdomen width to body length was greater in the freshwater fish. The anadromous male fish collected a mean of 2,638 eggs with a range of 1,119 to 4,052 eggs from about 6–7 females. In the three freshwater (the Yamayoke, the Tsuya and the Jizo) populations, males must have mated with about 9–22, 7–18 and 4–7 females respectively. It seems that theleiurus form increases its reproductive success by its much more mating opportunities and the parental efforts of nesting males as well as by spawning large eggs. Furthermore, among the freshwater populations, the Jizo one inhabiting the upper stream was clearly larger in body size, in egg size and in clutch size than the Yamayoke and the Tsuya ones which inhabit stable waterbodies with springs. It is possible that the Jizo population adopted the strategy of spawning a few large eggs as an adaptation to its habitat. The causal and functional explanations in reproductive characteristics among the four populations are discussed in regard to differences in the environmental conditions.     

Geographical variations in freshwater populations of the three-spined stickleback,Gasterosteus aculeatus,in Japan

This paper describes an investigation of morphological variation among six freshwater populations (dominantly low-plated morph) of the three-spined stickleback,Gasterosteus aculeatus, in Japan. Such populations are mainly distributed in restricted water areas within a band from Ise to Wakasa Bays, the most constricted part of Honshu Island. According to some differences in morphological variation, i.e. plate morph, number of lateral plates, body size, body shape and body colour, they were classified into two types corresponding to occurrence in the Ibi River and Lake Biwa water systems. The former type was monomorphic dominated by lowplated morphs in spring-fed water, whereas the latter was a dimorphic population consisting of low- and partially-plated morphs. My results suggest that the plate morph type is not correlated with climate nor predation but is related to geological isolation patterns during the course of the landlocking process. Conversely, variations in meristic (number of dorsal and anal fin rays and gill rakers) and morphometric (body shape and body colour) characteristics may have been related to different environmental conditions. This study also provides supporting evidence that the freshwater three-spined stickleback is a distinct species from the anadromous stickleback,G. aculeatus.     

Fecundity of the three-spined stickleback, Gasterosteus aculeatus (L.)

The number of eggs spawned by female three-spined sticklebacks Gasterosteus aculeatus (L.) was highly correlated with the size of the fish expressed either as total length or as weight after spawning. There were no significant correlations between the size of the eggs measured either as wet weight per egg or as dry weight per egg, and the size of the fish. Nor were there significant correlations between the calorific value of the eggs and the length or weight of the fish. Regressions relating egg production to the length and the weight of the fish are given. These results and an analysis of previous studies on the fecundity of the stickleback suggest that variations in fecundity are primarily a reflection of variation in the size of the fish at maturity, and that this size is related to the race of the stickleback and to environmental conditions such as food supply which influence the growth of sticklebacks.     

Ultraviolet light and visual behaviour in the three-spined stickleback, Gasterosteus aculeatus

The ability of various vertebrates to perceive visual information in the ultraviolet region of the electromagnetic spectrum (300-400 nm) is receiving increasing interest. To date, many of these studies have concentrated on the role of ultraviolet perception in mate choice, yet there are several ways in which the ability to perceive ultraviolet information may affect other behaviours. Until now, it has been widely assumed that colour in the three-spined stickleback can be quantified by methods appropriate to the human visual system. However, evidence does exist that suggests that, in some populations, sticklebacks are capable of perceiving ultraviolet wavelengths. Using a behavioural technique, we tested the ability of the stickleback to perceive ultraviolet light under full-spectrum conditions to establish whether such wavelengths are utilised within their normal behavioural repertoire. We tested this ability by assessing whether subjects could locate hidden food in a foraging task where food position was indicated by the position of landmarks. These landmarks differed only in their ultraviolet content, appearing identical when viewed across the human visible region of the spectrum. We found that sticklebacks were able to use ultraviolet perception to locate a foraging patch under full-spectrum conditions.     

