Comparative ultrastructural studies on the prolactin cells in the pituitary of nine-spined sticklebacks,Pungitius pungitius L., with and without adenohypophyseal cysts

Summary The ultrastructure of the prolactin cells in nine-spined sticklebacks (Pungitius pungitius L.) with and without large pituitary cysts was compared. Prolactin cells from animals with cysts were less active as regards hormone synthesis and release than those from animals without cysts. Rough endoplasmic reticulum, mitochondria, Golgi apparatus and exocytotic figures were all more prominent in animals without cysts. Lysosomes and mature secretory granules were more plentiful in animals with cysts. The correlation between inactive prolactin cells and pituitary cysts argues against the hypothesis that cysts store prolactin released by holocrine secretion. It is more likely that the origin of cysts from degenerating cells signals extreme inactivity in the rostral pars distalis.     

Factors Associated with the Seasonal Variation in the Incidence of Pituitary Cysts in Nine-Spined Sticklebacks,Pungitius pungitius L.

Cysts among the prolactin cells in the pituitary gland of the nine-spined stickleback, Pungitius pungitius L., are most common during and shortly after periods of high mortality that mark the end of the reproductive season. Fish are oldest at this time of year. Nevertheless, cysts may occur in breeding as well as spent fish, and their presence does not relate to changes in oocyte development. They can be induced by starving animals in February, but not in September—October. Starved fish always have small prolactin cells and nuclei. It is suggested that cysts form in older fish in response to stress, be this natural or induced. No correlation was noticed between the condition factor of fish caught in the wild, and the incidence of pituitary cysts.     

The origin of pituitary cysts in the rostral pars distalis of the nine-spined stickleback,Pungitius pungitius L.

Summary Pituitary cysts in the nine-spined stickleback, Pungitius pungitius L., develop next to blood vessels passing through the prolactin zone of the rostral pars distalis to the connective tissue capsule at its periphery. Cysts were most frequent when pituitaries were large compared with body lengths. However, the incidence of cysts could not be closely related to body length alone. As the rostral pars distalis is more highly vascularised in fish with large pituitaries, and as there was no evidence of accumulating secretion around these blood vessels, it is suggested that cysts develop when vascular demands (or supplies) become excessive. Associated with the greater vascularity of the rostral pars distalis in large pituitaries was a remarkable development of non-granulated cells. Indeed cysts may originate as enlarged intercellular spaces between such cells, as the latter often surround blood vessels. The non-granulated cells are also important in enlarging cyst cavities (by phagocytosing prolactin-cell debris) and perhaps for maintaining their structural integrity. It is suggested that the macrophages within cyst cavities are derived from the non-granulated cells.     

The Response of Prolactin,ACTH, and Growth Hormone Cells in the Pituitary Gland of the Three-Spined Stickleback,: Gasterosteus aculeatus L. form leiurus,to Increased Environmental Salinities

Benjamin, M. 1980. The response of prolactin, ACTH, and growth hormone cells in the pituitary gland of the three-spined stickleback, Gasterosteus aculeatus L. form leiurus, to increased environmental salinities. (Department of Anatomy, University College, Cardiff, Wales, U.K.) — Acta zool. (Stockh.) 61(1): 1–7. The time-sequence of response of the prolactin, ACTH and growth hormone cells in the pituitary gland of the leiurus form of the three-spined stickleback, Gasterosteus aculeatus L., to a transfer from freshwater to seawater, was studied by light microscopy. The appearance of the pituitary was compared with that of animals caught in brackish or sea water. The prolactin cells respond only slowly to seawater by cytological changes interpreted as signs of decreased secretory activity. It is thus suggested that prolactin is unlikely to be important for osmoregulation in this stickleback. The growth hormone cells are more active in all seawater-adapted animals, while the ACTH cells are less active—although only in laboratory experiments. The differing responses of the pituitaries of the leiurus and trachurus forms of G. aculeatus and Pungitius pungitius to high salinities are compared. Even in species whose pituitaries are virtually identical, the response to high salinites may differ widely.     

