Compensatory investment in zebra finches: females lay larger eggs when paired to sexually unattractive males

The classical version of the differential allocation hypothesis states that, when females reproduce over their lifetime with partners that differ in their genetic quality, they should invest more in reproduction with high-quality males. However, in species with lifetime monogamy, such as the zebra finch, partner quality will typically remain the same. In this case, the compensatory investment (CI) hypothesis predicts higher investment for low-quality males, because low genetic quality offspring are more dependent on maternal resources. Here, we show that female zebra finches invested more resources, both in terms of egg volume and yolk carotenoid content, when paired to a low genetic quality male, as judged from his previous ability to obtain extra-pair paternity in aviary colonies. We also found that females deposited slightly larger amounts of testosterone into eggs when paired to a low parental quality male, as judging from his previous success in rearing offspring. This is, to our knowledge, the first experimental support for the CI hypothesis in a species with lifetime monogamy. We stress that in more promiscuous species, the benefits of classical differential allocation may partly be neutralized by the supposed benefits of CI.     

Great tits provided with ad libitum food lay larger eggs when exposed to colder temperatures

The amount of nutrients deposited into a bird egg varies both between and within clutches of the same female. Larger eggs enhance offspring traits, but as a tradeoff, laying large eggs also infers energetic costs to the female. Income breeders usually lay larger eggs later in the season, when temperatures and food availability are higher. Egg size is thus affected by the daily amount of energy available to produce an egg under cold conditions, but it is less well known in how far temperature exerts direct effects on egg size. We show that great tit females Parus major with access to ad libitum food and breeding in climate‐controlled aviaries varied their egg investments. The size of an individual egg was best predicted by mean temperatures one week pre‐laying, with females laying larger, rather than smaller, eggs under colder conditions. Eggs increased in size over the season, but not significantly over the laying sequence. The degree of daily temperature fluctuation did not influence egg size. In addition to a substantial between‐female variation, sisters were more similar to each other than unrelated females, showing that egg size does also reflect heritable intrinsic female properties. Natural variation in egg size is thus not only determined by energy‐limitation, but also due to females allocating more resources to eggs laid in colder environments, thus increasing early survival of the chicks. That the positive correlation between temperature and egg investments that is found in a natural population is reversed under ad libitum food conditions demonstrates that wild great tits tradeoff own condition with survival prospects of their chicks as a function of available food, not ambient temperature.     

Reproductive allocation in Aidablennius sphynx (Teleostei, Blenniidae): females lay more eggs faster when paired with larger males

Individuals are expected to invest more in current reproductive effort when paired with a partner of higher than average quality. Aidablennius sphynx is an external fertilizing fish with paternal care in which females gain direct benefits from spawning with large males, but often 'make do' with small males. In this study, female reproductive responses to large and small males were investigated. When paired with large males, females spawned more eggs per unit time (i.e., at a faster rate). There was no difference in the size of the eggs spawned by females in relation to partner size. By ovipositing at a faster rate, females may have allocated more reproductive effort to large males. In addition, since small males are known to release far fewer sperm than large males, females may have reduced their spawn rate with small males as a tactic to ensure fertilization.     

Dark-bellied brent geese Branta b. bernicla breeding near snowy owl Nyctea scandiaca nests lay more and larger eggs

Several studies have demonstrated that snowy owls Nyctea scandiaca defend an area around their nests against predators, hereby inadvertently creating safe havens for breeding dark-bellied brent geese Branta b. bernicla . However, studies investigating brent goose breeding ecology within the predator-exclusion zones of the snowy owls are absent. In 1999 and 2005, years of high lemming abundance Lemmus sibiricus and Dicrostonyx torquatus , brent geese were primarily breeding in association with snowy owls in the Medusa river catchment on western Taimyr, Russia. Goose nest failure, either as a result of nest abandonment by the adult birds or of nest depredation, increased with increasing distance from the owl nests. Within the brent goose colonies, clutch size as well as egg size increased with decreasing distance from the snowy owl nest, indicating an increasing adult quality closer to owl nests. However, as a result of the abandonment of eggs and goslings, the increasing clutch size did not result in a higher nest success during this study. Apparently brent geese compete for breeding sites close to owl nests, but details of this process remain unknown.     

