A model for environmental sex reversal in fish

A mathematical model is presented which combines genetic XX-female/XY-male sex determination with environmental pressure for phenotypic sex reversal. This may occur when fishes are exposed to endocrine disrupters, specifically masculinization by exposure to androgens and feminization by exposure to estrogens. A generic model is derived for the sex ratio in successive generations and three special cases, with chronic and constant pressure to sex reverse, are discussed in detail. These show that, with extreme environmental pressure to masculinize, the male genotype is at risk of dying out but with less extreme pressure, masculinization will not be detectable since the proportion of phenotypic males becomes one-half. With feminization at any pressure to sex reverse, the male and female genotypes will be maintained in a stable sex ratio in which the proportion of genotypic males exceeds one-half and is close to one-half if YY offspring (eggs) are not viable. In converse, the model is also applicable to the genetic ZZ-male/ZW-female system of sex determination in fish. At present suitable data are not available with which to validate the model, but proposals are made for relevant experimental studies.     

Intra-sexual variation in male reproduction in teleost fish: a comparative approach

The occurrence of intra-sexual variation in reproduction is a widespread phenomenon in teleosts. One such form of variation consists in the occurrence of alternative male types: males that invest resources in mate attraction and males that exploit the investment of the former males, by trying to sneak fertilizations during spawning. These alternative reproductive tactics can be classified according to their plasticity during the life span of the individuals (i.e., fixed vs. sequential vs. reversible). Furthermore, the differences between morphs within a given species may involve a set of different traits, including reproductive behavior, the differentiation of male morphological traits, and the patterns of gonad tissue allocation and the differentiation of gonadal accessory glands. In this paper, we review the available data on four species exhibiting different types of intra-sexual plasticity in reproduction that have been studied in our lab. The data on the proximate mechanisms, androgens and forebrain arginine-vasotocin (AVT), underlying these alternative tactics suggest that between-morph differences in androgen levels, especially in 11-ketotestosterone, are especially present in species where the alternative male types have evolved morphological traits that are tactic-specific (i.e., sexual polymorphisms) and that differences in AVT appear to be related to between-morph differences in the expression of courtship behavior. Therefore, this comparative approach leads us to propose that the different endocrine systems are involved in the differentiation of different sets of traits that make up alternative phenotypes, and that the differentiation of alternative tactics is not controlled by a single endocrine system (e.g., androgens).     

Geographical and taxonomic biases in invasion ecology

Invasive alien species come from most taxonomic groups, and invasion biology is searching for robust cross-taxon generalizations and principles. An analysis of 2,670 papers dealing with 892 invasive species showed that all major groups of invaders are well studied, but that most information on the mechanisms of invasion has emerged from work on a limited number of the most harmful invaders. A strong geographical bias, with Africa and Asia understudied, inhibits a balanced understanding of invasion, because we might be lacking knowledge of specific invasion mechanisms from poorly studied, regionally specific habitats. International cooperation is required to achieve a more geographically balanced picture of biological invasions. Invasive species with the greatest impact are best studied, but more studies of species that are naturalized but not (yet) invasive are needed to improve understanding of the mechanisms acting during the naturalization phase of invasions and leading to successful invasion.     

Genetic variation in sex allocation in a parasitic wasp: variation in sex pattern within sequences of oviposition

In order to maximize their fitness under Local Mate Competition (LMC), arrhenotokous female wasps have to produce a precise sex ratio when encountering hosts. Recent progress in the theory of hymenopterous parasitoid reproduction suggest that they manage to do it by laying male and female eggs in a particular order and that such reproductive strategies are adaptive. Therefore, the determinism of such sequential patterns would be regulated by genetic control on which natural selection could act. To test this hypothesis, sequences of oviposition were recorded in a set ofTrichogramma brassicae Bezdenko (Hymenoptera; Trichogrammatidae) females and in their daughters by providing themEphestia kuehniella Zeller (Lepidoptera; Pyralidae) eggs. In order to describe accurately sex pattern within these oviposition sequences, I present a joined non-parametric and multivariate statistical method. It is shown thatT. brassicae females do not produce male and female eggs in random sequences. Moreover, the way they organize the sequence of the sexes in their progeny seems to be under a strong genetic control. The evolutionary consequences of such results are discussed.     

