Satiation,digestive tract evacuation and return of appetite in the European minnow,Phoxinus phoxinus (Cyprinidae) following short periods of pre-prandial starvation

Synopsis In adultPhoxinus phoxinus (Cyprinidae), starvation for 4 or 16 days had no significant effect on the rate of foregut evacuation of the first satiation meal consumed after the starvation period. Starvation for 4 and 16 days had no significant effect on the rate of return of appetite after that first meal. A model based on the rates of foregut evacuation and return of appetite predicted that in the presence of excess food, the foregut contents would represent about 4% of total body weight. This prediction was corroborated experimentally. The effect of the length of starvation on the size of the first meal was not consistent. The results suggest that the hyperphagia noted in minnows starved for 16 days is a consequence of the persistence on succeeding days of the rates of evacuation and return of appetite achieved on the first day of realimentation.     

Genetic variability in the European minnow,Phoxinus phoxinus (L.)

An electrophoretic study of genetic variation at 11 loci was performedfor a population of European minnows, Phoxinus phoxinus (L.). Ten loci, EST-1 ^*, EST-2 ^* EST-3 ^*,GPD-1 ^*,GPD-2 ^*,GPI-1 ^*,GPI-2 ^*,MPI ^*,6PGD ^* and PGM ^* were polymorphic. IDH ^*wasmonomorphic. The mean number of heterozygotic loci over all 176 fish was 3.05 ± 0.104(SE). Observed mean heterozygosity was 0.28±0.058(SE) and expected mean heterozygosity was 0.27±0.054(SE). EST-2 ^*, EST-3 ^* andPGM ^* were not in Hardy-Weinberg equilibrium. Length,condition, parasite numbers or male breeding characters, i.e. red colorationand tubercles, were not influenced by single enzyme loci.     

The genetic structure and microdistribution of shoals of Phoxinus phoxinus, the European minnow

Efficient foraging and a reduction in predation risk have been proposed as reasons for shoal formation. Some behaviours in cyprinid shoals are at first sight altruistic (e.g. predator inspection behaviour, reactions to alarm substance), such that kin selection may have been involved in their evolution. If shoaling behaviour does evolve through kin selection, then genetic differentiation is expected to be greater between shoals than within shoals. Such a hypothesis was tested here by examining shoal integrity and the relatedness of individuals within and between shoals in the European minnow, Phoxinus phoxinus , using nuclear and mitochondrial DNA markers. The breeding structure of 13 minnow shoals collected from Dorset and North Wales, U.K., was examined using allozymes. Genetic affinity within and between shoals was tested using mitochondrial DNA and multi-locus DNA fingerprinting. Shoals consisted of a random assortment of allozyme genotypes, shoal members did not share the same maternal mtDNA lineages and DNA fingerprint profiles were as varied within shoals as between them. The data indicate that it is unlikely that kin selection occurs in P. phoxinus and there is no apparent relationship between shoaling behaviour and genotype distribution in this species.     

Parasitism and Heterozygosity Influence the Secondary Sexual Characters of the European Minnow,Phoxinus phoxinus (L.) (Cyprinidae)

The Hamilton & Zuk hypothesis (Hamilton & Zuk 1982) was tested in the European minnow, Phoxinus phoxinus (L.), where the male breeding characters include tubercles, which are horny growths on the skin of the head, and a bright red abdomen. Two trematode parasite species, neither directly transmitted, were present: Diplostomum phoxini in the brains of 100% of the fish and Macrolecithus papilliger in the guts of 44%. The heterozygosity of the fish at 11 loci was determined. The number of M. papilliger was negatively related to redness and positively related to the size of tubercles. The number of D. phoxini was not related to redness but was also positively related to the size of tubercles. Heterozygosity was positively related to redness and negatively related to the size of tubercles. This is the first report of such inverse relationships with heterozygosity. The number of tubercles was positively related to the size and condition of a fish. We suggest that males unable to develop exaggerated characters to display to females, e.g. a bright red abdomen, have the alternative strategy of emphasizing weapons for use in male-male competition. This effect could explain other reported positive correlations between parasite burden and sexual display.     

Structure of component parasite communities in the grayling, Thymallus thymallus L. (Salmoniformes, Thymallidae), and minnow, Phoxinus phoxinus L. (Cypriniformes, Cyprinidae), from the upper reaches of the Pechora River

A comparative analysis of parasite communities in the grayling and minnow has been performed. These communities differ in the number of constituent species and in the fact that the community of the minnow is dominated by one allogenic species, whereas that of the grayling usually has two dominants, both being autogenic species. The values of species diversity indices characterizing these communities are also different. Differences between the parasite communities of these fish species reflect the position of their hosts in the hydrobiocenosis. Species prevailing in both communities are classified as specialists, which is characteristic of parasite communities of the boreal-piedmont faunal complex. Both these communities consist of three species groups distinguished by their proportions in the total biomass, which indicates that parasitic communities are structured in a certain way and that the distribution of species in them is not random.     