Differential allocation in a lekking bird: females lay larger eggs and are more likely to have male chicks when they mate with less related males

The differential allocation hypothesis predicts increased investment in offspring when females mate with high-quality males. Few studies have tested whether investment varies with mate relatedness, despite evidence that non-additive gene action influences mate and offspring genetic quality. We tested whether female lekking lance-tailed manakins (Chiroxiphia lanceolata) adjust offspring sex and egg volume in response to mate attractiveness (annual reproductive success, ARS), heterozygosity and relatedness. Across 968 offspring, the probability of being male decreased with increasing parental relatedness but not father ARS or heterozygosity. This correlation tended to diminish with increasing lay-date. Across 162 offspring, egg volume correlated negatively with parental relatedness and varied with lay-date, but was unrelated to father ARS or heterozygosity. Offspring sex and egg size were unrelated to maternal age. Comparisons of maternal half-siblings in broods with no mortality produced similar results, indicating differential allocation rather than covariation between female quality and relatedness or sex-specific inbreeding depression in survival. As males suffer greater inbreeding depression, overproducing females after mating with related males may reduce fitness costs of inbreeding in a system with no inbreeding avoidance, while biasing the sex of outbred offspring towards males may maximize fitness via increased mating success of outbred sons.     

Heat acclimation: phenotypic plasticity and cues to the underlying molecular mechanisms

Acclimation, in contrast to evolutionary adaptation, is a “within life time phenotypic adaptation” resulting in a widening of the dynamic regulatory range of body temperature. Increased efficiency and capacity of the thermoregulatory effectors, and delayed onset of the temperature threshold for thermal injury, contribute to the beneficial effects of acclimation. Reprogrammed gene expression and changes in cellular signaling underlie these responses. Constitutive elevation of the inducible heat shock protein (HSP) 72 kDa provides cytoprotection and delays thermal injury without the need for de novo HSP synthesis upon thermal stress. The time window for evocation of heat acclimation is the early phase of acclimation, the short-term heat acclimation (STHA), with accelerated sympathetic excitability and a drop in plasma thyroxin playing an essential role. An important consequence of thermal acclimation is the development of cross-tolerance between heat acclimation and ischemia/reperfusion insults. The beneficial implications of this feature are discussed.     

Phenotypic accommodation: adaptive innovation due to developmental plasticity

Phenotypic accommodation is adaptive adjustment, without genetic change, of variable aspects of the phenotype following a novel input during development. Phenotypic accommodation can facilitate the evolution of novel morphology by alleviating the negative effects of change, and by giving a head start to adaptive evolution in a new direction. Whether induced by a mutation or a novel environmental factor, innovative morphological form comes from ancestral developmental responses, not from the novel inducing factor itself. Phenotypic accommodation is the result of adaptive developmental responses, so the novel morphologies that result are not "random" variants, but to some degree reflect past functionality. Phenotypic accommodation is the first step in a process of Darwinian adaptive evolution, or evolution by natural selection, where fitness differences among genetically variable developmental variants cause phenotype-frequency change due to gene-frequency change.     

The beneficial acclimation hypothesis versus acclimation of specific traits: physiological change in water-stressed Manduca sexta caterpillars

