Variable helper effects, ecological conditions, and the evolution of cooperative breeding in the acorn woodpecker

The ecological conditions leading to delayed dispersal and helping behavior are generally thought to follow one of two contrasting scenarios: that conditions are stable and predictable, resulting in young being ecologically forced to remain as helpers (extrinsic constraints and the habitat saturation hypothesis), or that conditions are highly variable and unpredictable, leading to the need for helpers to raise young, at least when conditions are poor (intrinsic constraints and the hard life hypothesis). We investigated how variability in ecological conditions influences the degree to which helpers augment breeder fitness in the cooperatively breeding acorn woodpecker (Melanerpes formicivorus), a species in which the acorn crop, territory quality, and prior breeding experience all vary in ways that have important effects on fitness. We found that the relationship between ecological conditions and the probability that birds would remain as helpers was variable but that helpers generally yielded greater fitness benefits when ecological conditions were favorable, rather than unfavorable, for breeding. These results affirm the importance of extrinsic constraints to delayed dispersal and cooperative breeding in this species, despite its dependence on a highly variable and unpredictable acorn crop. Our findings also confirm that helpers can have very different fitness effects, depending on conditions, but that those effects are not necessarily greater when breeding conditions are unfavorable.     

Robust modeling of additive and nonadditive variation with intuitive inclusion of expert knowledge

We propose a novel Bayesian approach that robustifies genomic modeling by leveraging expert knowledge (EK) through prior distributions. The central component is the hierarchical decomposition of phenotypic variation into additive and nonadditive genetic variation, which leads to an intuitive model parameterization that can be visualized as a tree. The edges of the tree represent ratios of variances, for example broad-sense heritability, which are quantities for which EK is natural to exist. Penalized complexity priors are defined for all edges of the tree in a bottom-up procedure that respects the model structure and incorporates EK through all levels. We investigate models with different sources of variation and compare the performance of different priors implementing varying amounts of EK in the context of plant breeding. A simulation study shows that the proposed priors implementing EK improve the robustness of genomic modeling and the selection of the genetically best individuals in a breeding program. We observe this improvement in both variety selection on genetic values and parent selection on additive values; the variety selection benefited the most. In a real case study, EK increases phenotype prediction accuracy for cases in which the standard maximum likelihood approach did not find optimal estimates for the variance components. Finally, we discuss the importance of EK priors for genomic modeling and breeding, and point to future research areas of easy-to-use and parsimonious priors in genomic modeling.     

Transient response of aerobic and anoxic activated sludge activities to sudden substrate concentration changes

The state-of-the-art understanding of activated sludge processes as summarized in activated sludge models (ASMs) predicts an instantaneous increase in the biomass activity (which is measured, e.g., by the corresponding respiration rate OUR, NUR, etc.) under sudden substrate concentration changes. Experimental data (e.g., short-term batch respiration experiments under aerobic or anoxic conditions) collected for the calibration of the dynamic models (ASMs) often exhibit a transient phenomenon while attaining maximum activity, which cannot be explained by the current understanding of the activated sludge process. That transient phenomenon exhibits itself immediately upon addition of a substrate source to an endogenously respiring activated sludge sample and it usually takes a few minutes until the activated sludge reaches its maximum possible rate under given environmental conditions. This discrepancy between the state-of-the-art model and the experimental data is addressed in detail in this investigation. It is shown that the discrepancy is not caused by an error in the experimental set-up/data but it is rather due to model inadequacy. Among the hypotheses proposed, it appears that this transient response of the activated sludge most likely results from the sequence of intracellular reactions involved in substrate degradation by the activated sludge. Results from studies performed elsewhere with pure cultures (S. cerevisae and E. coli) support the hypothesis. The transient phenomenon can be described by a dynamic metabolic network model or by a simple first-order model, as adopted in this study. The transient phenomenon occurring in short-term batch respiration experiments is shown to interfere severely with parameter estimation if not modeled properly (2.8%, 11.5%, and 16.8% relative errors [average of three experiments] on Y(H), micro(maxH), and K(S), respectively). Proper modeling of this transient phenomenon whose time constant is on the order of minutes (1 to 3 min) is expected to contribute fundamentally to a better understanding and modeling of Orbal, carousel, and SBR-type treatment plants with fast-alternating process conditions, although such studies are beyond the scope of this report.     

