Should I stay or should I go? : Reproductive and dispersal strategies of site-specific liverwort species

The area of intensively managed forests, in which required conditions for several liverwort species are seldom found, has expanded over the forest landscape during the last century. Liverworts are very sensitive to habitat changes, because they demand continuously moist microclimate. Consequently, about third of the forest liverworts have been classified as threatened or near threatened in Finland. The general objective of this thesis is to increase knowledge of the reproductive and dispersal strategies of the substrate-specific forest bryophytes. A further aim was to develop recommendations for conservation measures for species inhabiting unstable and stable habitats in forest landscape. Both population ecological and genetic methods have been applied in the research. Anastrophyllum hellerianum inhabits spatially and temporally limited substrate patches, decaying logs, which can be considered as unstable habitats. The results show that asexual reproduction by gemmae is the dominant mode of reproduction, whereas sexual reproduction is considerably infrequent. Unlike previously assumed, not only spores but also the asexual propagules may contribute to long-distance dispersal. The combination of occasional spore production and practically continuous, massive gemma production facilitates dispersal both on a local scale and over long distances, and it compensates for the great propagule losses that take place preceding successful establishment at suitable sites. However, establishment probability of spores may be restricted because of environmental and biological limitations linked to the low success of sexual reproduction. Long-lasting dry seasons are likely to result in a low success of sexual reproduction and decreased release rate of gemmae from the shoots, and consequent fluctuations in population sizes. In the long term, the substratum limitation is likely to restrict population sizes and cause local extinctions, especially in small-sized remnant populations. Contrastingly, larger forest fragments with more natural disturbance dynamics, to which the species is adapted, are pivotal to species survival. Trichocolea tomentella occupies stable spring and mesic habitats in woodland. The relatively small populations are increasingly fragmented with a high risk for extinction for extrinsic reasons. The results show that T. tomentella mainly invests in population persistence by effective clonal growth via forming independent ramets and in competitive ability, and considerably less in sexuality and dispersal potential. The populations possess relatively high levels of genetic diversity regardless of population size and of degree of isolation. Thus, the small-sized populations inhabiting stable habitats should not be neglected when establishing conservation strategies for the species and when considering the habitat protection of small spring sites. Restricted dispersal capacity, also on a relatively small spatial scale, is likely to prevent successful (re-)colonization in the potential habitat patches of recovering forest landscapes. By contrast, random short-range dispersal of detached vegetative fragments within populations at suitable habitat seems to be frequent. Thus, the restoration actions of spring and streamside habitats close to the populations of T. tomentella may contribute to population expansion. That, in turn, decreases the harmful effects of environmental stochasticity.

Kin selection, social polymorphism, and reproductive allocation in ants

Social groups are common across animal species. The reasons for grouping are straightforward when all individuals gain directly from cooperating. However, the situation becomes more complex when helping entails costs to the personal reproduction of individuals. Kin selection theory has offered a fruitful framework to explain such cooperation by stating that individuals may spread their genes not only through their own reproduction, but also by helping related individuals reproduce. However, kin selection theory also implicitly predicts conflicts when groups consist of non-clonal individuals, i.e. relatedness is less than one. Then, individual interests are not perfectly aligned, and each individual is predicted to favour the propagation of their own genome over others. Social insects provide a solid study system to study the interplay between cooperation and conflict. Breeding systems in social insects range from solitary breeding to eusocial colonies displaying complete division of reproduction between the fertile queen and the sterile worker caste. Within colonies, additional variation is provided by the presence of several reproductive individuals. In many species, the queen mates multiply, which causes the colony to consist of half-sib instead of full-sib offspring. Furthermore, in many species colonies contain multiple breeding queens, which further dilutes relatedness between colony members. Evolutionary biology is thus faced with the challenge to answer why such variation in social structure exists, and what the consequences are on the individual and population level. The main part of this thesis takes on this challenge by investing the dynamics of socially polymorphic ant colonies. The first four chapters investigate the causes and consequences of different social structures, using a combination of field studies, genetic analyses and laboratory experiments. The thesis ends with a theoretical chapter focusing on different social interactions (altruism and spite), and the evolution of harming traits. The main results of the thesis show that social polymorphism has the potential to affect the behaviour and traits of both individuals and colonies. For example, we found that genetic polymorphism may increase the phenotypic variation between individuals in colonies, and that socially polymorphic colonies may show different life history patterns. We also show that colony cohesion may be enhanced even in multiple-queen colonies through patterns of unequal reproduction between queens. However, the thesis also demonstrates that spatial and temporal variation between both populations and environments may affect individual and colony traits, to the degree that results obtained in one place or at one time may not be applicable in other situations. This opens up potential further areas of research to explain these differences.