The sculpture and morphology of postcranial dermal armor plates and associated bones in gasterosteiforms and syngnathiforms inhabiting Estonian coastal waters

Lees, J., Märss, T., Wilson, M. V. H., Saat, T. and ?pilev, H. 2011. The sculpture and morphology of postcranial dermal armor plates and associated bones in gasterosteiforms and syngnathiforms inhabiting Estonian coastal waters. ―Acta Zoologica (Stockholm) 93 : 422–435. Five fish species inhabiting Estonian coastal waters (Gasterosteus aculeatus L., Pungitius pungitius (L.), and Spinachia spinachia (L.) of the order Gasterosteiformes and Syngnathus typhle L. and Nerophis ophidion (L.) of the order Syngnathiformes) are described on the basis of the sculpture and morphology of their postcranial dermal armor plates, as revealed and illustrated by SEM images. This study shows that the shapes of these superficial skeletal elements vary by species as well as by their position on the body, whereas the sculpture on the bones is taxon specific. The detailed features allow the identification of isolated fossil and subfossil remains and show promise for future systematics studies.     

Carotenoids in fish. XXVI. Pungitius pungitius (L.) and Gasterosteus aculeatus L. (Gasterosteidae)

The author investigated the presence of various carotenoids in the different parts of the body of Pungitius pungitius (L.) and Gasterosteus aculeatus L. by means of columnar and thin-layer chromatography. The investigations revealed the presence of the following carotenoids:

in Pungitius pungitius. α-carotene, β-carotene, β-cryptoxanthin, mutatochrome, zeaxanthin and astaxanthin;

in Gasterosteus aculeatus: β-carotene, β-cryptoxanthin, β-carotene epoxide, neothxanthin, canthaxanthin, mutatochrome, lutein, phoenicoxanthin, zeaxanthin, taraxanthin, tunaxanthin, astaxanthin, astaxanthin ester and α-doradexanthin. The total carotenoid content ranged from 2.229 to 138.504 µg/g wet weight.

     

Integumentary embryony in CMS sunflower line

Formation of integumentary embryos has been revealed in the ovules of the VIR 116 CMS line of sunflower (Helianthus annuus L.) pollinated by H. occidentalis, a perennial wild sunflower species. These embryos are observed within 7–9 days after the pollination in embryo sacs, where the normal fertilization did not occur. The embryos develop from somatic cells of a maternal organism (an integument) and can be one of the reasons of a matroclinal inheritance in interspecies hybrids.     

A test for within‐lake niche differentiation in the nine‐spined sticklebacks (Pungitius pungitius)

Specialization for the use of different resources can lead to ecological speciation. Accordingly, there are numerous examples of ecologically specialized pairs of fish “species” in postglacial lakes. Using a polymorphic panel of single nucleotide variants, we tested for genetic footprints of within‐lake population stratification in nine‐spined sticklebacks (Pungitius pungitius) collected from three habitats (viz. littoral, benthic, and pelagic) within a northern Swedish lake. Analyses of admixture, population structure, and relatedness all supported the conclusion that the fish from this lake form a single interbreeding unit.     

Small-sized euryhaline fish as intermediate hosts of the digenetic trematodeCryptocotyle concavum