Experimental evidence that age-specific reproductive success is independent of environmental effects

An age-specific improvement in reproductive performance has been reported in many iteroparous breeders. However, whether this is a consequence of intrinsic differences in competence amongst age classes or extrinsic differences in the environment they experience is unclear since the timing of breeding within a season generally also differs with age. To disentangle these effects, we experimentally manipulated the timing of breeding in shags, Phalacrocorax aristotelis. Old and young individuals thus reared their chicks at the same time both early and late in the breeding season. When breeding in the same environmental conditions, old pairs performed consistently better than young pairs. These data clearly demonstrate that the age-related differences in reproductive performance are not a result of environmental effects, but rather a consequence of intrinsic differences in brood rearing capacity.     

THE TIMING OF BIRDS' BREEDING SEASONS

Examination of survival rdtes of nestlings and fledglings of some species show that there is a strong tendency for those young which are hatched earliest in the season to have the greatest chance of surviving to breed. Since natural selection so strongly favours parents who leave many surviving young, the question arises as to why other birds breed later than the date at which they could most successfully raise their young.
It is suggested that the food supply for the breeding females immediately prior to the breeding season may limit their ability to form eggs and the females may thus not be able to lay at the time which would result in young being in the nest at the best time for raising them, but as soon after this time as the female is able to produce her eggs. Not all species are likely to be prevented, by food shortage, from breeding at the best time for raising young and the groups of birds most likely to be affected are discussed.     

Maternal correlates of brood sex ratio variation in the lekking lance-tailed manakin Chiroxiphia lanceolata

Theory predicts that overall population sex ratios should be around parity. But when individual females can receive higher fitness from offspring of one sex, they may benefit by biasing their brood sex ratios accordingly. In lekking species, higher variance in male reproductive success relative to that of females predicts that male offspring gain disproportionately from favorable rearing conditions. Females should therefore produce male-biased broods when they are in a position to raise higher quality offspring: i.e., in better body condition or when they reproduce earlier in the breeding season. To investigate these hypotheses, we studied brood sex ratios of lance-tailed manakins Chiroxiphia lanceolata . We found that overall sex ratios and mean brood sex ratios were not different from random expectation. Brood sex ratios were not related to laying date or female body condition. However, we detected a quadratic relationship between brood sex ratios and maternal age: both young (1–2 years) and old (8+ years) females produced female-biased brood sex ratios. This relationship was most clear in a year also distinguished by early rainy and breeding seasons. We suggest that breeding inexperience in young females and senescence in older females is the most plausible explanation for these results, and that the relationship between female age and brood sex ratio is mediated by environmental conditions.     

Theoretical contributions to biological control success

Ecologists have long tried, with little success, to develop ecological theory for biological control. Biological control illustrates how science often follows, rather than precedes, technological advances. A scientific theory of biological control remains a worthy and achievable goal. We need to (1) combine deductions from mathematical models with rigorous empiricism measuring and modeling the effects of abiotic and biotic environmental drivers on demography and population dynamics of real biological control systems in the field, (2) use a wider range of model systems to explore how population structure, movement, spatial heterogeneity, and external environmental conditions influence population and community dynamics, and (3) combine deductive and inductive approaches to address the day-to-day concerns of biocontrol scientists including how to rear and release a control organism, suppress the target organism, and minimize harm to non-target organisms. Further progress will require more fundamental research in population and community ecology directly relevant to biological control.     