Inheritance of vertebral number in the three-spined stickleback (Gasterosteus aculeatus)

Intraspecific variation in the number of vertebrae is taxonomically widespread, and both genetic and environmental factors are known to contribute to this variation. However, the relative importance of genetic versus environmental influences on variation in vertebral number has seldom been investigated with study designs that minimize bias due to non-additive genetic and maternal influences. We used a paternal half-sib design and animal model analysis to estimate heritability and causal components of variance in vertebral number in three-spined sticklebacks (Gasterosteus aculeatus). We found that both the number of vertebrae (h² = 0.36) and body size (h² = 0.42) were moderately heritable, whereas the influence of maternal effects was estimated to be negligible. While the number of vertebrae had a positive effect on body size, no evidence for a genetic correlation between body size and vertebral number was detected. However, there was a significant positive environmental correlation between these two traits. Our results support the generalization-in accordance with results from a review of heritability estimates for vertebral number in fish, reptiles and mammals-that the number of vertebrae appears to be moderately to highly heritable in a wide array of species. In the case of the three-spined stickleback, independent evolution of body size and number of vertebrae should be possible given the low genetic correlation between the two traits.     

The pineal complex of the three-spined stickleback,Gasterosteus aculeatus L.

Summary The pineal complex of the three-spined stickleback (Gasterosteus aculeatus L.) was investigated by light and electron microscopy, as well as fluorescence histochemistry for demonstration of catecholamines and indolamines. The pineal complex of the stickleback consists of a pineal organ and a small parapineal organ situated on the left side of the pineal stalk. The pineal organ, including the entire stalk, is comprised mainly of ependymal-type interstitial cells and photoreceptor cells with well-developed outer segments. Both unmyelinated and myelinated nerve fibres are present in the pineal organ. Nerve tracts from the stalk enter the habenular and posterior commissures. A small bundle of nerve fibres connects the parapineal organ and the left habenular body. The presence of indolamines (5-HTP, 5-HT) was demonstrated in cell bodies of both the pineal body and the pineal stalk, and catecholaminergic nerve fibres surround the pineal complex.     

Interspawning interval of wild female three-spined stickleback Gasterosteus aculeatus in Alaska

The interspawning interval, or spawning frequency, of wild three-spined stickleback, Gasterosteus aculeatus , was estimated using histological examination of postovulatory follicles (POF). Females in Alaskan lakes appeared to have as much as a 48 h delay between ovulation and ovoposition, yet the POF method could still be used to estimate the interspawning interval. In two Alaskan lakes the interspawning interval was estimated to range from 2·2 to 7·8 days among individual female G. aculeatus . These estimates were consistent with the range (2·5 to 5 days) of previous estimates among individual females from laboratory observations of spawning G. aculeatus , as well as anecdotal accounts of spawning intervals reported from wild populations in Canada (5–10 days). The interspawning interval of females increased during the course of the spawning season in Alaska, showing that the majority of female spawning activity occurred during the earliest portion of the approximate 6-week reproductive season. The increased interspawning interval appears to be related to a previously reported decrease in body condition in reproductive females during the breeding season. Thus, female G. aculeatus may be unable to sustain the initial rate of reproduction as energy stores that support the rapid growth of vitellogenic oocytes are depleted.     

Dyadic relationships in nesting males of the three-spined stickleback, Gasterosteus aculeatus

Multidimensional scaling analysis of nest site location, nesting interval, and aggressive and mating interactions revealed that male three-spined sticklebacks, Gasterosteus aculeatus, formed nesting clusters comprising five patterns of dyadic relationships. These patterns were defined by intervening variables such as size difference between the two males, nest site locations, synchronization of nesting cycles (egg presence in the nest), distance between two nests, and nest raiding behaviours. Several patterns of dyadic relationships were integrated among nesting males and their constellations indicated an expression of the social structure. Through an understanding the social structure, I described effects of social context or situations on individual reproductive success.     

Cost of egg production in the three-spined stickleback (Gasterosteus aculeatus L.)

In the interval between successive spawnings, female sticklebacks consumed approximately their own wet weight of Tubifex. On average, for 100 cal of food consumed, 26 cal of eggs, 3 cal of somatic growth and 11 cal of faeces were produced. In the absence of somatic growth, the efficiency of egg production was estimated as nearly 30%, and it was predicted that in the absence of egg production, the efficiency of somatic growth would range from 20% for a fish of 0.8 g wet weight to 3% for 2 g fish.
The weight of eggs produced at a spawning was a function of the weight of the female but not of the amount of food consumed. If food consumption was not sufficient to cover the cost of egg production, the fish lost weight. Total production (the summation of egg production and somatic growth) was a function of the amount of food consumed.     