Road to Royalty - Transition of Potential Queen to Queen in the Primitively Eusocial Wasp Ropalidia marginata

Ropalidia marginata, a primitively eusocial wasp, is different from typical primitively eusocial species in having docile queens who cannot be using dominance to maintain reproductive monopoly and instead appear to use a pheromone from the Dufour's gland to do so. When a docile queen is removed from her colony, one of the workers (potential queen, PQ) becomes highly aggressive, and if the queen is not returned, gradually loses her aggression and becomes the new docile queen within a few days. We hypothesized that the decrease in aggression of the PQ with time since queen removal should be correlated with her change in ovaries and pheromone profile. Because the Dufour's gland hydrocarbon composition in R.marginata can be correlated with fertility, this also gave us an opportunity to test whether PQ is different from workers in her Dufour's gland hydrocarbons. In this study, we therefore trace the road to royalty in R.marginata, that is, the transition of the PQ during queen establishment, in terms of her ovaries, aggression, and Dufour's gland hydrocarbons. Our study focuses on queen establishment, which is important for understanding how reproductive conflict can be manifested and resolved.

Dung as a potential medium for inter-sexual chemical signaling in Asian elephants (Elephas maximus)

Chemical signaling is a prominent mode of male-female communication among elephants, especially during their sexually active periods. Studies on the Asian elephant in zoos have shown the significance of a urinary pheromone (Z7-12:Ac) in conveying the reproductive status of a female toward the opposite sex. We investigated the additional possibility of an inter-sexual chemical signal being conveyed through dung. Sixteen semi-captive adult male elephants were presented with dung samples of three female elephants in different reproductive phases. Each male was tested in 3 separate trials, within an interval of 1-3 days. The trials followed a double-blind pattern as the male and female elephants used in the trials were strangers, and the observer was not aware of the reproductive status of females during the period of bioassays. Males responded preferentially (P < 0.005), in terms of higher frequency of sniff, check and place behavior toward the dung of females close to pre-ovulatory period (follicular-phase) as compared to those in post-ovulatory period (luteal-phase). The response toward the follicular phase samples declined over repeated trials though was still significantly higher than the corresponding response toward the non-ovulatory phase in each of the trials performed. This is the first study to show that male Asian elephants were able to distinguish the reproductive phase of the female by possibly detecting a pre-ovulatory pheromone released in dung. (C) 2012 Elsevier B.V. All rights reserved.

Reproductive queue without overt conflict in the primitively eusocial wasp Ropalidia marginata

Colonies of the primitively eusocial wasp Ropalidia marginata consist of a single egg layer (queen) and a number of non-egg-laying workers. Although the queen is a docile individual, not at the top of the behavioral dominance hierarchy of the colony, she maintains complete reproductive monopoly. If the queen is lost or removed, one and only one of the workers potential queen (PQ)] becomes hyperaggressive and will become the next queen of the colony. The PQ is almost never challenged because she first becomes hyperaggressive and then gradually loses her aggression, develops her ovaries, and starts laying eggs. Although we are unable to identify the PQ when the queen is present, she appears to be a ``cryptic heir designate.'' Here, we show that there is not just one heir designate but a long reproductive queue and that PQs take over the role of egg-laying, successively, without overt conflict, as the queen or previous PQs are removed. The dominance rank of an individual is not a significant predictor of its position in the succession hierarchy. The age of an individual is a significant predictor, but it is not a perfect predictor because PQs often bypass older individuals to become successors. We suggest that such a predesignated reproductive queue that is implemented without overt conflict is adaptive in the tropics, where conspecific usurpers from outside the colony, which can take advantage of the anarchy prevailing in a queenless colony and invade it, are likely to be present throughout the year.