Experimental manipulation of the timing of reproduction in the California vole

Summary The potential role of food and water in determining 1) the timing of the normal breeding season in the California vole, and 2) two frequently occurring exceptions to this normal pattern, are explored by using experimentally manipulated enclosed populations. Hypotheses derived from field observations were tested in the enclosed populations by the manipulation of green food, dry food, water, and vole density. The results of three experiments involving five populations are reported. In all populations densities were maintained at near-normal levels (475 to 1750 per ha). In all three experiments, it was shown that breeding was possible in the summer if sufficient food and water were available, although perhaps with reduced litter sizes and rates of growth and reproductive maturation of young. In two cases, severe weight losses and high mortality were induced by insufficient water. When water was provided, survivors recovered weight and reproductive competency in between five weeks and eight months. In three populations, starvation induced weight losses and mortality. In these cases recovery of the voles took only one month or less. In two of these last, the inadequate food situation was produced during the normally wet season by outbreaks of a European slug. It is concluded that available moisture is the most important factor in triggering both the end and beginning of the normal breeding season. Of course, inadequately fed mice do not breed at any season. Summer (dry season) breeding can occur when food and water conditions are adequate due to locally favorable conditions or very low population densities. Prolonged delays in growth and reproduction following the return of favorable conditions are most likely the consequence of chronic and severe water deprivation.     

An assessment of Gallistel's (2012) rationalistic account of extinction phenomena

Gallistel (2012) asserts that animals use rationalistic reasoning (i.e., information theory and Bayesian inference) to make decisions that underlie select extinction phenomena. Rational processes are presumed to lead to evolutionarily optimal behavior. Thus, Gallistel's model is a type of optimality theory. But optimality theory is only a theory, a theory about an ideal organism, and its predictions frequently deviate appreciably from observed behavior of animals in the laboratory and the real world. That is, behavior of animals is often far from optimal, as is evident in many behavioral phenomena. Hence, appeals to optimality theory to explain, rather than illuminate, actual behavior are misguided.     

Genetics of the guppy as a model for experiment in aquaculture

The guppy, Poecilia reticulata, is one of the most famous tropical ornamental fish in the world. There are many varieties and breeds which have been bred by aqua lists for many years. The guppy has frequently been used as model organism for experiments in fish genetics because of its short life cycle, ease of breeding and reproductive capacity. The laboratory strains which were created from local (Japanese) strains and varieties were used as an experimental genetic model for aquaculture. These laboratory strains were maintained as closed colonies for five to 20 years in our laboratory. One of the roles of model experiments in fish genetics to perform experiments which take a long time in which are commercially important, such as genetic drift, as the effect of inbreeding during the growth and maintenance of population. In this review, we mainly considered the results obtained in our laboratory relating to genetic drift, and the effects of inbreeding, heterosis and inheritance of quantitative traits.     

Influence of Gene Flow and Breeding Tactics on Gene Diversity within Populations

Expressions describing the accumulation of gene correlations within and among lineages and individuals of a population are derived. The model permits different migration rates by males and females and accounts for various breeding tactics within lineages. The resultant equations enable calculation of the probabilistic quantities for the fixation indices, rates of loss of genetic variation, accumulation of inbreeding, and coefficients of relationship for the population at any generation. All fixation indices were found to attain asymptotic values rapidly despite the consistent loss of genetic variation and accumulation of inbreeding within the population. The time required to attain asymptotic values, however, was prolonged when gene flow among lineages was relatively low (less than 20%). The degree of genetic differentiation among breeding groups, inbreeding coefficients, and gene correlations within lineages were found to be primarily functions of breeding tactics within groups rather than gene flow among groups. Thus, the asymptotic value of S. Wright's island model is not appropriate for describing genetic differences among groups within populations. An alternative solution is provided that under limited conditions will reduce to the original island model. The evolution of polygynous breeding tactics appears to be more favorable for promoting intragroup gene correlations than modification of migration rates. Inbreeding and variance effective sizes are derived for populations that are structured by different migration and breeding tactics. Processes that reduce the inbreeding effective population size result in a concomitant increase in variance effective population size.     

