No evidence for selective follicle abortion underlying primary sex ratio adjustment in pigeons

Primary sex ratio adjustment in birds has been extensively studied, yet the underlying physiological mechanisms are far from understood. Avian females are the heterogametic sex (ZW), and the future sex of the offspring is determined at chromosome segregation during meiosis I, shortly before the oocyte is ovulated. Assuming that the mother can detect the sex of the developing oocyte before ovulation, it has been suggested that a follicle of the un-preferred sex could selectively be induced to become atretic and regress instead of being ovulated (selective follicle abortion). This potential mechanism has been proposed to underlie biased primary sex ratios in birds, including the homing pigeon (Columba livia domestica), which produces a modal clutch size of two eggs. However, without replacement by an additional, already mature follicle, abortion of a preovulatory follicle would most likely result in either reduced clutch sizes or laying gaps, since a not-yet-recruited follicle still needed to undergo the whole maturation phase. In the current study we killed female pigeons, which were adjusting embryo sex of first eggs according to change in body mass. We examined ovaries for signs of follicle abortion but did not find any supporting evidence. All females produced one or two mature follicles but only two out of the 56 experimental birds produced an additional third mature follicle. Therefore, our results do not corroborate the hypothesis that pigeon mothers manipulate primary offspring sex by selectively aborting follicles of the un-preferred sex.     

The interplay between gonadal steroids and immune defence in affecting a carotenoid-dependent trait

The hypothesis that sexual ornaments are honest signals of quality because their expression is dependent on hormones with immune-depressive effects has received ambiguous support. The hypothesis might be correct for those signals that are carotenoid-dependent because the required carotenoid deposition in the signal, stimulated by testosterone, might lower the carotenoid-dependent immune defence of the organism. Two pathways underlying this androgen-dependent honest signaling have been suggested. Firstly, androgens that are needed for ornament expression may suppress immune defence, a cost that only high-quality animals can afford. Alternatively, immune activation may downregulate the production of androgens in low-quality individuals. Which of these alternatives is correct, and to what extent these effects are mediated by the different metabolites of androgens, remain open questions. To provide answers to these questions, we manipulated the levels of testosterone (T), 5α-dihydrotestosterone (DHT), and 17-β-estradiol (E2) in diamond doves Geopelia cuneata, a species in which both sexes exhibit a carotenoid-dependent, androgen-regulated red–orange periorbital ring of bare skin. On the first day of the experiment (day 0), we inserted steroid-releasing implants into groups of birds and on day 14, we subjected half of the birds to an immunological challenge by immunizing them with sheep red blood cells (SRBC). In females, but not in males, androgen but not estradiol treatments reduced antibody production to SRBC. In addition, the immunological challenge reduced redness and size of the trait as well as androgens levels in both sexes and in all treatments. This indicates that an immunological challenge can lower circulating T at the cost of the trait expression. These findings are in accordance with both pathways postulated in the immunocompetence-handicap hypothesis, but do not entirely support the idea that the immunosuppressive effect of androgens yields honest signaling since both T and DHT were not immunosuppressive in males, for which sexual signaling is supposed to be especially important.     

Development of an Ecologic Marine Classification in the New Zealand Region

We describe here the development of an ecosystem classification designed to underpin the conservation management of marine environments in the New Zealand region. The classification was defined using multivariate classification using explicit environmental layers chosen for their role in driving spatial variation in biologic patterns: depth, mean annual solar radiation, winter sea surface temperature, annual amplitude of sea surface temperature, spatial gradient of sea surface temperature, summer sea surface temperature anomaly, mean wave-induced orbital velocity at the seabed, tidal current velocity, and seabed slope. All variables were derived as gridded data layers at a resolution of 1 km. Variables were selected by assessing their degree of correlation with biologic distributions using separate data sets for demersal fish, benthic invertebrates, and chlorophyll-a. We developed a tuning procedure based on the Mantel test to refine the classification's discrimination of variation in biologic character. This was achieved by increasing the weighting of variables that play a dominant role and/or by transforming variables where this increased their correlation with biologic differences. We assessed the classification's ability to discriminate biologic variation using analysis of similarity. This indicated that the discrimination of biologic differences generally increased with increasing classification detail and varied for different taxonomic groups. Advantages of using a numeric approach compared with geographic-based (regionalisation) approaches include better representation of spatial patterns of variation and the ability to apply the classification at widely varying levels of detail. We expect this classification to provide a useful framework for a range of management applications, including providing frameworks for environmental monitoring and reporting and identifying representative areas for conservation.     