Cercariae of the trematodeCryptocotyle concavum, which encyst in skin and/or kidney of sticklebacks and gobies, were studied in the Schlei Fjord (western Baltic Sea). Mean incidence of dermal cysts was 48 % inGasterosteus aculeatus and 37 % inPungitius pungitius. No cysts were found in the kidneys of sticklebacks. While 97 % ofPomatoschistus microps had encysted metacercariae in the kidneys, only 2 % had cysts in the skin.Pomatoschistus minutus, however, showed hardly any cyst infestation of either skin or kidney. InP. microps the intensity of infestation by metacercariae was frequently more than 50 cysts; in contrast, sticklebacks rarely exhibited more than 5 dermal cysts. Infested fish were larger than 10 mm in total length, the incidence rate increasing with growth. Parasitic infestation depends on ambient salinity:C. concavum was not found at salinities below 4 . In contrast to the high incidence in fish, the first hosts — the snailsHydrobia stagnalis andH. neglecta — showed remarkably low infection rates (3 to 5 %). The findings reported are related to the distribution ofC. concavum, the mode of life of infested fish, the feeding habits of the final hosts and the infestation ofP. microps by other parasites. Evidently,P. microps represents an optimal second host forC. concavum.     

Diverse migratory histories in a brackish water type of the ninespine stickleback, <Emphasis Type="Italic">Pungitius pungitius</Emphasis>

The life history in a brackish water type of the ninespine stickleback, Pungitius pungitius, was studied by examining the strontium (Sr) and calcium (Ca) concentrations in the otoliths. The fluctuating patterns of Sr/Ca ratios along the life history transect in the otoliths varied widely among fish in spite of their identification as brackish water type as estimated by morphological characteristics. More than 70% fish showed the intermediate otolith Sr/Ca ratio throughout, averaging 5.23–7.71 × 10⁻³. Besides this brackish water resident life history type of P. pungitius, other sticklebacks had anadromous (25%) and freshwater amphidromous (2.5%) life history types. These findings clearly indicate that the migration of the ninespine stickleback between fresh and sea waters is obligatory but facultative having an ability to utilize the full range of salinity in its life history.     

Parasites of the nine-spined stickleback Pungitius pungitius (L)

Eleven species of parasite are reported from the nine-spined stickleback Pungitius pungitius (L). Nine of these, namely, Trichodina domerguei, T. tenuidens, Vorticella sp., Epistylis?, Diplostomum spathaceum, Cryptocotyle lingua, Proteocephalus filicollis, Schistocephalus solidus and glochidia of Anodonta cygnea, are new host records for Great Britain. Gyrodactylus spp. and Thersitina gasterostei were also found.     

Effects of water-level regulation on the nearshore fish community in boreal lakes

The fish community in the littoral areas of eight regulated lakes and five reference lakes in Finland was sampled by electrofishing. No significant effect of winter drawdown on species richness was recorded across lakes. Total fish density for stony bottoms of the regulated and reference lakes averaged 19.3 and 32.7 individuals per 100 m², respectively, but this difference was not statistically significant. The combined proportion of littoral fish species, including minnow (Phoxinus phoxinus), bullhead (Cottus gobio), alpine bullhead (Cottus poecilopus), nine-spined stickleback (Pungitius pungitius), and stone loach (Barbatula barbatula), supplemented with zoobenthos feeders ruffe (Gymnocephalus cernuus) and young burbot (Lota lota), was much lower in the regulated lakes than in the reference lakes. Besides winter drawdown, other variables, such as nutrient level and lake size, affected the fish community. Guest editors: K. M. Wantzen, K.-O. Rothhaupt, M. M?rtl, M. Cantonati, L. G.-Tóth & P. Fischer Ecological Effects of Water-Level Fluctuations in Lakes     

Optimal growth strategies under divergent predation pressure

The conditions leading to gigantism in nine‐spined sticklebacks Pungitius pungitius were analysed by modelling fish growth with the von Bertalanffy model searching for the optimal strategy when the model's growth constant and asymptotic fish size parameters are negatively related to each other. Predator‐related mortality was modelled through the increased risk of death during active foraging. The model was parameterized with empirical growth data of fish from four different populations and analysed for optimal growth strategy at different mortality levels. The growth constant and asymptotic fish size were negatively related in most populations. Optimal fish size, fitness and life span decreased with predator‐induced mortality. At low mortality, the fitness of pond populations was higher than that of sea populations. The differences disappeared at intermediate mortalities, and sea populations had slightly higher fitness at extremely high mortalities. In the scenario where all populations mature at the same age, the pond populations perform better at low mortalities and the sea populations at high mortalities. It is concluded that a trade‐off between growth constant and asymptotic fish size, together with different mortality rates, can explain a significant proportion of body size differentiation between populations. In the present case, it is a sufficient explanation of gigantism in pond P. pungitius.     