Evidence for morphological and adaptive genetic divergence between lake and stream habitats in European minnows (Phoxinus phoxinus, Cyprinidae)

Natural selection drives local adaptation, potentially even at small temporal and spatial scales. As a result, adaptive genetic and phenotypic divergence can occur among populations living in different habitats. We investigated patterns of differentiation between contrasting lake and stream habitats in the cyprinid fish European minnow (Phoxinus phoxinus) at both the morphological and genomic levels using geometric morphometrics and AFLP markers, respectively. We also used a spatial correlative approach to identify AFLP loci associated with environmental variables representing potential selective forces responsible for adaptation to divergent habitats. Our results identified different morphologies between lakes and streams, with lake fish presenting a deeper body and caudal peduncle compared to stream fish. Body shape variation conformed to a priori predictions concerning biomechanics and swimming performance in lakes vs. streams. Moreover, morphological differentiation was found to be associated with several environmental variables, which could impose selection on body and caudal peduncle shape. We found adaptive genetic divergence between these contrasting habitats in the form of 'outlier' loci (2.9%) whose genetic divergence exceeded neutral expectations. We also detected additional loci (6.6%) not associated with habitat type (lake vs. stream), but contributing to genetic divergence between populations. Specific environmental variables related to trophic dynamics, landscape topography and geography were associated with several neutral and outlier loci. These results provide new insights into the morphological divergence and genetic basis of adaptation to differentiated habitats.     

Comparison of compensatory growth responses of juvenile three-spined stickleback and minnow following similar food deprivation protocols

The compensatory growth responses of individual juveniles of two co- existing species were compared after identical periods of starvation to determine inter-specific similarities and differences. The carnivorous stickleback Gasterosteus aculeatus was compared with the omnivorous minnow Phoxinus phoxinus. Both species experienced 1 or 2 weeks of starvation before being re-fed ad libitum. The two species differed in their response to the starvation periods, with minnows showing a lower weight-specific loss. Both species showed compensatory responses in appetite, growth and to a lesser extent, growth efficiency. Minnows wholly compensated for 1 and 2 weeks of starvation. At the end of the experiment, sticklebacks starved for 2 weeks were still showing a compensatory response and had not achieved full compensation. The compensatory responses of the sticklebacks showed a lag of a week before developing in the re-feeding phase, whereas the response of the minnows was immediate. Analysis of lipid and dry matter concentrations suggested that the compensatory response restored reserve lipids while also bringing the fish back to the growth trajectory of continuously fed fish.     

Opsin-immunoreactive outer segments and acetylcholinesterase-positive neurons in the pineal complex of Phoxinus phoxinus (Teleostei,Cyprinidae)

Summary The pineal complex of the teleost, Phoxinus phoxinus L., was studied light-microscopically by the use of the indirect immunocytochemical antiopsin reaction and the histochemical acetylcholinersterase (AChE) method.Opsin-immunoreactive outer segments of photoreceptor cells were demonstrated in large numbers in all divisions of the pineal end-vesicle and in the pineal stalk. Moreover, they were found in the roof of the third ventricle, adjacent to the orifice of the pineal recess as well as scattered in the parapineal organ. These immunocytochemical observations provide direct evidence of the presence of an opsin associated with a photopigment in the photosensory cells of the pineal and parapineal organs of Phoxinus. By means of the AChE reaction (Karnovsky and Roots 1964) inner segments of pineal photoreceptors, intrinsic nerve cells, several intrapineal bundles of nerve fibers, and a prominent pineal tract were specifically marked. The pineal neurons can be divided into two types: one is located near the pineal lumen, the other near the basal lamina. The latter perikarya bear stained processes directed toward the photoreceptor layer. A rostral aggregation of two types of AChE-positive nerve cells occurs in the ventral wall of the pineal end-vesicle. The main portion of the AChE-positive pineal tract, which lies within the dorsal wall of the pineal stalk, can be followed to the posterior commissure where some of the nerve fibers course laterally. A few AChE-positive pineal nerve fibers run toward the lateral habenular nucleus via the habenular commissure. In the region of the subcommissural organ single AChE-positive neurons accompany the pineal tract. The nerve cells of the parapineal organ exhibit a moderate AChE activity.These findings extend the structural basis for the remarkable light-dependent activity of the pineal organ of Phoxinus phoxinus. To the memory of Professor Karl von Frisch, pioneer and master in the field of photoneuroendocrine systemsThis investigation was supported by grants from the Deutsche Forschungsgemeinschaft to A.O. (Ok 1/24; 1/25: Mechanismen biologischer Uhren) and to H.-W. K. (Ko 758/1; 758–2)On leave from the 2nd Department of Anatomy, SOTE, Budapest, Hungary     