Costs of breeding and their effects on the direction of sexual selection

A recent life-history model has challenged the importance of the operational sex ratio and the potential reproductive rates of males and females as the factors most important for the control of sexual selection, arguing that the cost of breeding, interpreted as the probability of dying as a consequence of the current breeding attempt, is the single most important factor that best predicts a mating system. In one species of bushcricket, the mating system can be reversed by resource manipulation. Here, we examine the costs of breeding in this system. Consistent with the model, increased costs of breeding can explain female competition and increased male choosiness under resource limitation. However, this is due to differences in the time required for a breeding attempt, rather than differences in breeding mortality which did not differ between the sexes. In general, males lived longer than females and we discuss the possible reasons behind this pattern of sex-biased non-breeding mortality.     

The breeding biology of equatorial vertebrates: reproduction in the fruit bat, Eidolon helvum, at latitude 0°20'N

The African fruit bat, Eidolon helvum Kerr, is known to occur only south of the Sahara where its distribution is localized. At 0°20' N, this bat exhibits seasonal and synchronized breeding the rhythm of which appears to correspond with the two rainfall peaks typical of Uganda. Quite unexpectedly and unprecedently, this bat exhibits delayed implantation lasting about three months. Evidence for this phenomenon comes from the histological examination of the adult female genital tracts which indicate that mating occurs only in April-June, but implanted embryos are only in evidence in October-November. The male sexual cycle corresponds to such a rhythm and shows a maximum testicular weight at the mating time. Births take place in February-March thus giving a gestation period of about four months. Only one young is born at a time and births occur just before the onset of the higher of the two rainfall peaks, presumably creating favourable conditions for the maximum survival of the young bats when weaned.     

The biotron breeding system: a rapid and reliable procedure for genetic studies and breeding in rice

Oryza sativa is widely used as a model organism for many aspects of research in monocots and cereals. However, it has certain disadvantages as a model species compared with Arabidopsis thaliana, the eudicot species most widely used in plant sciences: first, it has a long cultivation time; and second, it requires considerably more space for growth. Here, we introduce a biotron breeding system, which allows rapid and reliable rice cultivation using a well-equipped artificial environmental chamber. This system involves use of regulation of CO? levels, removal of tillers and embryo rescue to overcome the disadvantages of rice cultivation. The rice cultivars Nipponbare, Koshihikari, Taichung 65 and Kasalath all showed vigorous growth and sufficient seed production in the biotron breeding system with accelerated flowering time. Nipponbare, which was the earliest among these cultivars, flowered at about 50 d after sowing. The life cycle of these plants could be further shortened using an embryo rescue technique on immature seeds at 7 d after pollination, thereby avoiding the lengthy process of seed maturation. Overall, it was possible to shorten the life cycle of Nipponbare to about 2 months under the controlled conditions. Furthermore, controlled crosses, which can be difficult with conventional cultivation methods, were easy to perform as we could control the exact timing of anther dehiscence. Thus, our biotron breeding system offers a valuable new approach to genetic and breeding studies in rice.     

The importance of food quantity and quality for reproductive performance in alpine water pipits (Anthus spinoletta)

Studies relating reproduction to food availability are usually restricted to food quantity, but ignore food quality and the effects of habitat structure on obtaining the food. This is particularly true for insectivorous birds. In this study we relate measures of reproductive success, time of reproduction and nestling size of water pipits (Anthus spinoletta) to biomass, taxonomic composition and nutritional content of available food, and to vegetation structure and distance to feeding sites. Clutch size was positively correlated with the proportion of grass at the feeding sites, which facilitates foraging. This suggests that water pipits adapt their clutch size to environmental conditions. Also, pipits started breeding earlier and produced more fledglings when abundant food and a large proportion of grass were available, probably because these conditions allow the birds to gain more energy in less time. The number of fledglings was positively correlated with the energy content of available food. No significant relationships were found between feeding conditions and nestling size or the time that nestlings took to fledge. This suggests that water pipits do not invest more in individual nestlings when food conditions are favourable but rather start breeding earlier and produce more young. Taxonomic composition and nutritional content of prey were not correlated with any of the reproductive parameters, indicating that profitability rather than quality of food affects reproductive success. Received: 31 May 1996/Accepted: 12 August 1996     