Feeding activities of two euryhaline small-sized fish in a western Baltic brackish fjord

The daily food intake and feeding activities of the common gobyPomatoschistus microps (Krøyer) and the nine-spined sticklebackPungitius pungitius (L.) were investigated in the brackish Schlei fjord. At the investigation site of Olpenitz, salinities varied between 11 and 15‰, and water temperatures between 5 and 18°C during the period of in-situ experiments in 1981 and 1982. Common gobies sometimes attained a density of more than 100 individuals per square metre, nine-spined sticklebacks as much as 18 individuals per square meter. Their food changed depending on the supply of plankton or benthos. Regarding numbers, their food consisted mainly of harpacticoids, in springtimes of calanoids; with regard to weight, amphipods, polychaetes or chironomid larvae often prevailed. The total food ingestion, measured by means of its relation to fish weights (fullness index), was highest in spring and summer: 2.3% inP. microps and 2.6% inP. pungitius. Low fullness indices of 0.8% inP. microps and 0.3% inP. pungitius were found during times of low water temperatures. 24-h field investigations revealed that the adultP. microps presented clear diurnal rhythms with highest fullness indices after dawn and a further maximum at dusk. Only young gobies ingested some benthos at night.P. pungitius displayed similar rhythms in which maximum fullness indices varied during the day. This species seems to forage every four hours. At dusk, both species preferred benthos to plankton. The results are discussed with respect to the implications of diurnal and seasonal fluctuations in prey choice by small-sized fish populations.     

Genetic and morphological discrimination of three species of ninespined stickleback Pungitius spp. (Teleostei,Gasterosteidae) in France with the revalidation of Pungitius vulgaris (Mauduyt, 1848)

The taxonomy of French ninespined sticklebacks (Pungitius spp.) has long been controversial. To clarify the taxonomy in this group, we use mitochondrial (COI) and nuclear (RNF213) sequence markers, as well as morphological data. In France, both genetic markers discriminate three evolutionary lineages. Morphological analysis on fresh and type specimens supports the different lineages and the existence of three species in France. Pungitius pungitius, occurring in the North of France and Rhone basin, is characterized by specimens longer than 35 mm SL, by a flat head with a straight or slightly concave snout, typically 9–10 dorsal spines, 10–11 dorsal soft rays, 9–10 anal soft rays, 0–12 scutes on the caudal peduncle with a keel reaching the last anal‐fin ray, longer pelvic fin, post‐dorsal and caudal peduncle lengths, and a slim caudal peduncle (caudal peduncle depth/length 11.8%–21.9%). Pungitius laevis, occurring in France, in the English Channel basins and Loire drainage, differs from the other species by a head rounded with concave snout in specimens longer than 35 mm SL, accentuating the impression of fleshy lips, 0–4 scutes on the caudal peduncle and a higher caudal peduncle depth/length ratio (15.7%–34.5%). Finally, Pungitius vulgaris, endemic to the Vienne River and rivers of south‐western France as far north as the Garonne estuary, is differentiated by a rounded head with a straight or slightly convex snout, the absence of scutes on the caudal peduncle and by having 11 pectoral‐fin rays. Our data confirm the existence of a hybridization zone in the North of France between P. pungitius and P. laevis. As a result, Pungitius lotharingus is invalid, as it was described based on hybrid specimens. A lectotype for P. laevis was designated because the syntypes included hybrids. This revision provides new perspectives for evolutionary biology studies and will have consequences for Pungitius conservation in France.     