Accumulation and effects of aluminium in the minnow (Phoxinus phoxinus L.) at different pH levels

The accumulation and effects of 150 pg All^-1 on minnows ( Phoxinus phoxinus L.) were studied in soft water at pH 7, 6 and 5. The fish were kept in a flow-through water system for up to 48 days. Addition of aluminium to the water resulted in poor appetite, passivity and a tendency for the fish to gather as far as possible from the aluminium inlet. The mortality was high at pH 5 and also after addition of aluminium at pH 6. Severe lesions were noted in the gills and olfactory organs after exposure to aluminium. The concentration of aluminium in the gills, liver and kidney was increased at low pH. A large proportion of the aluminium was located on the surface of the gill epithelium. The recovery capacity was pronounced when the aluminium exposure was terminated and the pH was increased from 5 to 7. No further mortality occurred and gill and olfactory organ structure returned to near normality in about 36 days.     

Grouping fails to affect the growth and energy budget of a cyprinid, Phoxinus phoxinus (L.)

The effects of grouping on the growth and some components of the energy budget of the minnow, Phoxinus phoxinus (L.), were tested in the laboratory. No effects of grouping were detected on absorption efficiency, the proportion of food energy lost in nitrogenous excretion, maximum rate of food consumption and conversion efficiency. Grouping had no significant effect on either specific growth rate or the dry matter content of individual fish.     

The effects of spinal nerve section on responses of melanophores in the minnow, Phoxinus phoxinus(L.)

A continuous observation apparatus was used to study the responses of Phoxinus phoxinus melanophores to illuminated black/white backgrounds and their reversal. The fish. Although confined, showed maximum melanosome dispersion (MI 5) and maximum melanosome aggregation (MI 1) when exposed to illuminated black and white backgrounds respectively. Melanophores affected by spinal nerve section showed full melanosome dispersion and the affected area appeared as a black band. The affected melanophores marginally and gradually aggregated their melanosomes if the fish was exposed to an illuminated white background for about a week. The responses of these melanophores to illuminated black and white backgrounds and their reversal indicates that the dispersal of their melanosomes in response to a black background is much faster than their aggregation in response to a white background. It is concluded that an active mechanism is involved and possible factors controlling it are discussed.     

Component communities of parasites in the grayling Thymallus thymallus (L.) (Salmoniformes, Thymallidae) and the minnow Phoxinus phoxinus (L.) (Cypriniformes, Cyprinidae) from the Pechora River

Component communities of parasites of Thymallus thymallus and Phoxinus phoxinus are described. The communities differ in the number of species and in the number of dominate species, which is one in P. phoxinus and usually two in T. thymallus. In the first case the dominate species is allogenic, while in the second case dominate species are autogenic. Values of the species diversity indexes are also different for these communities. The differences between parasite communities of T. thymallus and P. phoxinus underline positions of these hosts in hydrobiocenose. Species specialists take first place in the parasite communities both of the hosts, that is characteristic for the parasite communities of the fishes from the boreal submountain faunistic complex. Parasite communities in both of the hosts consist of three groups of species discriminated by the ratio of their biomasses, that suggests forming of the groups by a coordination of the species biomasses.     

Interactions between developmental processes,growth, and food selection in the larvae and juveniles of Rutilus rutilus (L.) (Cyprinidae)

Summary The aim of this study was to assess the effects of developmental events, occurring in fish during the first weeks after hatching, on the quantity and quality of the ingested food and on growth. The investigation was carried out with the larvae and juveniles of Rutilus rutilus, the single cyprinid species occurring in an oligotrophic subalpine lake in Tirol, Austria. Comparison between availability of prey in the water and gut contents suggests that the selection of food by the young fish is strongly influenced by developmental processes. For example, the prevalence of indigestible phytoplankton in the gut of young larvae can be taken as a sign of the not yet fully developed sensory and locomotory capacities of the young fish (El-Fiky et al. 1987). Furthermore, quantitative and qualitative changes in the gut contents correlate strongly with changes in the form and relative length of the gut, but reflect only weakly the availability of prey in the water. In the Seefelder See population of R. rutilus the switch from a phytoplankton to a cladoceran dominated diet is accompanied by an increase in relative growth rate by nearly one order of magnitude (Wieser et al. 1988).     