Enameloid in the teleost fish Lophius

Summary The ultrastructural and chemical differences between mature enameloid and dentine in the teleost fish Lophius were demonstrated using transmission (TEM) and scanning (SEM) electron microscopy, X-ray diffraction analysis, electron diffraction analysis and electron microprobe analysis. An enameloid cap (about 50 m thick) and a thin layer (1 m thick) of collar enameloid were revealed in the scanning (SEM) and transmission (TEM) electron microscopes. Collar enameloid, structurally different from dentine both in SEM and TEM, proved to be less mineralized than the cap as revealed by electron microprobe analysis. The enameloid cap was seen to differ from dentine by (i) its electron density in TEM; (ii) the structural arrangement perpendicular to the underlying dentine, observed in SEM and TEM and corroborated by electron diffraction data, which showed a preferred orientation of enameloid crystallites; (iii) its higher F content decreasing gradually from the apex of the tooth to the junction with dentine and falling suddenly below this junction; and (iv) its lower Mg content. Establishment of a classification of enameloids is proposed, similar to that for dentines, in which selachian enameloid would be a reference standard.

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Résumé La microscopie électronique à transmission, la microscopie électronique à balayage, la diffraction X, la diffraction électronique et l'analyse par microsonde ont prouvé l'existence d'un émailoïde, ultrastructuralement et chimiquement différent de la dentine, chez la baudroie. La microscopie électronique à balayage et la microscopie électronique à transmission ont mis en évidence un chapeau d'émailoïde, d'environ 50 m d'épaisseur, et une couche fine (1 m) d'émailoïde du collet. L'émailoïde du collet, dont l'ultrastructure différente de celle de la dentine a été mise en évidence en microscopie électronique à transmission et en microscopie électronique à balayage, apparaît, après analyse par microsonde, moins minéralisé que le chapeau. Le chapeau d'émailoïde diffère de la dentine par (i) sa densité aux électrons en microscopie électronique à transmission; (ii) sa structure perpendiculaire à celle de la dentine sous-jacente, visible en microscopie électronique à transmission et en microscopie électronique à balayage et confirmée par la diffraction électronique, qui montre une orientation préférentielle des cristaux; (iii) son contenu en fluor plus élevé, qui diminue progressivement de l'apex à la jonction avec la dentine et chute brutalement au-dessous de cette jonction; (iv) sa teneur plus basse en magnésium. Les auteurs proposent que soit établie une classification des émailoïdes analogue à celle des dentines et dans laquelle l'émailoïde de Sélaciens constituerait un standard de référence.

     

Mechanisms of rapid sympatric speciation by sex reversal and sexual selection in cichlid fish

Mechanisms of speciation in cichlid fish were investigated by analyzing population genetic models of sexual selection on sex-determining genes associated with color polymorphisms. The models are based on a combination of laboratory experiments and field observations on the ecology, male and female mating behavior, and inheritance of sex-determination and color polymorphisms. The models explain why sex-reversal genes that change males into females tend to be X-linked and associated with novel colors, using the hypothesis of restricted recombination on the sex chromosomes, as suggested by previous theory on the evolution of recombination. The models reveal multiple pathways for rapid sympatric speciation through the origin of novel color morphs with strong assortative mating that incorporate both sex-reversal and suppressor genes. Despite the lack of geographic isolation or ecological differentiation, the new species coexists with the ancestral species either temporarily or indefinitely. These results may help to explain different patterns and rates of speciation among groups of cichlids, in particular the explosive diversification of rock-dwelling haplochromine cichlids.     

Induction of XY sex reversal by estrogen involves altered gene expression in a teleost, tilapia

To clarify the importance of endogenous estrogens during sex differentiation in a teleost fish, the Nile tilapia, we examined the target events for endogenous estrogens and their role during gonadal sex differentiation. The expression of CYP19a (P450arom) precedes any morphological gonadal sex differentiation. Further to these findings, the treatment of XX fry with non-steroidal aromatase inhibitor (AI), Fadrozole, from seven to 14 days after hatching caused complete sex reversal to functional males. The XX sex reversal induced by AI was rescued completely with simultaneous estrogen treatment. We also found that XY fry treated with estrogen, before the appearance of morphological sex differences, caused complete sex reversal from males to females. Taken together, these results suggest that endogenous estrogens are required for ovarian differentiation. To identify the down-stream gene products of estrogen during ovarian differentiation, we performed subtractive hybridization using mRNA derived from normal and estrogen treated XY gonads. Two out of ten gene products were expressed in germ cells, whereas the others were expressed in somatic cells.     

Stress and interpopulation variation in glycolytic enzyme activity and expression in a teleost fish Fundulus heteroclitus

Northern populations of Fundulus heteroclitus have twofold greater activity of lactate dehydrogenase-B (LDH-B) than southern populations, but exposure to stress increases LDH-B in southern populations, abolishing this difference. To test whether differences in the activity of other hepatic glycolytic enzymes between populations are sensitive to stress, we injected fish with a pharmacological dose of cortisol in coconut oil (400 microg g(-1)) or exposed them to handling stress and measured the activities of all the glycolytic enzymes. At rest, liver phosphofructokinase (PFK) and aldolase (ALD) activities were greater in southern fish, whereas LDH-B activity was greater in northern fish. No other glycolytic enzymes differed in activity between populations in control fish. Cortisol injection and handling stress decreased PFK and ALD and increased LDH activities in the southern but not the northern population, such that the populations no longer differed in the activity of any enzyme following treatment. Unlike Ldh-B mRNA, Pfk and Ald mRNA levels did not parallel enzyme activity, suggesting complex kinetics or regulation at multiple levels. Plasma cortisol did not differ between populations at rest but was significantly different between populations in treated fish. These data suggest that differences in liver enzyme activity may be related to differences in stress hormone physiology between populations.     