An integrated assessment of the continued spread and potential impacts of the colonial ascidian, Didemnum sp. A, in U.S. waters

Didemnum sp. A is a colonial ascidian or “sea squirt” of unknown geographic origin. Colonies of Didemnum sp. A were first documented in U.S. waters in 1993 at Damariscotta River, Maine and San Francisco Bay, California. An alarming number of colonies have since been found at several locations in New England and along the West Coast of the contiguous continental United States. Originally believed to be restricted to artificial structures in nearshore habitats, such as ports and marinas, colonies of Didemnum sp. A have also been discovered on a gravel-pavement habitat on Georges Bank at depths of 40-65m. The wide distribution of Didemnum sp. A, the presence of colonies on an important offshore fishing ground, and the negative economic impacts that other species of noninidigenous ascidians have had on aquaculture operations have raised concerns about the potential impacts of Didemnum sp. A. We reviewed the available information on the biology and ecology of Didemnum sp. A and potentially closely related species to examine the environmental and socioeconomic factors that may have influenced the introduction, establishment and spread of Didemnum sp. A in U.S. waters, the potential impacts of this colonial ascidian on other organisms, aquaculture, and marine fisheries, and the possibility that it will spread to other U.S. waters. In addition, we present and discuss potential management objectives for minimizing the impacts and spread of Didemnum sp. A. Concern over the potential for Didemnum sp. A to become invasive stems from ecological traits that it shares with other invasive species, including the ability to overgrow benthic organisms, high reproductive and population growth rates, ability to spread by colony fragmentation, tolerance to a wide range of environmental conditions, apparent scarcity of predators, and the ability to survive in human dominated habitats. At relatively small spatial scales, species of Didemnum and other nonindigenous ascidians have been shown to alter the abundance and composition of benthic assemblages. In addition, the Canadian aquaculture industry has reported that heavy infestations of nonindigenous ascidians result in increased handling and processing costs. Offshore fisheries may also suffer where high densities of Didemnum sp. A may alter the access of commercially important fish species to critical spawning grounds, prey items, and refugia. Because colonial ascidian larvae remain viable for only 12–24hrs, the introduction and spread of Didemnum sp. A across large distances is thought to be predominantly human mediated; hull fouling, aquaculture, and ballast water. Recent studies suggest that colony growth rates decline when temperatures exceed 21 ºC for 7 consecutive days. Similarly, water temperatures above 8 to 10 ºC are necessary for colony growth; however, colonies can survive extended periods of time below this temperature threshold as an unidentified overwintering form. A qualitative analysis of monthly mean nearshore water temperatures suggest that new colonies of Didemnum will continue to be found in the Northeast U.S., California Current, and Gulf of Alaska LMEs. In contrast, water temperatures become less favorable for colony establishment in subarctic, subtropical, and tropical areas to the north and south of Didemnum’s current distribution in cool temperate habitats. We recommend that the Aquatic Nuisance Species Task Force serve as the central management authority to coordinate State and Federal management activities. Five objectives for a Didemnum sp. A management and control program focusing on preventing the spread of Didemnum sp. A to new areas and limiting the impacts of existing populations are discussed. Given the difficulty of eradicating large populations of Didemnum sp. A, developing strategies for limiting the access of Didemnum sp. A to transport vectors and locating newly established colonies are emphasized. (PDF contains 70 pages)

Engineered underdominance as a method of insect population replacement and reproductive isolation

Insect vector-borne diseases, such as malaria and dengue fever (both spread by mosquito vectors), continue to significantly impact health worldwide, despite the efforts put forth to eradicate them. Suppression strategies utilizing genetically modified disease-refractory insects have surfaced as an attractive means of disease control, and progress has been made on engineering disease-resistant insect vectors. However, laboratory-engineered disease refractory genes would probably not spread in the wild, and would most likely need to be linked to a gene drive system in order to proliferate in native insect populations. Underdominant systems like translocations and engineered underdominance have been proposed as potential mechanisms for spreading disease refractory genes. Not only do these threshold-dependent systems have certain advantages over other potential gene drive mechanisms, such as localization of gene drive and removability, extreme engineered underdominance can also be used to bring about reproductive isolation, which may be of interest in controlling the spread of GMO crops. Proof-of-principle establishment of such drive mechanisms in a well-understood and studied insect, such as Drosophila melanogaster, is essential before more applied systems can be developed for the less characterized vector species of interest, such as mosquitoes. This work details the development of several distinct types of engineered underdominance and of translocations in Drosophila, including ones capable of bringing about reproductive isolation and population replacement, as a proof of concept study that can inform efforts to construct such systems in insect disease vectors.