Putative cholinergic elements in the photosensory pineal organ and retina of a teleost,Phoxinus phoxinus L. (Cyprinidae)

Summary Putative cholinergic neurons in the photosensory pineal organ of a cyprinid teleost, the European minnow, were studied by use of choline acetyltransferase (ChAT) immunocytochemistry and acetylcholinesterase (AChE) histochemistry. Pinealofugally projecting neurons were visualized using retrograde HRP-filling through their cut axons. For comparison, the distribution of choline acetyltransferase immunoreactivity (ChAT-IR) and AChE-positive elements in the retina was investigated.While the distributional patterns of ChAT-IR and strongly AChE-positive perikarya in the retina are similar and may represent the same neuronal population, ChAT-IR and AChE-positive elements in the pineal organ appear to belong to separate populations. In the retina, small- to medium-sized perikarya in the inner nuclear layer, and small perikarya in the ganglion cell layer are ChAT-IR and AChE positive. The entire inner plexiform layer is AChE positive, while only sublaminae 1, 2 and 4 are ChAT-IR. No indication of cholinergic activity was observed in the optic axon layer.In the pineal organ, ChAT-IR is restricted to small perikarya situated rostrally and dorsally in the pineal end-vesicle. AChE-positive neurons are present throughout the pineal end-vesicle and the pineal stalk. The pineal tract (the pinealofugally projecting axons of intrapineal neurons) is strongly AChE positive, but displays no ChAT-IR. The distribution of pinealofugally projecting neurons, labeled with retrogradely transported HRP, is markedly dissimilar to that of the ChAT-IR elements. It is proposed that the photosensory pineal organ transmits photic information to the brain via a non-cholinergic pathway. The possibility that the ChAT-IR neurons represent small local interneurons is discussed in the light of comparative physiological and anatomical findings.     

黄山九龙峰保护区尖头的年龄、生长和繁殖

于2011年5月至2012年4月逐月采集标本, 对黄山九龙峰保护区尖头 (Phoxinus oxycephalus)的年龄、生长和繁殖等生活史特征进行了研究. 尖头 雌、雄鱼性比为0.77:1, 与1:1差异显著. 根据体长频率分布来确定年龄组成,结果显示雌鱼和雄鱼的最大寿命分别为3龄和2龄, 均以1龄和2龄为优势年龄组. 1龄时的体长生长速度最大, 至2龄或3龄逐渐变缓. 雌、雄鱼的初次性成熟年龄均为2龄, 对应体长分别为93.0 mm和76.9 mm. 尖头 繁殖时间为4-7月; 绝对繁殖力较低(174-2151卵粒), 且在2、3年龄组间显著差异, 但相对繁殖力无显著的年龄组间差异. 尖头 的上述生活史特征, 可能体现了对局域环境条件(如营养贫乏、环境动荡及水文周期等)的生态适应性. 尖头 在我国东南部的高海拔地区呈零星分布, 种群数量小, 加之繁殖投入低下, 因此该物种在面临人类干扰和环境变化时极易濒危和灭绝.       

Effects of steroid hormones and other substances on alarm substance cells and mucous cells in the epidermis of the European minnow, Phoxinus phoxinus (L.), and other Ostariophysi (Pisces)

The alarm substance cells and mucous cells of various species of Ostariophysi do not react uniformly to treatment with androgen and other steroid hormones. This indicates that the mechanism controlling formation of alarm substance is not uniform in the Ostariophysi and three exist at least two such mechanisms, one of which is operated by androgens and perhaps other steroid hormones of the gonads. Steroid hormones from the adrenal cortex proved ineffective. In the European minnow, anti-androgen (androcur®, cyproteroneacetate) had an efrect opposite to that of androgen, while androgen treatment upon this species led to an increase in the size of mucous cells and a reduction in the alarm substance cells. The epidermis of androgen-treated European minnows contained neither alarm substance cells nor alarm substance, and skin extracts of such fish proved ineffective in behaviour experiments upon fish schools. The effect of further hormone treatments are discussed and the reactions of the European minnow are compared to those of other species.     