No effects of estrogen receptor overexpression on gonadal sex differentiation and reversal in medaka fish

In order to elucidate a possible role of estrogen receptor in the gonadal sex differentiation and the sex reversal with sex steroids, we examined for the formation of testis or ovary in transgenic medaka fish overexpressing the medaka estrogen receptor under the constitutive medaka beta-actin promoter. The transgenic fish underwent the genetically determined gonadal differentiation and showed the same sex-reversal rates as those of wild-type non-transgenic fish after treatments with estrogen and androgen. These results present invaluable data to reconsider the role of estrogen receptor in the gonadal sex determination.     

Photokinesis in the teleost fish Danio malabaricus

Summary Photokinesis as shown by the rapidly moving fish Danio malabaricus was examined in two experiments.In Experiment I, which involved sequences of short periods of light and darkness, the Ss for the most part showed a positive relationship between light intensity and level of activity. This reaction included a well differentiated response to abrupt intensity change, the effect of the transition from darkness to light being marked. Level of activity in both light and dark periods tended to fall; after a certain point, however, it rose again in the light but continued to fall in the darkness.In Experiment II, successive increases in the total area of lighted space available were matched by increases in the total length of time spent there by the Ss. The overall reaction, however, was initially photonegative and became more so as the experiment progressed. The range of most frequent periods of time spent in the lighted area increased gradually, while the length of entries into the shaded area gradually decreased, in the course of the experiment. But for a lighted area the Ss' length of entry was shorter than for the corresponding dark area, the choice of period was more precise, there were fewer longer periods, and the range was less. In the last condition, the Ss divided their time approximately equally between the lighted 7/8 and the shaded 1/8 of the area available. The number of times the Ss crossed from one area to the other almost doubled during the experiment.The author wishes to express his gratitude to Prof. G. Thinès for his help and encouragement. He was supported by a research grant from N. A. T. O., administered through the Science Research Council, London. The findings reported formed part of a study submitted as a doctoral dissertation at the Université de Louvain. The raw data is tabulated in this study and may be consulted at the Institut de Psychologie et des Sciences Pédagogiques. Address after 1 October: Department of Psychology, Bryn Mawr College, Bryn Mawr, Pennsylvania, U.S.A.     

Genome evolution and biodiversity in teleost fish

Teleost fish, which roughly make up half of the extant vertebrate species, exhibit an amazing level of biodiversity affecting their morphology, ecology and behaviour as well as many other aspects of their biology. This huge variability makes fish extremely attractive for the study of many biological questions, particularly of those related to evolution. New insights gained from different teleost species and sequencing projects have recently revealed several peculiar features of fish genomes that might have played a role in fish evolution and speciation. There is now substantial evidence that a round of tetraploidization/rediploidization has taken place during the early evolution of the ray-finned fish lineage, and that hundreds of duplicate pairs generated by this event have been maintained over hundreds of millions of years of evolution. Differential loss or subfunction partitioning of such gene duplicates might have been involved in the generation of fish variability. In contrast to mammalian genomes, teleost genomes also contain multiple families of active transposable elements, which might have played a role in speciation by affecting hybrid sterility and viability. Finally, the amazing diversity of sex determination systems and the plasticity of sex chromosomes observed in teleost might have been involved in both pre- and postmating reproductive isolation. Comparison of data generated by current and future genome projects as well as complementary studies in other species will allow one to approach the molecular and evolutionary mechanisms underlying genome diversity in fish, and will certainly significantly contribute to our understanding of gene evolution and function in humans and other vertebrates.     

Sexual determination and differentiation in teleost fish

The present work reviews the latest information on the cellular, molecular and physiological aspects of sexual determination and differentiation in teleost fish. The group exhibits a large variety of mechanisms of sexual determination. These may be genetic, or depend on environmental conditions such as temperature, pH, and social factors, all of which can influence the proportion of the sexes. Additionally, sex steroids play an important role in the regulation of sexual differentiation. The patterns of gonadal sexual differentiation are diverse, and species may be hermaphroditic or gonochoristic, some of the latter displaying juvenile hermaphroditism. In recent years, several genes involved in the sexual determination and differentiation pathways in vertebrates, particularly in mammals, have also been characterized in teleosts. Conserved as well as diversified functions have been proposed.     