A review of IATTC research on the early life history and reproductive biology of scombrids conducted at the Achotines Laboratory from 1985 to 2005

English: For nearly a century, fisheries scientists have studied marine fish stocks in an effort to understand how the abundances of fish populations are determined. During the early lives of marine fishes, survival is variable, and the numbers of individuals surviving to transitional stages or recruitment are difficult to predict. The egg, larval, and juvenile stages of marine fishes are characterized by high rates of mortality and growth. Most marine fishes, particularly pelagic species, are highly fecund, produce small eggs and larvae, and feed and grow in complex aquatic ecosystems. The identification of environmental or biological factors that are most important in controlling survival during the early life stages of marine fishes is a potentially powerful tool in stock assessment. Because vital rates (mortality and growth) during the early life stages of marine fishes are high and variable, small changes in those rates can have profound effects on the properties of survivors and recruitment potential (Houde 1989). Understanding and predicting the factors that most strongly influence pre-recruit survival are key goals of fisheries research programs. Spanish: Desde hace casi un siglo, los científicos pesqueros han estudiado las poblaciones de peces marinos en un intento por entender cómo se determina la abundancia de las mismas. Durante la vida temprana de los peces marinos, la supervivencia es variable, y el número de individuos que sobrevive hasta las etapas transicionales o el reclutamiento es difícil de predecir. Las etapas de huevo, larval, y juvenil de los peces marinos son caracterizadas por tasas altas de mortalidad y crecimiento. La mayoría de los peces marinos, particularmente las especies pelágicas, son muy fecundos, producen huevos y larvas pequeños, y se alimentan y crecen en ecosistemas acuáticos complejos. La identificación los factores ambientales o biológicos más importantes en el control de la supervivencia durante las etapas tempranas de vida de los peces marinos es una herramienta potencialmente potente en la evaluación de las poblaciones. Ya que las tasas vitales (mortalidad y crecimiento) durante las etapas tempranas de vida de los peces marinos son altas y variables, cambios pequeños en esas tasas pueden ejercer efectos importantes sobre las propiedades de los supervivientes y el potencial de reclutamiento (Houde 1989). Comprender y predecir los factores que más afectan la supervivencia antes del reclutamiento son objetivos clave de los programas de investigación pesquera.

Reproductive biology of female Rikuzen sole (Dexistes rikuzenius)*

The annual ovarian cycle, mode of maturation, age at maturity, and potential fecundity of female Rikuzen sole (Dexistes rikuzenius) from the North Pacific Ocean off the coast of Japan were studied by 1) histological examination of the gonads, 2) measurement and observation of the oocytes, and 3) by otolith aging. The results indicated that ovulation occurs from September to December and peaks between September and October. Vitellogenesis began again soon after the end of the current season. Maturity was divided into eight phases on the basis of oocyte developmental stages. Mature ovaries contained developing oocytes and postovulatory follicles but no recruiting oocytes, indicating that this species has group-synchronous ovaries and is a multiple spawner. Almost all females matured first at an age of 1+ year and spawned every year until at least age 8+ years. Potential fecundity increased exponentially with body length and the most fecund fish had 15 times as many oocytes as the least fecund fish. Potential fecundity and relative fecundity were both positively correlated with age from 1 to 6+ years, but were negatively correlated, probably because of senescence, in fish over 7 years. These results emphasize that the total productivity of a D. rikuzenius population depends not only on the biomass of females older than 1+ but also on the age structure of the population.