Direct male-male competition can facilitate invasion of new colour types in Lake Victoria cichlids

The possibility that disruptive sexual selection alone can cause sympatric speciation is currently a subject of much debate. The initial difficulty for new and rare ornament phenotypes to invade a population, and the stabilisation of the resulting polymorphism in trait and preference make this hypothesis problematic. Recent theoretical work indicates that the invasion is facilitated if males with the new phenotype have an initial advantage in male-male competition. We studied a pair of sympatric incipient species of cichlids from Lake Victoria, in which the red (Pundamilia nyererei) and blue males (P. pundamilia) vigorously defend territories. Other studies suggested that red phenotypes may have repeatedly invaded blue populations in independent episodes of speciation. We hypothesised that red coloration confers an advantage in male-male competition, assisting red phenotypes to invade. To test this hypothesis, we staged contests between red and blue males from a population where the phenotypes are interbreeding morphs or incipient species. We staged contests under both white and green light condition. Green light effectively masks the difference between red and blue coloration. Red males dominated blue males under white light, but their competitive advantage was significantly diminished under green light. Contests were shorter when colour differences were visible. Experience of blue males with red males did not affect the outcome of a contest. The advantage of red over blue in combats may assist the red phenotype to invade blue populations. The apparently stable co-existence of red and blue incipient species in many populations of Lake Victoria cichlids is discussed.Communicated by C. St. Mary     

Experiential Science; Towards an Integration of Implicit and Reflected Practitioner-Expert Knowledge in the Scientific Development of Organic Farming

For further development of organic agriculture, it will become increasingly essential to integrate experienced innovative practitioners in research projects. The characteristics of this process of co-learning have been transformed into a research approach, theoretically conceptualized as “experiential science” (Baars 2007, Baars and Baars 2007). The approach integrates social sciences, natural sciences, and human sciences. It is derived from action research and belongs to the wider field of transdiscliplinary research. In a dialogue-based culture of equality and mutual exchange the principal of a “bottom-up” experiential learning process can be stimulated and fully reflective. It provides an opportunity to develop organic agriculture as multiple best-practices based on transdisciplinary projects, cases studies, and case series. The aim of the article is to describe the methodological characteristics and the theoretical and practical potential of experiential science for research in and development of organic farming. Three characteristic projects are outlined to illustrate the main elements of the methodology: the retrospective reflection on intuitive and experiential knowledge held by farmers; the knowledge derived from on-farm experimentation; the exchange of knowledge and experiences between farming pioneers within a “masterclass” setting. The study concludes that experiential science offers an important philosophical reconciliation process whereby a synthesis of different approaches to research becomes possible in solving real-life problems: quantitative and qualitative, subjective and objective, reductionistic and holistic, practice and science. Recognizing that there are multiple elements contributing to the process of acquiring knowledge, experiential science draws on a broad field of scientific methods thereby integrating the hermeneutic approach of social sciences and the Humanities with the established methods of contemporary natural science.     

Setting information priorities for remediation decisions at a contaminated-groundwater site.

Many sites of contamination due to inappropriate disposal of hazardous materials or wastes have been found. These sites have the potential of damaging the environment and human health and thus need to be evaluated as to whether and what actions should be initiated. In the decision on whether a contaminated site should be subject to management, the knowledge concerning important parameters that would influence the decision will be beneficial to planning of data collection to support the decision. This paper presents a case study of contaminated site located in northern Taiwan, where the groundwater is contaminated by chlorinated hydrocarbons including trichloroethylene (TCE) and tetrachloroethylene (PCE). A site-specific multimedia risk assessment is performed to estimate the total risk resulting from the contamination. In addition, Monte Carlo simulation, rank correlation coefficients, and decision criteria are combined to develop a methodology for assessing the important of parameters in terms of their influence on the decision. It is found that TCE concentration, vegetable yield, deposition interception fraction of vegetables, and plant surface loss constant, are the four parameters important to the decision-making of the case problem.     