Sacciform cells in the skin of teleost fish

The fine structure of sacciform gland cells of the epidermis is described in the number of species of teleost fish. In some of these the cell type had either not been found, or not recognized as such, before. Some histological and histochemical results are also reported. Despite considerable differences in the histochemistry and in the morphology of the sacciform cells over the range of species studied, some features of the fine structure are constant and can be used as diagnostic characters. The nucleus is peripheral, and there is a large membrane-limited lumen, into which the secretion is released from membrane-bounded vacuoles at the margin of the cytoplasm. It is probable that the secretion originates mainly in channels of endoplasmic reticulum which become swollen to form the vacuoles. Most sacciform cells open at the surface of the skin by an apical pore, but some have not been seen to open. The classification of the various unicellular glands of teleosts is discussed and it is concluded that attempts to categorise them by the nature of the secretion alone are unsatisfactory.     

The Egyptian fruit bat Rousettus aegyptiacus (Chiroptera: Pteropodidae) in the Palaearctic: Geographical variation and taxonomic status

Two metrically defined subspecies have traditionally been recognised within the Palaearctic distribution range of the Egyptian fruit bat Rousettus aegyptiacus; the larger R. a. aegyptiacus in Egypt and the northern part of the Middle East and the smaller R. a. arabicus in the southern Middle East. An extensive material of R. aegyptiacus from all parts of this area, i.e. the Levant (incl. Turkey and Cyprus), Egypt (incl. Sinai), northern Sudan, Yemen, Oman, Iran, and Pakistan, as well as comparative samples from its sub-Saharan range, were tested using both morphological and genetic approaches in order to revise the species’ taxonomic status. The results indicated two possible processes, depending on the method used. Genetic analysis of the mitochondrial genome (nd1 and cytb) indicated low variation (< 2.0% of genetic distance) and lack of geographical structure while morphometric analysis indicated significant metric differences. Two basic size morphotypes were found within the Palaearctic range, with a rather mosaic-like geographical distribution and a lack of clear size distinction between the two categories, though intermediate types were detected. Thus, we suggest that all Palaearctic populations of R. aegyptiacus represent one form, the nominotypical subspecies, which is uniform in genetic traits but plastic in metric traits.     

Neuropharmacology of adrenergic neurons in teleost fish

Although this brief review is based on relatively few types of experiments in few species of teleosts, it is possible to summarize some points of interest regarding the similarities and differences in the mechanisms of adrenergic neurotransmission in fish compared to the higher vertebrates. 1. There is a substantial mixing of cranial autonomic ("parasympathetic") and spinal autonomic ("sympathetic") pathways in the cranial nerves. This close relationship between the two systems and the differences in the nature of the neurons of cranial origin (cholinergic, and non-adrenergic, non-cholinergic) and spinal origin (adrenergic, cholinergic and mixed "polynergic") gives a basis in fish also for a complex pattern of innervation of the various organs. 2. Adrenaline is the major transmitter substance in the adrenergic neurons of most teleosts studied, but there are exceptions within the same species. For instance, in the swimbladder mucosa of the cod, noradrenaline dominates, while adrenaline is the major catecholamine in most other organs innervated by adrenergic neurons. The reasons for the regional differences are not known and further studies of the rate of catecholamine turn-over in the adrenergic neurons of fish are clearly indicated. 3. Adrenoceptors of both the alpha- and the beta-type show great similarities with those of mammals. Some differences in the potencies of certain compounds (e.g., clonidine and methoxamine) exist and receptor binding studies should add valuable information about the adrenoceptors of teleosts. The existence of a subtype of beta-adrenoceptor (beta 2) has been proposed and further work is needed to confirm or deny the applicability of the beta 1/beta 2 adrenoceptor terminology in fish. 4. There appears to be some differences in the mode of action of the so called "indirectly acting amines", such as tyramine, between teleosts and mammals. While the uptake of tyramine into the nerve terminals in mammals appears to take place via the cocaine-sensitive neuronal uptake system which is also responsible for catecholamine uptake (uptake 1), tyramine uptake in cod neurons appears to be via a separate pathway. 5. Presynaptic supersensitivity of the type seen in mammals has also been demonstrated in teleost adrenergic neurons. Both denervation (chemical or surgical) and blockade of the neuronal uptake mechanism by cocaine or desipramine produce this type of supersensitivity, while post-synaptic supersensitivity has so far not been described in teleosts. The effects of removal of the uptake system shows that the uptake process may be as important in teleosts as in mammals in the removal of adrenergic transmitter from the synaptic cleft. 6. In the total picture of adrenergic functions in fish, the circulating catecholamines take a special role...