Potential Impact of PCB's on Bluefish, Pomatomus saltatrix, Management

Since 1979, anglers along the U.S. Atlantic coast have landed by weight more bluefish, Pomatomus saltatrix, than any other marine species. A fishery management plan has been developed jointly by three fishery management councils and the Atlantic States Marine Fisheries Commission to preserve the bluefish resource. Major objectives of the plan include prevention of recruitment overfishing and reduction in waste of bluefish. In 1985, a Federal survey found PCB concentrations in larger bluefish (over 500 mm fork length) that exceeded the U.S. Food and Drug Administration tolerance level of 2 parts per million. Harvest strategies are presented in this article to protect the reproductive capability of bluefish while minimizing human health risks associated with dietary intake of PCB's.

Reproductive biology of Black Anglerfish (Lophius budegassa) in the northwestern Mediterranean Sea

Gonadal morphology and reproductive biology of the Black Anglerfish (Lophius budegassa) were studied by examining 4410 specimens collected between June 2007 and December 2010 in the northwestern Mediterranean Sea. Ovaries and testes presented traits common among fishes of the order Lophiiformes. Spawning occurred between November and March. Size at first maturity (L50) was 33.4 cm in total length (TL) for males and 48.2 cm TL for females. Black Anglerfish is a total spawner with group-synchronous oocyte development and determinate fecundity. Fecundity values ranged from 87,569 to 398,986 oocytes, and mean potential fecundity was estimated at 78,929 (standard error of the mean [SE] 13,648) oocytes per kilogram of mature female. This study provides the first description of the presence of 2–3 eggs sharing the same chamber and a semicystic type of spermatogenesis for Black Anglerfish. This new information allows for a better understanding of Black Anglerfish reproduction—knowledge that will be useful for the assessment and management of this species.

An experimental assessment of the aquaculture potential of the brown mussel, Modiolus metcalfei

A study was conducted at Banate Bay, Iloilo, from November 1975 to March 1976. Trials were conducted using spat collectors of four types, but no Modiolus metcalfei spat settled on any of the experimental collectors during the entire study period. Instead they attached to the exposed posterior half of the living Modiolus collected for reproductive cycle studies. The aquaculture potential of the brown mussel is considered to be low. Improvement of its production potential should be approached along the line of resource management rather than aquaculture. This management should be aimed at two objectives: (1) maintenance of enough adults on settlement surfaces, and (2) provision of space to allow new recruitments to grow. A possible solution, therefore, is controlled harvesting or thinning after the peak in the settlement season. In this manner, the chances of the mussel bed recovering year after year may be enhanced.

Biomarker responses and reproductive toxicity of the effluent from a Chinese large sewage treatment plant in Japanese medaka (Oryzias latipes)

The present study was conducted to assess the potential toxicity of the effluent from a large sewage treatment plant (GBD-STP) in Beijing. Japanese medakas (Oryzias latipes) at reproduction active period were exposed to a serial of graded concentrations of the effluent or 100 ng l(-1) of 17-alpha-ethinylestradiol (EE2, positive control). Growth, gonadosomatic index (GSI), hepatosomatic index (HSI), reproductive success, induction potency of vitellogenin (VTG) in male fish and that of 7-ethoxyresorufin-o-deethylase activity (EROD) in male fish liver were used as test endpoints. The growth suppression of fish was observed in a dose-dependent manner, resulting in significant differences in both body length and body weight of medaka above 5% effluent. This effluent can inhibit the growth of gonad of medakas and are more sensitive to male than to female. At exposure concentration of 40% and higher, there was an unexpected decrease of HSI values, which may be resulted from sub-lethal toxicity of effluent to fish liver. VTG of plasma in males were induced in all exposure concentration levels, but not in a dose-dependent manner. The concentration of 5% effluent would be the lowest observed adverse effect level (LOAEL) affecting reproductive success when examining fertile individuals, fecundity and fertilization rate. The overt CYP1A response and higher reproductive toxicity may be indicative of low process efficiency of this STP. (c) 2004 Elsevier Ltd. All rights reserved.