Species–environment relationships of fish and map‐based variables in small boreal streams: Linkages with climate change and bioassessment

Species–environment relationships were studied between the occurrence of 13 fish and lamprey species and 9 mainly map‐based environmental variables of Finnish boreal small streams. A self‐organizing map (SOM) analysis showed strong relationships between the fish species and environmental variables in a single model (explained variance 55.9%). Besides basic environmental variables such as altitude, catchment size, and mean temperature, land cover variables were also explored. A logistic regression analysis indicated that the occurrence probability of brown trout, Salmo trutta L., decreased with an increasing percentage of peatland ditch drainage in the upper catchment. Ninespine stickleback, Pungitius pungitius (L.), and three‐spined stickleback, Gasterosteus aculeatus L., seemed to benefit from urban areas in the upper catchment. Discovered relationships between fish species occurrence and land‐use attributes are encouraging for the development of fish‐based bioassessment for small streams. The presented ordination of the fish species in the mean temperature gradient will help in predicting fish community responses to climate change.     

Morphological divergence of North‐European nine‐spined sticklebacks (Pungitius pungitius): signatures of parallel evolution

Parallel evolution is characterised by repeated, independent occurrences of similar phenotypes in a given habitat type, in different parts of the species distribution area. We studied body shape and body armour divergence between five marine, four lake, and ten pond populations of nine‐spined sticklebacks [Pungitius pungitius (Linnaeus, 1758)] in Fennoscandia. We hypothesized that marine and lake populations (large water bodies, diverse fish fauna) would be similar, whereas sticklebacks in isolated ponds (small water bodies, simple fish fauna) would be divergent. We found that pond fish had deeper bodies, shorter caudal peduncles, and less body armour (viz. shorter/absent pelvic spines, reduced/absent pelvic girdle, and reduced number of lateral plates) than marine fish. Lake fish were intermediate, but more similar to marine than to pond fish. Results of our common garden experiment concurred with these patterns, suggesting a genetic basis for the observed divergence. We also found large variation among populations within habitat types, indicating that environmental variables other than those related to gross habitat characteristics might also influence nine‐spined stickleback morphology. Apart from suggesting parallel evolution of morphological characteristics of nine‐spined sticklebacks in different habitats, the results also show a number of similarities to the evolution of three‐spined stickleback (Gasterosteus aculeatus Linnaeus, 1758) morphology. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 403–416.     

Life history variation among years in a brackish water type of the Ninespine Stickleback, <Emphasis Type="Italic">Pungitius pungitius</Emphasis>

The life history variation in a brackish water type of the Ninespine Stickleback Pungitius pungitius collected over 3 years was studied by examining the strontium (Sr) and calcium (Ca) concentrations in the otoliths. The life histories showed the fish were either brackish water residents or anadromous throughout those years. 74–78% of the fish had the former life history and 22–26% the latter. The composition of each life history was constant throughout the 3 years, and the occurrence of the “anadromous” novel life history found previously is a general phenomenon in this species.     

Reproductive tactics of four sympatric sticklebacks (Gasterosteidae)

Synopsis The life history traits (longevity, growth, age at maturity, fecundity, egg size and length of breeding season) were compared for four sympatric sticklebacks (Gasterosteidae). Significant differences occurred for growth, longevity, age at maturity and length of breeding season, but not for egg diameters. Differences in life history traits are interpreted in terms of mechanisms permitting the coexistence of these closely-related territorial fish. Egg numbers were lowest inApeltes quadracus and highest inGasterosteus aculeatus withGasterosteus wheatlandi andPungitius pungitius having similar and intermediate fecundities.     

Orosensory food testing in fish: Chronology of behavior

The orosensory food testing behavior in fish was studied using the nine-spined stickleback Pungitius pungitius as an example. The patterns of the chronology of manipulations performed by fish in testing food objects were identified. The existence of two stereotypical patterns of feeding behavior was confirmed, and their new characteristics were obtained. The relationship between the responsiveness to food and the predisposition of fish to feeding was revealed.