Do organisms make beneficial physiological adjustments in response to environmental change? We examined this question by measuring the effects of short-term (12-36 h) and long-term (larval lifetime) hydric stress on the tobacco hornworm, Manduca sexta. Larvae were reared from the first instar on low-water (69%) or high-water (80%) artificial diets and then transferred early in the fifth instar to the same or opposite diet (2x2 design). Within the subsequent 36 h, we measured 24-h growth rates and three primary determinants of the water budget: water gain via consumption and water loss via evaporation and defecation. Larvae preexposed to low-water diet grew less rapidly on low-water diet than those switched acutely to low-water diet from high-water diet, showing that larvae preexposed to a particular environment do not necessarily acclimate beneficially to that environment. Our data on water fluxes to and from larvae, however, strongly suggest that water-stressed larvae did make beneficial physiological adjustments. Larvae responded to short-term hydric stress by minimizing rates of water excretion, primarily by increasing rates of rectal water absorption. Larvae responded to chronic water stress by significantly reducing rates of evaporative water loss; they also showed additional reductions in fecal water excretion, but these decreases were due to lowered consumption and not to further increases in rate of rectal water absorption. This mismatch between maladaptive acclimation of organismal performance and beneficial adjustment of suborganismal traits can be reconciled by recognizing that organismal physiology is hierarchical: fitness-related performance traits represent the aggregate outcome of numerous, more mechanistic physiological traits. Although chronic exposure to an environment may depress the aggregate effect of these mechanistic traits on performance, organisms are not precluded from making beneficial adjustments to individual traits contributing to performance.     

Adult neural stem cells: plasticity and developmental potential.

Stem cells play an essential role during the processes of embryonic tissue formation and development and in the maintenance of tissue integrity and renewal throughout adulthood. The differentiation potential of stem cells in adult tissues has been thought to be limited to cell lineages present in the organ from which they derive, but there is evidence that somatic stem cells may display a broader differentiation repertoire. This has been documented for bone marrow stem cells (which can give rise to muscle, hepatic and brain cells) and for muscle precursors, which can turn into blood cells. The adult central nervous system (CNS) has long been considered incapable of cell renewal and structural remodeling. Recent findings indicate that, even in postnatal and adult mammals, neurogenesis does occur in different brain regions and that these regions actually contain adult stem cells. These cells can be expanded both in vivo and ex vivo by exposure to different combinations of growth factors and subsequently give rise to a differentiated progeny comprising the major cell types of the CNS. Almost paradoxically, adult neural stem cells display a multipotency much broader than expected, since they can differentiate into non-CNS mesodermal-derivatives, such as blood cells and skeletal muscle cells. We review the recent findings documenting this unforeseen plasticity and unexpected developmental potential of somatic stem cells in general and of neural stem cells in particular. To better introduce these concepts, some basic notions on the functional properties of adult neural stem cells will also be discussed, particularly focusing on the emerging role of the microenvironment in determining and maintaining their peculiar characteristics.     

Parthenogenesis in non-rodent species: developmental competence and differentiation plasticity

An oocyte can activate its developmental process without the intervention of the male counterpart. This form of reproduction, known as parthenogenesis, occurs spontaneously in a variety of lower organisms, but not in mammals. However, it must be noted that mammalian oocytes can be activated in vitro, mimicking the intracellular calcium wave induced by the spermatozoon at fertilization, which triggers cleavage divisions and embryonic development. The resultant parthenotes are not capable of developing to term and arrest their growth at different stages, depending on the species. It is believed that this arrest is due to genomic imprinting, which causes the repression of genes normally expressed by the paternal allele. Human parthenogenetic embryos have recently been proposed as an alternative, less controversial source of embryonic stem cell lines, based on their inherent inability to form a new individual. However many aspects related to the biology of parthenogenetic embryos and parthenogenetically derived cell lines still need to be elucidated. Limited information is available in particular on the consequences of the lack of centrioles and on the parthenote's ability to assemble a new embryonic centrosome in the absence of the sperm centriole. Indeed, in lower species, successful parthenogenesis largely depends upon the oocyte's ability to regenerate complete and functional centrosomes in the absence of the material supplied by a male gamete, while the control of this event appears to be less stringent in mammalian cells. In an attempt to better elucidate some of these aspects, parthenogenetic cell lines, recently derived in our laboratory, have been characterized for their pluripotency. In vitro and in vivo differentiation plasticity have been assessed, demonstrating the ability of these cells to differentiate into cell types derived from the three germ layers. These results confirmed common features between uni- and bi-parental embryonic stem cells. However data obtained with parthenogenetic cells indicate the presence of an intrinsic deregulation of the mechanisms controlling proliferation vs. differentiation and suggest their uni-parental origin as a possible cause.     