An experimental study on the causal relationships between (ecto-)parasites, testosterone and sexual signalling

Bird song is considered to have evolved via sexual selection and should as such honestly signal aspects of the quality of its bearer. To ensure honesty, the immunocompetence handicap hypothesis proposed a dual role of testosterone, having positive effects on sexual signalling but suppressive effects on immune function. However, recent studies showed that it is rather an immune activation that suppresses the androgen production. This reversed chain of causation may significantly alter the pathways, which translate the effects of parasites and pathogens into changes in the expression of male sexual traits. We infested male canaries with Ixodes ricinus tick nymphs to investigate the causal relationships between (ecto-)parasites, testosterone and sexual signalling, here singing behaviour. We focused on flexible song traits, which may quickly reflect changes in the infestation status, and tested whether these effects relate to changes in the plasma testosterone levels or health state. The experimental tick infestation altered the males’ song performance by reducing song consistency, a trait that had previously been identified to reflect male quality. The tick infestation lowered the plasma testosterone levels and had a negative effect on the health status in terms of a reduced hematocrit. Our pathway analysis then revealed that it is the parasite-induced reduction of the plasma testosterone levels but not of the health state that caused the changes in song consistency. Thus, our study supports the view that it is the effect of parasites and immune activation on plasma testosterone levels that generates the trade-off between immunocompetence and sexual signalling.     

An adaptive annual rhythm in the sex of first pigeon eggs

When the reproductive value of male and female offspring varies differentially, parents are predicted to adjust the sex ratio of their offspring to maximize their fitness (Trivers and Willard, Science 179:90–92, 1973). Two factors have been repeatedly linked to skews in avian offspring sex ratio. First, laying date can affect offspring sex ratio when the sexes differ in age of first reproduction, such that the more slowly maturing sex is overproduced early in the season. Second, position of the egg in the laying sequence of a clutch may affect sex ratio bias since manipulating the sex of the first eggs may be least costly to the mother. We studied both factors in two non-domesticated pigeon species. Both the Wood pigeon (Columba palumbus) and the Rock pigeon (Columba livia) have long breeding seasons and lay two-egg clutches. In the field, we determined the sex of Wood pigeon nestlings. In Rock pigeons, housed in captivity outdoors, we determined embryo sex after 3 days of incubation. On the basis of their sex-specific age of first reproduction, we predicted that males, maturing at older age than females, should be produced in majority early and females later in the year. This was confirmed for both species. The bias was restricted to first eggs. Rock pigeons produced clutches throughout the year and show that the sex of the first egg followed an annual cycle. To our knowledge, this study presents the first evidence of a full annual rhythm in adaptive sex allocation in birds. We suggest that this reflects an endogenous seasonal program in primary sex ratio controlled by a preovulatory mechanism.     

Testosterone increases siblicidal aggression in black-legged kittiwake chicks (Rissa tridactyla)

To compete for parental food deliveries nestling birds have evolved diverse behaviors such as begging displays and sibling aggression. Testosterone has been suggested to be an important mechanism orchestrating such competitive behaviors, but evidence is scarce and often indirect. Siblicidal species provide an interesting case in which a clear dominance hierarchy is established and the dominant chicks lethally attack siblings. We experimentally elevated testosterone in chicks of a facultatively siblicidal species, the black-legged kittiwake, Rissa tridactyla, and showed that testosterone-treated chicks were more aggressive toward their sibling than were control chicks. In such facultatively siblicidal species, chicks normally exhibit intense aggression only when threatened by starvation. Indeed, we found that chicks in relatively poorer condition were more aggressive than were chicks in better condition, even among testosterone-treated chicks, suggesting the action of an additional signal modulating aggression. Relatively larger siblings were also more aggressive than were relatively smaller siblings, confirming the importance of size advantage in determining dominance hierarchies within the brood. In addition, testosterone increased aggression toward a simulated predator, indicating that in kittiwakes testosterone can increase aggression in contexts other than siblicide. Testosterone promoted aggression-mediated dominance, which increased begging although testosterone treatment did not have a significant separate effect on begging. Therefore, testosterone production in the kittiwake and most likely other siblicidal species seems an important fitness mediator already early in life, outside the sexual context and not only manifesting itself in aggressive behavior but also in dominance-mediated effects on food solicitation displays toward parents.     

Environmental equity in the vicinity of Amsterdam Airport: The interplay between market forces and government policy

This paper examines the impact of the interplay between market forces and government policy on environmental equity in the vicinity of Amsterdam Airport in the Netherlands. Despite pressure on space and the presence of the international airport, exposure to the higher levels of traffic noise, NO₂ and external safety risks, and accumulation of traffic noise and NO₂ (‘bads’) was rather limited and generally distributed fairly equally among income categories, probably as a result of environmental regulations. However, lower-income categories had less access to lower levels of traffic noise, NO₂ and public green space (‘goods’) than higher-income categories, which may be explained by both historical processes and market forces. Policy makers seem to accept a higher environmental burden on the regional population because of the benefits for the national economy generated by Amsterdam Airport.