气象因子对落叶松毛虫种群数量的影响

为探究落叶松毛虫种群数量与气象因子的关系,利用大兴安岭地区落叶松毛虫发生年度监测数据与气象数据构建线性回归模型,以赤池信息量准则为评价依据,通过逐步回归的方式从拟合优度和复杂度两方面权衡选取最优模型.采用相对权重法确定影响落叶松毛虫种群数量的关键气象因子,并量化了关键气象因子对松毛虫种群数量的影响.结果表明: 低龄幼虫期和繁殖期是影响落叶松毛虫种群数量的关键时期,而中龄幼虫期和高龄幼虫期的影响较弱; 低龄幼虫期日相对湿度均值、低龄幼虫越冬期日均温小于-22 ℃积温和繁殖期降水总量是影响落叶松毛虫种群数量的关键气象因子; 低龄幼虫期日相对湿度均值和繁殖期降水总量的标准差每增加1倍,落叶松毛虫种群数量的标准差分别减少62%和35%;低龄幼虫越冬期日均温小于-22 ℃积温的标准差每增加1倍,落叶松毛虫种群数量的标准差增加40%.表明全球气候变暖趋势下落叶松毛虫的爆发特点等可能呈现出新的形式,建立长期的种群数量监测体系尤为重要.     

A review of spatial computational models for multi-cellular systems, with regard to intestinal crypts and colorectal cancer development

Colon rectal cancers (CRC) are the result of sequences of mutations which lead the intestinal tissue to develop in a carcinoma following a “progression” of observable phenotypes. The actual modeling and simulation of the key biological structures involved in this process is of interest to biologists and physicians and, at the same time, it poses significant challenges from the mathematics and computer science viewpoints. In this report we give an overview of some mathematical models for cell sorting (a basic phenomenon that underlies several dynamical processes in an organism), intestinal crypt dynamics and related problems and open questions. In particular, major attention is devoted to the survey of so-called in-lattice (or grid) models and off-lattice (off-grid) models. The current work is the groundwork for future research on semi-automated hypotheses formation and testing about the behavior of the various actors taking part in the adenoma–carcinoma progression, from regulatory processes to cell–cell signaling pathways.     

FACTORS AFFECTING THE AGE OF FIRST BREEDING OF THE KITTIWAKE RISSA TRIDACTYLA

At a Kittiwake colony in Northumberland, 80% of those birds which returned to their natal colony to breed were males and these supplied 52% of all male recruits. More females breed away from their natal colony than males. There was no differences in the proportions of young fledged from sites in the centre or at the edge of the colony, or by parents of different experience, which returned to breed. Kittiwakes breed for the first time at ages from 3 to 8 years, but most at 4 or S years old. Males arrive back at the colony at an earlier age than females and breed for the first time one year earlier. Males obtaining sites at the centre of the colony first breed at an earlier age than those at the edges. Neither the age nor the area of first breeding appear to be transmitted from parent to offspring. Males breeding first aged 4 years or younger produced more young than those which first bred aged 5 years or older, despite their partners laying smaller clutches. This difference was most marked among those males recruited to sites in the centre of the colony. The advantage of this earlier breeding is counteracted by a lower survival rate among those males which start to breed at the younger ages. In all breeding Kittiwakes, annual reproductive output increases with experience while annual survival rates decrease. Once they had started to breed, many birds failed to breed in one subsequent season. Nearly 60% of these cases of intermittent breeding occurred in the year following first breeding. Intermittent breeding was most frequent among young birds and among females. It is suggested that each breeding involves a cost to the individual in terms of reduced survival, and that deferred and intermittent breeding are means of guarding survival. A model is proposed whereby the age at which a bird starts to breed, the nesting site which it obtains, and its subsequent breeding strategy result in each individual producing an optimal number of reproducing offspring in its lifetime, relative to its quality.