Lens regeneration in axolotl: new evidence of developmental plasticity

Background

Among vertebrates lens regeneration is most pronounced in newts, which have the ability to regenerate the entire lens throughout their lives. Regeneration occurs from the dorsal iris by transdifferentiation of the pigment epithelial cells. Interestingly, the ventral iris never contributes to regeneration. Frogs have limited lens regeneration capacity elicited from the cornea during pre-metamorphic stages. The axolotl is another salamander which, like the newt, regenerates its limbs or its tail with the spinal cord, but up until now all reports have shown that it does not regenerate the lens.

Results

Here we present a detailed analysis during different stages of axolotl development, and we show that despite previous beliefs the axolotl does regenerate the lens, however, only during a limited time after hatching. We have found that starting at stage 44 (forelimb bud stage) lens regeneration is possible for nearly two weeks. Regeneration occurs from the iris but, in contrast to the newt, regeneration can be elicited from either the dorsal or the ventral iris and, occasionally, even from both in the same eye. Similar studies in the zebra fish concluded that lens regeneration is not possible.

Conclusions

Regeneration of the lens is possible in the axolotl, but differs from both frogs and newts. Thus the axolotl iris provides a novel and more plastic strategy for lens regeneration.

     

Specific versus placebo effects in biofeedback training: a critical lay perspective

Recent interchanges on the question of how to evaluate biofeedback have been cast in terms of a researcher versus clinician dichotomy. This tends to make the arguments ad hominem and focuses attention on minutiae that are of limited general interest. Accordingly, one purpose of the present paper is to state the specific-effects approach to biofeedback evaluation from a critical lay, rather than a research, perspective. The logic of the specific-effects approach to treatment evaluation is first illustrated by a hypothetical example (the Minefield Parable), and it is then suggested that the approach is appropriate for the evaluation of any treatment, be it physical, psychological, or some complex combination. The other purpose of the paper is to further clarify the specific-effects position by responding to some difficulties that have been raised by critics of the position. Some of these difficulties are based on misrepresentations of the position, while others are genuine. However, even for the genuine difficulties, practical solutions are available. The paper concludes that the question of whether a particular class of treatments works is one that is properly raised by the intelligent consumer, and that, for the answer to that question, only the facts, based on adequately controlled clinical studies, will do.     

Specific versus placebo effects in biofeedback training: A critical lay perspective

Recent interchanges on the question of how to evaluate biofeedback have been cast in terms of a researcher versus clinician dichotomy. This tends to make the arguments ad hominem and focuses attention on minutiae that are of limited general interest. Accordingly, one purpose of the present paper is to state the specific-effects approach to biofeedback evaluation from a critical lay, rather than a research, perspective. The logic of the specific-effects approach to treatment evaluation is first illustrated by a hypothetical example (the Minefield Parable), and it is then suggested that the approach is appropriate for the evaluation of any treatment, be it physical, psychological, or some complex combination. The other purpose of the paper is to further clarify the specific-effects position by responding to some difficulties that have been raised by critics of the position. Some of these difficulties are based on misrepresentations of the position, while others are genuine. However, even for the genuine difficulties, practical solutions are available. The paper concludes that the question of whether a particular class of treatments works is one that is properly raised by the intelligent consumer, and that, for the answer to that question, only the facts, based on adequately controlled clinical studies, will do.The preparation of this paper was supported by a grant from the National Science and Engineering Research Council of Canada. I am indebted to Hal Scher and Donna Shulhan for comments on an earlier draft.     