Genetic evidence for male-biased dispersal in the three-spined stickleback (Gasterosteus aculeatus)

Sex-biased dispersal is capable of generating population structure in nonisolated populations and may affect adaptation processes when selective conditions differ among populations. Intrasexual competition for local resources and/or mating opportunities predicts a male-biased dispersal in polygynous species and a female bias in monogamous species. The patterns of sex-biased dispersal in birds and mammals are well explained by their respective mating systems, but the picture emerging from fish studies is still mixed. Using neutral genetic markers, we investigated whether there is any evidence for sex-biased dispersal among Baltic Sea populations of the three-spined stickleback ( Gasterosteus aculeatus ). The null hypothesis of non sex-biased dispersal was rejected in favour of male-biased dispersal in this species. As the three-spined stickleback has a polygynous mating system, the observed male bias in dispersal is consistent with the hypothesis that local mate competition might drive the observed pattern. Although more research both on the proximate and ultimate causes behind the observed pattern is needed, our results serve as a first step towards understanding patterns of sex-biased dispersal in this species.     

Reduction of sexual dimorphism in stream-resident forms of three-spined stickleback Gasterosteus aculeatus

Sexual dimorphism in geometric body shape and external morphology was compared between marine and stream-resident forms of three-spined stickleback Gasterosteus aculeatus collected from North America and Japan. Some aspects of sexual dimorphism were shared between ecotypes: males had larger heads than females with no significant effect of ecotype on the magnitude of sexual dimorphism. By contrast, a significant sex-by-ecotype interaction was found for body depth. Males tended to have deeper bodies than females in both forms, but the magnitude of sexual dimorphism was reduced in stream-resident forms. Although females were generally larger in standard length and had larger pelvic girdles, significant sexual dimorphism in these traits was not consistently found across populations or ecotypes. These results suggest that some aspects of sexual dimorphism were shared between ecotypes, while others were unique to each population. The results further suggest that ecology may influence the evolution of sexual dimorphism in some external morphological traits, such as body depth.     

Detecting Holocene divergence in the anadromous-freshwater three-spined stickleback (Gasterosteus aculeatus) system

The anadromous-freshwater three-spined stickleback (Gasterosteus aculeatus) system allows for inferring the role of adaptation in speciation with a high level of accuracy because the freshwater ecotype has evolved multiple times from a uniform anadromous ancestor. A cause for concern is that independent evolution among drainages is not guaranteed in areas with a poorly resolved glacial history. This is the case for the west European great rivers, whose downstream valleys flanked the southern limit of the late Pleistocene ice sheet. We tested for independent and postglacial colonization of these valleys hypothesizing that the relationships among anadromous and freshwater sticklebacks correspond to a raceme structure. We compared the reduction in plate number accompanying this colonization to the genetic differentiation using 13 allozyme and five microsatellite loci in 350 individuals. Overall microsatellite differentiation (F(ST) = 0.147) was twice as large as allozyme differentiation (F(ST) = 0.066). Although habitat-specific gene flow may mask the ancestral relationships among both ecotypes, levels of microsatellite differentiation supported the hypothesis of raceme-like divergence, reflecting independent colonizations rather than the presence of two distinct evolutionary clades. Under an infinite alleles model and in the absence of gene flow, the observed freshwater divergence might be reached after 440 (microsatellites) to 4500 (allozymes) generations. Hence, the anadromous-freshwater stickleback system most likely diverged postglacially. We conclude that the reduction in plate number in two freshwater basins probably occurred independently, and that its considerable variation among populations is not in agreement with the time since divergence.     

Studies on the toxicity of cadmium to the three-spined stickleback Gasterosteus aculeatus L.

Cadmium was found to be lethal to sticklebacks at all concentrations from 100.0 to 0.001 mg Cd 1^–1, in water of 103–111 mg 1^–1 hardness as CaCO₃. The pattern of mortality as shown by the time-concentration curve suggests that toxicity is not due to a single mechanism but changes with concentration. Fish were found to accumulate cadmium, the whole body levels increasing from 0.90 μg/g fresh weight at 0.001 mg Cd 1^–1 exposure concentration to 51.0 μg/g at 100 mg Cd 1^–1. The concentration factor was shown to decrease with increasing exposure concentration from 0.51 at 100 mg Cd 1^–1 to 511 at 0.001 mg Cd 1^–1. The plerocercoid parasite Schistocephalus solidus in the host's perivisceral cavity contained less cadmium than the tissues of its host.