Depolarization of sperm membrane potential is a common feature of men with subfertility and is associated with low fertilization rate at IVF

STUDY QUESTION. Are significant abnormalities in outward (K+) conductance and resting membrane potential (Vm) present in the spermatozoa of patients undertaking IVF and ICSI and if so, what is their functional effect on fertilization success? SUMMARY ANSWER. Negligible outward conductance (≈5% of patients) or an enhanced inward conductance (≈4% of patients), both of which caused depolarization of Vm, were associated with a low rate of fertilization following IVF. WHAT IS KNOWN ALREADY. Sperm-specific potassium channel knockout mice are infertile with defects in sperm function, suggesting that these channels are essential for fertility. These observations suggest that malfunction of K+ channels in human spermatozoa might contribute significantly to the occurrence of subfertility in men. However, remarkably little is known of the nature of K+ channels in human spermatozoa or the incidence and functional consequences of K+ channel defects. STUDY DESIGN, SIZE AND DURATION. Spermatozoa were obtained from healthy volunteer research donors and subfertile IVF and ICSI patients attending a hospital assisted reproductive techniques clinic between May 2013 and December 2015. In total, 40 IVF patients, 41 ICSI patients and 26 normozoospermic donors took part in the study. PARTICIPANTS/MATERIALS, SETTING, METHODS. Samples were examined using electrophysiology (whole-cell patch clamping). Where abnormal electrophysiological characteristics were identified, spermatozoa were further examined for Ca2+ influx induced by progesterone and penetration into viscous media if sufficient sample was available. Full exome sequencing was performed to specifically evaluate potassium calcium-activated channel subfamily M α 1 (KCNMA1), potassium calcium-activated channel subfamily U member 1 (KCNU1) and leucine-rich repeat containing 52 (LRRC52) genes and others associated with K+ signalling. In IVF patients, comparison with fertilization rates was done to assess the functional significance of the electrophysiological abnormalities. MAIN RESULTS AND THE ROLE OF CHANCE. Patch clamp electrophysiology was used to assess outward (K+) conductance and resting membrane potential (Vm) and signalling/motility assays were used to assess functional characteristics of sperm from IVF and ICSI patient samples. The mean Vm and outward membrane conductance in sperm from IVF and ICSI patients were not significantly different from those of control (donor) sperm prepared under the same conditions, but variation between individuals was significantly greater (P< 0.02) with a large number of outliers (>25%). In particular, in ≈10% of patients (7/81), we observed either a negligible outward conductance (4 patients) or an enhanced inward current (3 patients), both of which caused depolarization of Vm. Analysis of clinical data from the IVF patients showed significant association of depolarized Vm (≥0 mV) with low fertilization rate (P= 0.012). Spermatozoa with electrophysiological abnormities (conductance and Vm) responded normally to progesterone with elevation of [Ca2+]i and penetration of viscous medium, indicating retention of cation channel of sperm (CatSper) channel function. LIMITATIONS, REASONS FOR CAUTION. For practical, technical, ethical and logistical reasons, we could not obtain sufficient additional semen samples from men with conductance abnormalities to establish the cause of the conductance defects. Full exome sequencing was only available in two men with conductance defects. WIDER IMPLICATIONS OF THE FINDINGS. These data add significantly to the understanding of the role of ion channels in human sperm function and its impact on male fertility. Impaired potassium channel conductance (Gm) and/or Vm regulation is both common and complex in human spermatozoa and importantly is associated with impaired fertilization capacity when the Vm of cells is completely depolarized.

The reproductive biology of the soft shell clam Mya arenaria in Ireland and the possible impacts of climate variability

Little is known about the biology of the softshell clam in Europe, despite it being identified as a potential species to culture for food in the future. Monthly samples of the softshell clam, Mya arenaria, were collected intertidally from Co. Wexford, Ireland, over a period of sixteen months. The mean weight of sampled individuals was 7 4 ± 4 . 9  g and mean length was 8 . 2 ± 0 . 2  cm. Histological examination revealed a female-to-male ratio of 1 : 1.15. In 2010, M. arenaria at this site matured over the summer months, with both sexes either ripe or spawning by August. A single spawning event was recorded in 2010, completed by November. Two unusually cold winters, followed by a warmer-than-average spring, appear to have affected M. arenaria gametogenesis in this area, potentially affecting the time of spawning, fertilisation success, and recruitment of this species. No hermaphrodites were observed in the samples collected, nor were any pathogens observed. Timing of development and spawning is compared with the coasts of eastern North America and with other European coasts.