Great tits, Parus major, lay too many eggs: experimental evidence in mid-boreal habitats

This study shows that great tits lay too large clutches in mid‐boreal habitats. First, breeding success, measured with number of fledglings or proportion of eggs that produce fledglings, in northern Finland (65°N) is much poorer than in central and western Europe. Second, brood size manipulations (ca ±30% of the natural mean) revealed that reduced broods produced equal numbers of and larger‐sized fledglings than control and enlarged broods, giving thus the best fitness value for reduced broods. Third, parents of enlarged broods could not adjust (i.e. increase) their feeding effort to the greater number of nestlings. Fourth, extra feeding (about 1/3 of the theoretical maximal needs of the nestlings) during the nestling period resulted in more numerous and larger‐sized fledglings in comparison to control broods. We suggest that the ultimate explanation for the too large clutches is gene flow from the southern population, which prevents local adaptations in the north. Consequently, the main reason for food limitation during the nestling period is that northern great tits apply “southern” decision rules for timing of breeding, clutch size and foraging behaviour. Thus, they tend to breed too early in comparison to the food abundance peak, lay too large clutches in comparison to the level of resources and, perhaps, forage on a too narrow diet (75% caterpillars). Since the late broods that matched the local food abundance peak did not succeed better than the mismatched earlier ones, the most crucial fault of northern great tits seems to be that they overestimate food abundance during peak demands and lay too large clutches. Another explanation for this could be that northern great tits have adopted a brood reduction strategy. However, the long‐term data reveal that years of high breeding success, which would maintain large clutches in the population, are very rare in the north. Therefore, it is unlikely that a brood reduction strategy per se could explain the phenomenon. Instead, it could work together with the gene flow against local adaptation for clutch adjustment.     

Plant Proteus: brown algal morphological plasticity and underlying developmental mechanisms

Brown algae are multicellular photosynthetic marine organisms, ubiquitous on rocky intertidal shores at cold and temperate latitudes. Nevertheless, little is known about many aspects of their biology, particularly their development. Given their phylogenetic distance (1.6 billion years) from other plant organisms (land plants, and green and red algae), brown algae harbor a high, as-yet undiscovered diversity of biological mechanisms governing their development. They also show great morphological plasticity, responding to specific environmental constraints, such as sea currents, reduced light availability, grazer attacks, desiccation and UV exposure. Here, we show that brown algal morphogenesis is rather simple and flexible, and review recent genomic data on the cellular and molecular mechanisms known to date that can possibly account for this developmental strategy.     

Cellular 'neoteny': a possible developmental basis for chromaffin cell plasticity

Adrenal medullary chromaffin cells, SIF cells and sympathetic neurons are derived from the sympatho-adrenal sublineage of the neural crest, and represent a range of cellular phenotypes extending from endocrine to neuronal. It is suggested here that these cell types may represent different stages of developmental 'arrest' along a linear pathway whose endpoint is a cholinergic sympathetic neuron. This model explains the 'transdifferentiation' of mature cells seen in this system as simply a delayed realization of transitions that normally occur between these stages during development. Such a 'linear model' of phenotypic diversification may be applicable to other developing systems that generate closely related but distinct cell types.     

Deer mouse aerobic performance across altitudes: effects of developmental history and temperature acclimation

Aerobic physiology at high altitudes has been studied in many animals. Prior work on laboratory-bred deer mice (a species with a wide altitudinal range) showed depression of aerobic capacity at high altitude, even after acclimation. However, wild deer mice show no reduction in thermogenic performance at high altitude, and performance limits seem to be due to physiological and anatomical adjustments to environmental temperature and not to oxygen availability. We asked whether across-altitude performance differences exist in deer mice after accounting for temperature acclimation (approximately 5 degrees and 20 degrees -25 degrees C) and prenatal and neonatal development altitude (340 vs. 3,800 m). We measured maximal thermogenic oxygen consumption (VO2sum) in cold exposure and ran mice on a treadmill to elicit maximal exercise oxygen consumption (VO2max). We found a 10% reduction in VO2max at 3,800 m compared with that at 340 m; thus, the mice were able to compensate for most of the 37% reduction in oxygen availability at the higher altitude. Development altitude did not affect VO2max. There was no effect of test altitude or development altitude on VO2sum in warm-acclimated animals, but both test and development altitude strongly affected VO2sum in cold-acclimated mice, and compensation for hypoxia at 3,800 m was considerably less than that for exercise.