Feeding and growth of larval herring,Clupea harengus,in relation to density of copepod nauplii

Synopsis Feeding and growth rates of 1–3 wk old herring larvae from four different stocks were compared in laboratory experiments (8°C). For most of the larval groups, feeding rate was saturated at nauplii (Acartia tonsa, nauplii stages 3–5) densities over 301^–1 (5 g d.w. 1^–1). Specific growth rate increased asymptotically with nauplii density, and reached about 6% d^–1 at densities over 120 nauplii 1^–1. The growth rates attained in the laboratory were similar to field measured growth rates of similarly aged herring larvae at comparable food densities. Since food particles were homogenously distributed in the laboratory tanks, patches of dense plankton concentrations are, thus, apparently not necessary for larval growth and survival in the sea. Growth efficiency differed between larval groups, with large sized larvae being the most efficient in transforming ingested matter into growth. The difference probably relates to different sizes rather than to the different geographical origins of the larvae.     

Effects of impoundment on populations of bullhead Cottus gobio L. and minnow, Phoxinus phoxinus (L.), in the basin of Cow Green Reservoir

The distribution, growth and reproduction of bullhead, Cottus gobio , L., and minnow, Phoxinus phoxinus (L.), were studied at Cow Green Reservoir, Upper Teesdale, for 10 years following impoundment in 1970. Comparisons were made with pre-impoundment results. The length-for-age of bullheads increased after impoundment. The length-for-age of minnows decreased. In winter both species were confined mainly to deep water or to the stone facing of the earth dam. In summer they were found throughout the reservoir though bullheads were scarce in the littoral zone. Female bullheads became sexually mature at an earlier age (84% at age-group I) than before impoundment (33% at age-group I) but no change was apparent amongst males (67–62% in age-group I). The post-impoundment sex ratio did not differ from 1:1 and individual fecundity did not change after impoundment. There was a significant reduction in individual fecundity of the minnow. The mean instantaneous rate of mortality ( Z ) for bullhead of age-groups II to IV was 0.96 ± 0.34 year⁻¹, compared with 0.85 ± 0.29 year in the Tees before impoundment.     

食物浓度对方形臂尾轮虫发育历期与实验种群增长参数的影响

为探讨富营养化水体中大量发生的轮虫与微藻之间的关系 ,在实验室条件下 ,观测了不同藻类食物浓度下方形臂尾轮虫生命周期中各发育阶段的历时以及种群增长参数的变化。实验选用蛋白核小球藻为食物 ,浓度梯度为 0 1× 10 6、 0 5× 10 6、 1 0× 10 6、 2 0× 10 6、 4 0× 10 6、 8 0× 10 6、 12× 10 6cells/ml。结果表明 :食物浓度对方形臂尾轮虫的胚胎发育时间与繁殖前期历时、产卵量、世代净生殖率、世代时间和种群的内禀增长率均有明显影响。在食物浓度为 2 0× 10 6cells/ml时 ,方形臂尾轮虫的繁殖前期最短 ,产卵量最大 ,世代净生殖率和种群增长率最高 ,世代时间最短。方形臂尾轮虫可在高于其最适食物浓度的较宽的食物浓度范围内 ,保持较高的种群增长率 ,表现出较强的对环境中高浓度藻类的适应     

The occurrence of tumors in the minnow Phoxinus phoxinus (L.), their influence upon the parasite fauna, component community of parasites, and organism of the host

The occurrence of tumors, their influence upon the organism of Phoxinus phoxinus (L.), its parasite fauna, and parasite component community were investigated in the upstream of the Pechora River. According to the data obtained, tumors could occur in the fishes of every age group, but one-year (0+) or two-year (1+) old minnow is affected by tumors more frequently. The tumors lesion extensiveness ranges from 0.02 to 3 %. From 1 to 3 tumors were recorded on one fish specimen. The investigated tumors were in progressive stage (Georgiev, 2000), since the vascular ingrowth and dissemination (in few cases) of the tumors were observed. Tumors are colored in intensive-black and taupe. The taupe tumors usually have a compact capsule at its peripheries, which isolates affected tissue from muscle fibers. In the intensive-black tumors the invasion of tumor cells to the adjacent transversal striated musculature is observed. Distinct symptoms of necrosis are revealed in all slides of the new growths. Blood vessels are formed in most tumors, and the blood flow is recorded before the completion of the vessels forming, that apparently supplies the tumors feeding. Metastases in different organs revealed in several minnow specimens. Tumor affected individuals of the minnow has parasite species complex practically identical (by species list and quantity) with the same of the even-aged unaffected fishes. However, the parasite component communities of the affected individuals are characterized by 4 groups of species, while the parasite component communities of the intact individuals--by 3 groups. The parasite communities of affected and unaffected one-year fishes are similar by the number of the groups of species, but differ in the number of species.